V \>

<P. > 1

1

Quaest

lones

entomologicae

published at the Department of Entomology, Uni-

versity of Alberta, Edmonton, Canada.

VOLUME II

NUMBER 1

JANUARY 1966

QUAESTIONES ENTOMOLOGICAE

A periodical record of entomological investigations, published at

the Department of Entomology, University of Alberta, Edmonton, Alberta.

Volume 2 Number 1 3 January 1966

CONTENTS

Book Reviews 1

Salama, Voss, Tinga - Effects of microwaves

on Periplaneta americ ana and Tribolium confusum 3

McDonald - The genitalia of North American

Pentatomoidea (Hemiptera : Heteroptera) 7

Book Reviews

DAVIS, H.S., Editor. MEILLON, BOTHA DE, HARRINGTON, J. S.

and KALK, MARGARET, Associate Editors. 1964. Ecological Studies

inSouthern Africa. Monographiae Biologicae. Vol. XIV. Dr. W. Junk.

The Hague, xxiv + 415 pp. 23 plates. 47 figs. Cloth bound. Price -

60 Guilders ($18.00 Canadian).

This volume is intended as a companion volume to Biogeography

and Ecology in Australia, vol. VIII in the same series*. It contains 28

papers on ecological studies in southern Africa of which eight deal directly

or indirectly with insects. The general intention is to make available to

workers outside southern Africa a summary of ecological work being

carried out there and in this it succeeds. However it does not coverall

aspects of the ecology of southern Africa and so is not as useful as its

companion volume on Australia.

Most of the papers are general in scope and some of them are

rather superficial. Papers of especial interest are that by Cooke on the

Pleistocene environment in southern Africa, two papers by Vesey-

FitzgCrald and A. Leu on locusts and a paper by Brynard on the effects

of veld burning on the vegetation and game in the Kruger National Park,

the first scientific study I have seen of this very controversial subject.

The paper by Alcock on the advance of the deserts is also of considerable

interest.

An introductory chapter integrates the papers and gets them into

the perspective of other research in southern Africa. Useful biblio-

graphies accompany each paper.

The book is printed on high quality paper and has an attractive

and durable dust cover. The quality of the illustrations is good and that

of the photographic plates is excellent.

This book should be referred to by all per sons interested in African

ecology but cannot be considered worth its extremely high cost.

SMXT, BERNARD. 1964. Insects in Southern Africa : How to control

them. Oxford University Press . Cape Town. South Africa, xiv + 399pp.

Price - Rand $4. 95 ($8. 75 Canadian).

This is a small volume written for "Students, Health Officers,

Gardeners and Farmers" rather than Entomologists, by one of South

Africa's most respected entomologists. The work integrates and sum-

marizes all that is known about insect pests and their control in South

Africa in a manner that can be readily under stood by laymen. It is written

in a friendly, breezy style laced with anecdotes from the experiences of

the author and others. Technical terms are kept toaminimum and those

that are nesses sary are explained in the text obviating the need for a

glossary.

Although the book is mainly concerned with pests, beneficial

insects, especially those used in biological control, receive a fair share

of the space and conspicuous insects of little economic importance such

as mayflies and dragonflies are not excluded.

The sections on control are refreshing at the present time, when

most books on this topic tend to stress chemical controls to the exclusion

of all others. One frequently finds advice against spraying when natural

enemies are present and considerable weight is given to cultural controls

such as proper rotational grazing and the proper timing of agricultural

practices.

The older Pre World War II insecticides are also recommended

when the author believes they are still adequate or superior to the newer

ones.

Persons with no background in entomology may find the taxonomic

arrangement of the book confusing and it may prove difficult for an un-

trained person to run down the source of his trouble from it. This is to

some extent offset by a visual index in which figures of all major groups of

insects and terrestrial arthropods are indexed to the appropriate text,

but remains a major fault.

In many ways this work is a model of extension writing and although

it refers exclusively to southern Africa should be read by all persons

connected with entomological extension work.

Peter Graham

3

EFFECTS OF MICROWAVES ON PER1PLANETA AMERICANA AND

TRIBOL1UM CONFUSUM

H.S. SALAMA, W.A.G. VOSS, andW. TINGA

Department of Entomology Quaestiones entomologicae

University of Alberta 2:3 — 5. 1966

The cockroach, Periplaneta americana L. and the confused flour beetle, Tribolium

confusum Duv. can be killed by low-level microwave power radiation at a frequency of 2,450

megacycles (mcs) with electric field strengths less than 65 volts per cm (V /cm) in a plane wave

transmission system, as well as in multimode resonant cavities (electronic ovens) supplied with

1 or 2 kilowatts (kw) of power. The lethal effect of radiation at this frequency is mainly due to

heat. In P. americana, localised internal molecular heating occurs, due to the size of the insect.

With the smaller T. confusum, direct heating of the insect does not appear to be a predominant

factor but it is shown that these insects can be killed in large volumes of flour, with microwave

power used to raise the temperature of the material. However, the exposure time required de-

pends on the insect’s size in relation to the wavelength. Effects on the activity of both species

were observed and adverse effects on reproduction are established with T. confusum adults.

INTRODUCTION

Frings (1952), and Whitney et al. (1961) have discussed the effec-

tiveness of radio frequency electromagnetic waves in killing certain insect

species. Their results show that the mortality is predominantly a thermal

effect, energy being absorbed by the insects in prefer ence to surrounding

media such as flour and grain. Reproduction was not affected in those

specimens that survived the treatment (Whitney et al. , 1961).

This investigation was made to determine the effectiveness of far

higher frequencies, in the radar range (2,450 mcs), at comparable in-

cident power levels (0. 5 to 2. 0 kw) on the cockroach Periplaneta americana L.

and the confused flour beetle, Tribolium confusum Duv.

The generation and transmission of microwave power has been

described by Okress et al. (1964). The energy transfer systems and the

thermal effects in relation to the properties of heating dielectrics have

been discussed by Cop son (1962) and Voss (1965). For the work described

here, microwave power was generated at 2,450 mcs by a 2 kw magnetron

used to power a plane wave transmission system 72 cm wide, the

metal plates of which were separated by 5 cm in the direction of the

electric field. In the latter case, the power density was variable be-

tween 1.25 and 5 w/ cm^, measured by a calibrated power meter on the

coupling system to the magnetron. A power density of 5 w/ cm^ corres-

ponds to an average transverse field strength of 65 V/cm in the trans-

mission system.

Insect and environment temperatures were measured with a

needle-point ir on-constantan thermocouple referenced against melting

ice. Standard rearing techniques were used for both insect species.

4

Effects of Microwaves

RESULTS

P. americana L.

Female roaches were placed individually in small enclosures

drilled in nonpower absorbent plastic foam blocks in petri dishes, and

treated in the plane wave transmission system. Various exposure times

and power densities were used. After exposure, the internal body tem-

perature was recorded in three places. Insects that survived the exposure

were kept in glass observation jars and supplied with food.

It was found that an incident power density of 1. 25 w/ cm^ kills

the insects in 90 seconds after which the average body temperature was

60 C; a power level of 5 w/cm^ kills the insects in 5 seconds with a body

temperature of 72°C. Halving the above exposure times caused the insects

to be knocked downand they died within three to five days. Theminimum

body temperature recorded for a specimen which died at once was 37°C ,

after treatment with 2.5 w/cm^ for fifteen seconds. About 60 roaches

were used in these tests. R oaches heated in a conventional oven (at 60° C

for seven minutes) survived and behaved normally (average body temper-

ature 38° C). After twenty minutes in the same oven death resulted,

with a body temperature of 41° C.

Although the above results are not completely determinant, it is

postulated that either localised heating or other physiological changes

occur within the insect at low energy densities. The problem is par-

ticularly complex as an unknown proportion of the power density incident

on the insect is transmitted through it. The known behaviour of dielectrics

containing water suggests that the transmitted power would increase with

dielectric (insect) temperature. Further, it was found that legless speci-

mens heat more slowly than normal ones, whereas the relative orien-

tation of the body to the electric field had no observable effect.

T. confusum Duv.

Different developmental stages of the confused flour beetle were

treated in a carefully designed 48. 5 x 40 x 38. 5 cm multimode cavity,

excited by a magnetron at 2, 450 mcs. In one test, 1. 2 kw of power was

supplied to ten pounds of flour in the cavity, in which different stages of

the insect were confined to specimen tubes distributed uniformly through-

out the volume. After 2. 5 minutes a fairly uniform temperature of the

flour was recorded at 42°C (ambient 25°C). Mortality rates were zero.

The same procedure was followed using 25 pounds of flour. The tem-

peratures were recorded as 32°C (uniform) after 2 minutes, 45-75° C

and 55-90°C after 4. 5 and 7 minutes respectively, the variation of tem-

perature being due to non-uniform heating in still flour, caused by vari-

ations in the electric field strength and moisture content (average value

12%). Specimens were removed from various areas at the times indi-

cated. After the 4.5 minute interval, some insects survived in the lower

temperature zones, all others were killed. Further tests indicated that

the reproduction of T. confusum was affected by this treatment. In one test

designed specifically to investigate this, a large number of insects,

lightly covered with flour were exposed to 1.2 kw in the cavity system

for4minutes. The initial mortality was zero but 90% failed to reproduce.

Salama, Voss, & Tinga

5

DISCUSSION

In view of the technical feasibility of generating high levels of

microwave power at low cost, the method may prove to be economically

feasible for sterilizing flour infested with T. confusum and may have other

applications. Of more fundamental interest is the possibility of isolating

the thermal and physiological effects induced by high frequency electric

fields using the techniques described in this paper.

ACKNOWLEDGEMENTS

The writers are indebted to Professor B. Hocking for suggesting

the project; and to the Departments of Entomology and Electrical

Engineering of the University of Alberta in Edmonton for providing the

facilities.

REFERENCES

Copson, D. 3 1962. Microwave Heating, Avi Publ. Co. Westport, Conn.

Frings, H., 1952. Factors determining the effects of radio frequency

electromagnetic fields on insects and materials they infest. J.

econ. Ent. , 45 : 396-408.

Okress, E. C. et al. , 1964. Microwave Power Engineering, Spectrum 1,

No. 10, 76-100.

Voss, W.A. G. , 1965. Factors affecting the operation of high power

microwave heating systems for lumber processing. Institution

of Electrical and Electronic Engineers, 7th Biennial Electric

Heating Conference, Cleveland, Ohio.

Whitney, W. K. , Nelson, S. O. and Walkden, H.H., 1961. Effects of

high frequency electric fields on certain species of stored grain

insects, U.S.D.A. Marketing R es . Rep. 455. pp. 1-52.

(

THE GENITALIA OF NORTH AMERICAN PENTATOMOIDEA

(HEMIPTERA : HETEROPTERA)

7

F.J.D. McDonald .

Department of Entomology Quaestiones Entomologicae

University of Alberta 2:7—150. 1966

The male genitalia of 85 and the female genitalia of 80 species of North American

pentatomoid bugs are described. The male genitalia were found to vary very widely in the tribes

Pachycorini and Odontoscelini of the Scutellerinae. The female genitalia were less variable.

Species in the tribe Scutellerini are very easily defined on the basis of the male genitalia. The

Pentatominae, Asopinae, and Podopinae are very uniform in the structure of the genitalia and are

clos ely related to one another. The spermatheca of all species examined in the above sub-

families except Trichopepla semivittata (Say) (Pentatominae), has an elongate membraneous

dilation with a central sclerotized rod. Median penal lobes occur only in the Pentatominae,

Asopinae and Podopinae with the exception of one scutellerine, Symphylus carribeanus (Kirhaldy).

The Cydnidae exhibit great diversity of form both in the male and female genitalia. The status

of this family will remain obscure until further species have been examined. The Acantho-

somidae posses pentatomine type genitalia. The genitalia of Piezosternum subulatum (Thunberg)

do not resemble those of other species of the Tessaratomidae so far described. On the basis of

this work it is suggested that the Scutellerinae be accorded family status; the Asopinae and

Podopinae should be reduced to tribes within the Pentatominae; the Acanthosomidae reduced to

subfamily status within the P entatomidae and Piezosternum should be raised to subfamily status

within the Tessaratomidae. Phylogenetic ally the Pentatomoidea show some relationship to the

lygaeoid group, but this relationship is not close. The two groups are probably derived from a

common ancestor. The Tessaratomidae are an early offshoot of the hypotheti cal pentatomoid

ancestor. The main stock then developed into the Scutelleridae and the P entatomidae with the

A canthosominae a very early offshoot of the latter group.

INTRODUCTION

This study was undertaken with the hope of showing more clearly

the interrelationships of the North American genera of the Pentatomoidea.

Though many more problems have been raised than solved, I hope that

some basis is provided for a thorough taxonomic revision of this group

in the future. Both the male and female genitalia provide good taxonomic

characters and they will no doubt be used with increasing frequency,

especially where large number s of character s are required for analytical

purposes as in the rapidly developing field of numerical taxonomy.

Fairly detailed descriptions have beenmade of the male genitalia

while the female genitalia have been treated rather more generally. A

discussion of the results in each section of the work has been given with

a final overall synthesis of all points raised. The classification I have

proposed cannot be regarded as final any more than any other classifi-

cation, but, in general, it supports accepted classifications. A satis-

factory classification will depend on a thorough analysis of this super-

family on a world- wide basis.

MATERIALS AND METHODS

The specimens usedinthis workwere selectedfrom driedmuseum

material; males of 85 species and females of 80 species of pentatomoid

bugs were studied and a total of 256 specimens were examined. R epresen-

tatives of the type species of each genus were chosen wherever possible.

Material dissected out of each specimen has been placed in a microvial

and attached under the specimen. All material has been returned to the

United States National Museum, Washington.

8

North American Pentatomoidea

The genitalia were studied after treatment with 10% KOH in the

usual way. The terminalia were cleared in polyvinyl lactophenol, methyl

salicylate, or glycerine. Wherever necessary, chlorazol black was used

as a stain for membraneous structures. In some cases the internal

structure of the vesica could only be studied after thorough bleaching in

chlorine (McDonald 1961). A technique described by Kumar (1964) was

used to check the connections of the internal ducts of the vesica. This

method was not found to be entirely satisfactory.

Transverse sections of the vesica of Lampromicra senator (Fabricius)

and Cantao par en turn (White) weremade. The vesica was embedded inpara-

plast, sectioned at 6|i, stained in Mallory's triple stain and mounted in

Canada balsam.

Observations were made with a Wild and a Leitz stereoscopic

microscope with magnifications of up to 50X and 150X respectively.

Diagrams were drawn to scale using a squared ocular grid and squared

paper. Stippling where used indicates sclerotization. The conjunctival

appendages have been numbered in sequence from the dorsal to ventral

surface; the third are thus always ventral in position. The diagrams of

the vesica have nearly all been orientated so that the seminal duct is

ventral.

The classification of the Pentatomoidea throughout the descrip-

tions and discussion sections follows that of Leston (1953c). The keys

are arranged according to the proposed classification I have set out on

page 68. The generic and specific names followed are those of VanDuzee

(1917), with Kirkaldy (1909) as a second source of reference.

TERMINOLOGY

Male genitalia

The basic nomenclature used is that of Pruthi (1925) with slight

modifications. I have retained the term median penal lobe used by Baker

(1931) for the inner sclerotized lobes surrounding the vesica in Penta-

tominae, Asopinae and Podopinae. The homology of these lobes is un-

certain. The term endophallic duct is used for the duct between the

ejaculatory line which I used wrongly in previous papers (McDonald 1961,

1963). I feel this is a more basic term than conducting canal 2 of Kumar

(1964). I also cannot accept the latter author ‘s term conducting chamber

in place of ejaculatory reservoir, because outside the highly specialized

Scutellerini this chamber is usually well developed and probably does

act as a reservoir for sperm. Figure 1 is a general diagram showing

the terminology used in this paper.

Female genitalia

The nomenclature used in this section is that of Scudder (1959)

and Pendergrast (1957) for the spermatheca. For the purposes of this

study the spermatheca has been considered as part of the female genitalia.

McDonald

9

MORPHOLOGY OF THE MALE GENITALIA

The external features of the male genitalia are easily observed.

The appendages of the aedoeagus are sometimes difficult to expand and

wherever these have not been fully expanded this is stated in the text.

The male genitalia are situated in the ninth segment which is

modified into a cup-like structure, the pygophore (fig. 4). Within the

pygophore is a pair of hook-like structures, the claspers (fig. 5) and a

tube-like structure, the proctiger bearing distally the anus. Lying be-

neath the proctiger is the aedoeagus (fig. 1) attached by means of the

basal plate to the ventral surface of the pygophore.

The internal structure of the vesica has been worked out and

drawn as accurately as possible. However, without actually sectioning

the specimens interpretation of the internal ducts is subject to error and

cannot be regarded as final until sections of all species have beenmade.

Even so, the gross morphological details are quite readily observable

and these serve adequately for studies in homologies between genera at a

tribal level and above.

The tribe Scutellerini, represented in North America by the genus

Augocoris has a peculiarly developed convoluted duct passing back from

the entrance of the seminal duct into the ejaculatory reservoir (fig. 83).

Kumar (1964) states that this duct (conducting canal 1) is composed of

two sets of canals and that the seminal duct passes directly into this duct.

Cross sections of Lampromicra senator and Cantao parentum (Australian scutel-

lerines) show quite clearly that this duct is in fact single but of a highly

convoluted nature (fig. 3). Sections also showthat the seminal duct opens

into the base of the endophallic duct as does the convoluted duct. Un-

fortunately, sections could not be made of the genitalia of Augocoris gomesii

as very few specimens were available for study.

Pentatomidae -Scutellerinae

Odontoscelini

Fokkeria producta (Van Duzee), 1904

Pygophore (fig. 4) with dorsal border deeply and evenly arched;

ventral border U - shaped. Pygophoral opening with wide dorsal and lateral

flanges. A number of small setae on dorsal and lateral borders.

Claspers (fig. 5) small stem basally wide; apically narrowing

into a shallow hook; inner margin finely serrate. A number of setae on

mid region of stem.

Theca (fig. 6) conical, not heavily sclerotized. Three pairs of

conjunctival appendages borne on a common membraneous base: first,

conical, membraneous with a small sclerotized apex; second bifid,

consisting of two heavily sclerotized horns borne on a short membraneous

base; third large oblong structures bluntly rounded apically, heavily

sclerotized throughout and covered with numerous flat, blunt teeth.

Vesica strap-like, dor soventr ally compressed, base wide, sclero-

tized, fused to ventral margin of theca, tapering distally, apex mem-

braneous and bent through 90°. Seminal duct (fig. 7) leading into a small

10

North American Pentatomoidea

globose ejaculatory reservoir, walls of latter thickened; endophallic

duct connected to anterior end of reservoir, apically terminating within

membraneous apex of vesica.

Euptychodera corrugata (Van Duzee), 1904

Pygophore small (fig. 8), opening with dorsal and lateral flanges,

ventral border emarginate, a number of small setae scattered along the

lateral margins.

Claspers (fig. 9) shallow hooks, no differ entiation between apical

hook and stem. A number of stout setae along stem, inner basal margin

bearing many minute spines. Dorsal surface of hook scalloped.

Theca (fig. 10) small, conical. Three pairs of conjunctival ap-

pendages: first basally wide, membraneous, apically produced into

heavily sclerotized points; second consisting of heavily sclerotized

curved horns attached to a short membraneous base fused to common

base; third, (figs. 10, 11) large, sclerotized plate-like structures, outer

surface covered with numerous short stout spines; appendages normally

folded within common base beneath second conjunctival appendages.

Vesica (fig. 10) narrow and flattened dor so-ventrally, basally

attached to theca; sclerotized except for apical third. Seminal duct

(fig. 12) leading ventrally into anterior portion of bilobed ejaculatory

reservoir. Posterior lobe forming a small chamber lying somewhat on

top of larger anterior chamber . A wide endophallic duct connecting with

ejaculatory reservoir, apical opening membraneous.

These genitalia resemble very closely those of Fokkeria producta

especially in the form of the conjunctival appendages. I think that

Euptychodera is probably congeneric with Fokkeria .

V anduzeeina halli (Van Duzee), 1905

Pygophoral opening with a large flattened flange (fig. 13) laterally

on each side; dorsal margin wide; ventral margin narrow, flattened,

bearing a number of small fine setae. Proctiger with numerous long

fine setae on apex.

Claspers, scythe-like (fig. 14); stem continuous with apical hook

and covered with stout setae along outer margin; a number of longer

setae found at base of hook.

Theca (fig. 15) small, conical, not heavily sclerotized. Three

pairs of conjunctival appendages (fig. 16) present, fused onto wide mem-

braneous conjunctiva: first, basallv membraneous and wide; apically

produced into a small curved heavily sclerotized point, second completely

enclosed by their membraneous common base within theca when not ex-

panded; apex of each appendage terminating in a very large flattened,

heavily sclerotized horn; third, minute structures situated at bases of

second conjunctival appendages; apically sclerotized and pointed.

Vesica, small narrow and flattened, basally sclerotized andfused

to conjunctiva; medianly divided into two rounded sclerotized projections

lying one on each side (fig. 17) of a membraneous tube, within which is

the endophallic duct, projections bearing a number of teeth on their

apices. Apex of vesica with a wide flange. Internally, seminal duct

McDonald

11

passing ventrally up base of vesica and into small trilobed ejaculatory

reservoir; endophallic duct straight, basally merging into apex of reser-

voir.

Phimodera binotata ( P. torpida ) (Say), 1824

Pygophore with dorsal margin (fig. 18) very broad, covered

with fine setae; two small spine-like projections found laterally one on

either side on ventral border above bases of claspers. Ventral border

bearing numerous short stout setae.

Claspers (fig. 19) small, stem drawn out into a blunt apex; a

square projection lying below apex forming a shallow hook. Twelve to

seventeen setae found along apical half of stem; a number of very minute

setae found on under surface of apex.

Theca (fig. 20) cylindrical, apical mar gin merging into conjunctiva

when latter fully expanded. One pair of membraneous cylindrical con-

junctival appendages (fig. 22) attached ventrally into base of endophallic

duct; a ventral canal leads back into a large ejaculatory reservoir from

which a second dorsal canal opens into base of endophallic duct; latter

basally thickened and thrown into a number of loops, finally widening and

opening at secondary gonopore.

Eurygas trini

Eurygaster alternata (Say), 1828

Pygophore with dor so-lateral border (fig. 23) rounded, extending

down on each side to fuse with flattened and plate-like ventral margin.

Fine setae along margins of dor so-lateral border.

Claspers T-shaped (fig. 24) with a thick stem tapering basad.

A number of fine scallopings found on inner surface of each arm of cross-

piece and several small setae on each side of stem at its junction with

cross arm.

Theca conical, very slightly sclerotized, bearing on the ventral

margin centrally a long cylindrical membraneous process (fig. 25), apex

slightly sclerotized, pointed. Three pairs of conjunctival appendages

present; first membraneous basally, apically sclerotized forming a stout

curved horn; second heavily sclerotized, horn-like; third, very small

sclerotized horns.

Vesica, consisting of a long cylindrical tube, apically mem-

braneous, hook-shaped, upper margin of hook bearing a fringe of hairs;

basally sclerotized and bearing on dorsal surface a pair of leaf-like

vesical processes (fig. 26). Seminal duct opening ventrally into an an-

terior sinus to which posteriorly is attached a small elongate and heavily

sclerotized reservoir. Endophallic duct originating from anterior sinus

and terminating in a wide membraneous secondary gonopore lying within

invaginated apex of vesica.

12

North American Pentatomoidea

Pachycorini

Camirus moestus (Stal), 1862

Dorsal border of pygophore evenly arched (fig. 27), laterally with

two rounded prominences, ventral border almost straight.

Claspers (fig. 28) simple hook- like, shallow, stem fairly long.

Theca squat, cylindrical, not heavily sclerotized. Two pairs of

conjunctival appendages: first (fig. 29) entirely membraneous bag-like,

apically rounded; second bifid, ventral arm short, flat, sclerotized and

disc-like; dorsal arm membraneous, cylindrical, tapering apically to a

blunt point; both arms borne on a common partially sclerotized stem.

Vesica (fig. 30), complex; endophallic duct apically surrounded

by a large oblong membraneous sheath covered with very fine spines.

Seminal duct very fine, passing ventrally into base of ejaculatory duct.

A wide thickened convoluted duct extending back from entrance of seminal

duct, widening posteriorly into a large sinus; latter communicating by

means of a valve-like arrangement with a large dor sal ejaculatory reser-

voir. A long funnel-like duct connecting ejaculatory reservoir with a

narrow ventral chamber latter leading anteriorly into a short endophallic

duct. The vesica resembles that found in the Scutellerini (McDonald,

(1961) particularly in possessing a long convoluted duct.

Pachychoris torridus (Scopoli), 1772

Previously described by Kumar (1965). Pygophore with dorsal

margin membraneous, bearing medianly a narrow, heavily sclerotized

band produced into a broad median process (fig. 32), apically acute. A

pair of cylindrical pygophoral appendages lying one on either side of

median process; each appendage apically with two spines (fig. 33) outer-

most spine being single, innermost bifid. Lateral margins of pygophore

somewhat flattened; ventral border flattened and shelf-like.

Claspers (fig. 34) small; stem short, stout, merging into a broad

flattened hook; a number of fine setae at base of hook.

Theca (fig. 35) cone-shaped. Two pairs of conjunctival appen-

dages: first, large completely sclerotized, horn-like structures fused

to margin of theca; second cylindrical, rodlike, apically smoothly

rounded bearing half way along ventral mar gin a stout spine, below which

is a deep notch.

Vesica (fig. 36) extremely small and simple in construction;

lying between bases of second conjunctival appendages; apically opening

into a longitudinal groove. Seminal duct opening into a small tube expanded

medianly into an anterior sinus, narrowing distally into a very short

endophallic duct. The vesica is unusual in not possessing a free apical

portion as in other Scutellerinae.

Chelysomide a guttata (Her rich-Schaeffer )? 1839

Dor sal border of pygophore (fig. 37) arcuate; practically obsolete

medianly, laterally produced on each side into a stout sclerotized point.

Ventral margin flattened, border almost straight. Proctiger very dis-

tinct, membraneous except for a narrow dorsal median sclerite apically

McDonald

13

produced into a short curved median spine, lateral margins each produced

into a long spine; anal opening lying between this triad of spines. A

narrow band of very fine spicules on lateral margins on each side from

base of lateral spine.

Claspers scythe-shaped (fig. 38), stem broad, a number of very

small teeth found along inner margin of hook.

Theca (fig. 39) membraneous, hardly differentiated from con-

junctiva. Two pairs of conjunctival appendages: first stout, L- shaped

horns, heavily sclerotized; second long, heavily sclerotized, apically

flattened, and blade-like.

Vesica simply constructed, seminal duct (fig. 40) opening ven-

trally into a wide sclerotized tube, posteriorly dilated forming ejaculatory

reservoir; anteriorly extended as a wide endophallic duct, latter apically

membraneous .

Homaemus aeneifrons (Say), 1824

Pygophoral opening roughly hexagonal (fig. 41), dorsal margin

broad bearing a number of fine setae, laterally dorsal margin bears a

small pointed projection on each side. Ventral margin flattened some-

what medianly.

Claspers (fig. 42) hook-shaped, with a stout stem, a number of

long setae at base of hook.

Theca (fig. 43) small, conical. Three pairs of conjunctival ap-

pendages: first (fig. 44) basally voluminous membraneous structures,

apically bearing a heavily sclerotized horn; second, small, elongate,

membraneous structures; third broad and flattened, moderately sclero-

tized and with numerous small teeth scattered over outer surface (fig.

45).

Base of vesica resembling a nautilus shell (fig. 46), coiled and

with a number of pseudopartitions; central portion of coil attached on

either side to common base of conjunctival appendages. Apex of vesica

broadly rounded, armed with a large number of small spines. Seminal

duct extending ventrally into apical half of vesica and into a wide endo-

phallic duct, latter bent through 90°, apically membraneous and opening

into a membraneous pouch on mid dorsal surface of vesica. Ejaculatory

reservoir small and continuous with endophallic duct. The coiled struc-

ture at the base of the vesica may enable fluids to be pumped into the

appendages, thereby expanding them. The vesica is very similar to that

of H. aeneifrons consors examined by Kumar (1965).

Tetyra antillarum (Kirkaldy), 1909

Pygophore with dorsal border acutely arched (fig. 47), laterally

bearing two smooth sausage- shaped calluses one on each side lying just

above apex of claspers. Ventral border sinuous bearing a rounded ridge

on ventral surface. Lateral margins with numerous short setae; ventral

border with long fine setae.

Claspers (fig. 48) hook- shaped, bifid at apex, outer tooth acute,

inner tooth blunt, both heavily sclerotized. Stem short, squat, bearing

a number of long setae on a slight promontory at junction with hook.

14

North American Pentatomoidea

Theca (fig. 49) small* cone-shaped. Two pairs of fairly large

conjunctival appendages first (fig. 50) fused basally for about half their

length, apically bearing a small heavily sclerotized point; second also

basally fused, apical two-thirds free and capped with a long heavily

sclerotized horn.

Vesica, long and narrow, heavily sclerotized; dorsal margin

bearing a triangular sclerotized supra vesical process (fig. 51). Seminal

duct merging ventr ally into a wide S- shaped endophallic duct; ejaculatory

reservoir, small crescent-shaped, opening into base of endophallic duct.

Sphyrocoris obliquus (Germar), 1839

Pygophoral opening surrounded by a wide flange dor sally and

laterally (fig. 52): ventral margin flattened forming a lip with a slight

median indentation. Fine setae found on lateral mar gins and along ventral

border .

Claspers (fig. 53) scythe- shaped, stem short and stout; base of

hook bearing a number of large stout setae. Inner surface of hook scal-

loped.

Theca (fig. 54) small, cup- shaped. Two pairs of conjunctival

appendages: first membraneous broadly rounded at apex and fused to

mar gin of theca; second basally membraneous, apically bearing a small

heavily sclerotized horn.

Vesica (fig. 55) divided into two parts, ventral portion widely

V-shaped, apex blunt bearing a large number of barbs. Dor sally and in

apposition is a wide supra-vesical process marked with a number of

striae, upper margin with a groove and bearing a number of small sharp

teeth; this process fused to base of vesica.

Endophallic duct (fig. 56) opening into a small anterior sinus at

the base of the ejaculatory duct, latter fairly straight, situated along

ventral arm of vesica opening at its apex. Ejaculatory reservoir pear-

shaped, lying above and directly connected to anterior sinus.

Stethaulax marmoratus (Say), 1831

Dorsal border of pygophore (fig. 57) diffuse with a small notch

just above base of clasper on each side, ventral margin flattened.

Claspers (fig. 58) small club-like structures, apically produced

into a small beak-like point, a number of long setae laterally and beneath

the apex on the stem, apex minutely scalloped on both sides.

Theca (fig. 59) small, cup-shaped with a shallow median groove

on the ventral margin, lateral surfaces with a number of very minute

spines scattered in a band just below the margin. Two pairs of conjunc-

tival appendages: first long, cylindrical, basally membraneous, apically

with a heavily sclerotized bluntly rounded tip; second basally mem-

braneous, broad, apically produced into a long heavily sclerotized horn.

Vesica (fig. 60), apex broad, flattened in a dor so-ventral plane.

Seminal duct (fig. 61) opening ventrally into a small sinus, lying above

and connected to latter is a small elongate dorsal reservoir. Extending

back from sinus is a wide duct (figs. 61, 62) becoming convoluted and

thickened, posteriorly widening into dorsal chamber of ejaculatory

McDonald

15

reservoir, lying beneath is a ventral chamber connected to upper chamber

by a narrow passage. Endophallic duct L- shaped originating from en-

trance of seminal duct.

Symphylus carribeanus (Kirdaldy), 1909

Dorsal margin of pygophore broad (fig. 63), extending laterally

and merging into flattened ventral margin. Ventral and lateral margins

with numerous long fine setae, proctiger distally also covered with fine

setae.

Claspers (fig. 64) heavily sclerotized hammer- shaped, stem

centrally swollen, fused at right angles to cross arm one side of which is

longer than the other; longer arm bearing two small teeth apically, one

on upper mar gin and one on lower; short arm bearing one tooth on lower

margin. Lateral surfaces of head finely scalloped, a number of stout

setae on stem below junction with head.

Theca (fig. 65) small, globose. Two pairs of conjunctival appen-

dages: first blade-like moderately sclerotized apically acute; second

very long cylindrical membraneous, apically bearing a heavily sclerotized

horn. Median penal lobes present, dor sally fused together around base

of vesica, apically free forming two broad flat plates (fig. 66) on either

side of apex of ejaculatory duct; ventral margin with a number of peg-

like teeth (fig. 65).

Seminal duct (fig. 67) opening ventrally into a small bilobed an-

terior sinus; from latter a wide duct opening into dorsal chamber of

ejaculatory reservoir. Dorsal chamber connected to a large ventral

chamber by means of a narrow passage. Endophallic duct extending

from anterior sinus, short and straight, apically terminating a short

distance beyond the margins of the median penal lobes.

Diolcus irroratus (Fabricius), 1775

Dorsal border of pygophore U-shaped (fig. 68), laterally with

two C-shaped indentations lying adjacent to apices of claspers on each

side. Ventral border narrow, flattened, slightly sinuate. Fine setae

on lateral and ventral margins.

Claspers (fig. 69) with stout stem, apically produced into a short

blunt hook. Inner margin of hook scalloped. A few stout setae at base

of hook and along outer lateral margin of stem: numerous very small

fine spines along the inner lateral margin.

Theca (fig. 70) small, conical, dorsal surface greatly enlarged

and produced into two large flat horns one on each side with a wide U-

shaped emargination between them. One pair of cylindrical conjunctival

appendages, membraneous, apically bearing a heavily sclerotized gently

curved horn.

Vesica situated in a large oblong membraneous conjunctiva, (fig.

71) long narrow and sclerotized; seminal duct (fig. 72) passing straight

into ejaculatory duct; no ejaculatory reservoir . A small ventrally pro-

jecting apodeme attached at junction of endophallic duct and seminal duct.

A deep sclerotized pit borne dor sally within conjunctiva; beneath this

pit and immediately above basal half of vesica is a band of muscle fibres.

This whole structure may be some type of pumping device.

16

North American Pentatomoidea

Acantholomidea porosa (Germar), 1839

Pygophore (fig. 73) somewhat oblong in outline, opening surroun-

ded on dor sal and lateral margins by a fairly wide flange, ventral mar gin

narrow centrally. Proctiger heavily sclerotized, antero- dor sally exten-

ded into a V-shaped process lying on top of ventral margin of pygophore.

A number of minute setae on ventral margin and a number of stout setae

on posterior margin of proctiger.

Claspers (fig. 74) with long stem, apically with a shallow hook,

a few stout setae situated at base of hook.

Theca (fig. 7 5) small squat broader than long. Three pairs of

conjunctival appendages (fig. 76): first large horn-like structures,

sclerotized almost to base which is membraneous; second smaller,

membraneous squat structures, bearing apically a pair of stout heavily

sclerotized curved spines; third narrow cylindrical, sclerotized struc-

tures, apically acute.

Seminal duct (fig. 78) entering into a wide endophallic duct latter

extremely short, not extending past margin of ejaculatory reservoir.

From endophallic duct posteriorly is a wide duct opening into an S- shaped

ejaculatory reservoir; latter divided by a septum into a large dorsal

chamber and a smaller ventral chamber.

Scutellerini

Augocoris gomesii (Burmeister ), 1835

Pygophore with dorsal border (fig. 79) smoothly rounded; ventral

border with two deep V-shaped emar ginations on either side of a stout

blunt median process, lateral mar gins produced into blunt points . Dorsal

and ventral margins covered with long fine setae.

Theca heavily sclerotized, cylindrical with two small protuber-

ances on anter o-dor sal margin (fig. 81) one on each side of mid-line.

Two pairs of conjunctival appendages: first (fig. 82) bifid, one branch

completely membraneous, cylindrical, blunt at apex, the other sclero-

tized, broadly rounded at apex, common base membraneous; a sclero-

tized band round base of first conjunctival appendages, probably rep-

resents remains of second (Leston 1952); third typically scutellerine,

heavily sclerotized, cylindrical, and apically acute.

Seminal duct (fig. 83) connected directly into base of endophallic

duct; a long convoluted duct leading back from entrance of seminal duct,

expanding dor sally into an elongate ejaculatory reservoir, latter con-

nected by a canal to a large dorsal sinus; a short stout endophallic duct

attached apically to sinus.

Pentatomidae - Pentatominae

P entatomini

Pentatoma rufipes (Linnaeus), 1758

Described and figured by Piotrowski (1950). However, because

his description is in Polish a second description in English is not out of

McDonald

17

place.

Dorsal border of pygophore (fig. 84) broadly arched medianly

with a small bilobed superior ridge. Ventral border deeply concave

(fig. 85) centrally with a narrow U-shaped inferior ridge, internally

forming two ridges (superior rests of Leston 1954).

Claspers (fig. 86) C-shaped and strap-like; divided into two

arms, upper arm apically divided into a proximal broadly rounded lobe

and a distal elongate process, lower arm fused to margin of pygophore,

apically heavily sclerotized and produced into a broad flattened flange.

Theca (fig. 87) long, cylindrical. One pair of conjunctival appen-

dages; cylindrical very lightly sclerotized; apically produced into two

blunt lobes. Median penal lobes (fig. 88) fused to form a cone around

apex of vesica with lateral margins somewhat thickened and produced

into blunt points dor sally.

Seminal duct (fig. 89) merging apically into a canal found along

ventral and dorsal margins of box-like ejaculatory reservoir ; reservoir

with an oval anterior chamber (fig. 89) separated by an incomplete septum;

canal opening into this chamber dor sally. Base of endophallic duct in-

serted into reservoir ventrally; duct short, slightly kinked, apically

terminating between median penal lobes.

Dendrocoris humeralis (Uhler), 1877

Dorsal border of pygophore (fig. 90) with a narrow superior ridge;

lying one on either side is a pair of oblong genital plates (fig. 91) upper

surfaces finely scalloped. Ventral border produced into two large trian-

gular platforms one on each side with a deep median trough between them,

outer m sir gin of trough with a deep median U-shaped emargination (fig.

92).

Claspers (fig. 93) flattened, broad, apex emarginate, heavily

sclerotized, dorsal margin apically terminating in a point, ventr o-apical

margin broadly rounded. Clasper somewhat C-shaped in outline inner

apical margin finely scalloped.

Theca elongate, cylindrical. One pair of conjunctival appendages

(fig. 94), membraneous baggy structures, apically broadly rounded (not

fully inflated in fig. ), an elongate cylindrical dorsal conjunctival lobe

present and a second broadly rounded balloon-like lobe found ventrally

(fig. 95), enclosed by conjunctival appendages, probably representing

greatly modified median penal lobes.

Ejaculatory reservoir large, divided by means of septae into a

series of ducts. Seminal duct (fig. 95) merging ventrally into a long

canal leading round base of reservoir to apico-dorsal region and into a

duct which in turn opens info a central sinus. Base of endophallic duct

continuous with central sinus, duct wide, U-shaped, bearing a small

flange ventro-basally.

Piezodorus lituratus (Fabricius), 1794

Claspers, aedoeagus and vesica figured by Pruthi (1925).

Pygophore with dorsal border (fig. 96) widely U-shaped, opening

of pygophore small. Ventral border with a shallow median emargination,

gently sloping dorsad on either side. Ventral surface beneath border

18

North American Pentatomoidea

almost vertical with two shallowmedian depressions. A number of stout

setae found along margins.

Claspers (fig. 97) with fairly broad stem apically bent at right

angles forming a short triangular head, outer lateral margin finely

scalloped.

Theca small, oval in outline. Two pairs of conjunctival appen-

dages (fig. 98): first membraneous, cylindrical, apically tapering

slightly to a sclerotized blunt apex, basally produced into four conjunc-

tival lobes; second basally membraneous, cylindrical, apically bearing

a heavily sclerotized horn. Median penal lobes (fig. 99) flattened and

leaf-like, fused along their dorsal margins, and to the sub-apical portion

of the vesica.

Ejaculatory reservoir (fig. 100) divided into two chambers by

means of a stout sclerotized septum. Seminal duct merging ventrally

into a narrow canal passing round posterior margin of ejaculatory reser-

voir to open into anterior chamber. Endophallic duct opening out from

posterior chamber of ejaculatory reservoir, moderately long and sinuous,

apically terminating between median penal lobes.

Solubea pugnax (Fabricius), 1775

Pygophore (fig. 101) and claspers (fig. 102) described by Sailer

(1944).

Theca (fig. 103) cylindrical with a short membraneous hinge

attaching it to basal plates. One pair of conjunctival appendages, small,

moderately sclerotized throughout, apically broadly rounded. Median

penal lobes forming a cylindrical sheath round apex of vesica (fig. 104),

apical lateral margins heavily sclerotized forming a flat plate ventrally

on each side, dorsally lobes fused together by means of a membrane.

Seminal duct (fig. 105) enclosed in a thick membraneous sheath,

opening ventrally into base of endophallic duct; latter moderately long

enclosed in a stout cylindrical sheath, apically bent through 90°, basally

widening into a small bulb-like ejaculatory reservoir.

O

Peribalus limbolarius (Stal), 1872

Pygophore (fig. 106) and claspers (fig. 107) described by Baker

(1931).

Theca (fig. 108) vasiform expanded anteriorly into a large thecal

shield; two pairs of processes on each side of theca; anterior processes

smaller, heavily sclerotized, broadly rounded apically; posterior pro-

cesses smaller, heavily sclerotized, broadly rounded. One pair of mem-

braneous conjunctival appendages divided into two broad lobes. Median

penal lobes (fig. 109) flattened laterally into two wide sclerotized plates,

ventro-apical margins finely serrate, basally united by a narrow cross-

bar one third distance from their bases, lobes not enclosing apex of

vesica.

Ejaculatory reservoir (fig. 110) somewhat oblong with a canal

round posterior surface into which seminal duct enter s ventrally. Endo-

phallic duct narrow, sinuous, basally opening into apex of reservoir.

McDonald

19

Trichopepla semivittata (Say), 1832

Pyg°ph°re with dorsal border (fig. Ill) deeply and evenly arched,

bearing a small protuberance on each side; ventral border (fig. 112)

widely U-shaped. Two flat leaf-like genital plates one on either side

lying beneath the protuberance on the dorsal border.

Claspers (fig. 113) C- shaped, bifid at apex, no differentiation

between stem and apex; five fine setae on inner margin.

Theca lightly sclerotized, cylindrical and tapering apically. One

pair of conjunctival appendages (fig. 114), membraneous except for a

line of scler otization along ventral margin, apically broadly rounded.

Median penal lobes absent.

Seminal duct ventrally opening into a simple globular ejaculatory

reservoir. Endophallic duct heavily sclerotized, attached to reservoir

dor sally, duct long, thin and sinuous, apically a fine needle-like point.

Mormidea lugens (Fabricius), 1775

Pygophoral opening small (fig. 115), surrounded by wide mar gins,

dorsal border deeply arched; ventral border medianly U-shaped with

two acute prominences one on either side of emargination.

Claspers (fig. 116) very small, stout oblong structures, apically

very broadly rounded.

Theca (fig. 117) small, squat. One pair of membraneous balloon-

like conjunctival appendages. A large sheath-like structure present,

probably fused median penal lobes; moderately sclerotized, ventrally

with a deep cleft, basally narrowed and forming a short cylindrical stem;

whole structure surrounding apex of vesica.

Endophallic duct (fig. 118) moderately long bearing two sclero-

tized flanges, medianly wide, rounded, apically tapering, basally endo-

phallic duct expanding into a small bulb-like ejaculatory reservoir.

Seminal duct connecting with base of ejaculatory duct ventrally.

Brepholoxa heidemanni (Van Duzee), 1904

Opening of pygophore dorsad (fig. 119) and somewhat triangular.

Dorsal border shallowly concave with a narrow superior ridge not clearly

differentiated from border proper. Ventral border with two elongate

calluses forming a V; a deep U-shaped notch formed between apices of

calluses. Two further notches found, one on either side between junction

of dorsal and ventral borders.

Claspers (figs. 120, 121) with short, stout stem, C-shaped, bear-

ing apically a small heavily scler otized triangular pad, finely scalloped,

a second triangular pad found below apical one also finely scalloped.

Theca small, cylindrical. One pair of conjunctival appendages

(fig. 122) divided into three broad membraneous lobes, ventral most

fused together forming a platform beneath apex of vesica; dorsal lobes

largest, somewhat leaf-like.

Ejaculatory reservoir (fig. 123) membraneous, oval, with a pair

of septa attached to dorsal surface, reaching mid way into reservoir

forming an upper chamber. A canal leads from apico-ventral surface

round posterior mar gin of reservoir to open apically into an upper cham-

ber. Seminal duct inserted directly into this canal. Endophallic duct

20

North American Pentatomoidea

originating from apex of lower chamber of reservoir, short and curved

through 90°.

Arvelius albovunctatus ^DeGeer), 1773

Dorsal border of pygophore deeply concave (fig. 124) with two

rounded projections laterally, (fig. 125) one on each side; ventral border

also deeply concave, margin flattened into a lip bearing two longitudinal

ridges.

Claspers (fig. 126) F-shaped, distal arm apically bluntly bilobed.

Theca cylindrical with two small sharply pointed processes on

dorsal margin (fig. 127). One pair of conjunctival appendages composed

of three membraneous rounded lobes fused onto a common base and a

dorsal median conjunctival lobe. Median penal lobes consisting of a flat

pair of plates apically truncate, lobes fused along their lower margins

forming a trough around apex of vesica.

Seminal duct (fig. 128) opening medianly into a wide canal round

base of ejaculatory reservoir and connected to a dorso-apical chamber;

ejaculatory reservoir oval incompletely divided by means of a stout septum

into two chambers. Endophallic duct S-shaped basally, originating from

apico-ventral half of ejaculatory reservoir.

Aelia americana (Dallas), 1851

Pygophore (fig. 129) and claspers (fig. 130) described by Baker

(1931).

Theca (fig. 131) cylindrical somewhat diamond- shaped indorso-

ventral plane due to two lateral conical projections, one on each side;

dorsal margin produced into a thecal shield. One pair of conjunctival

appendages, broad, membraneous lobes rounded apically; when fully

inflated balloon-like. A large membraneous conjunctival lobe present.

Median penal lobes (fig. 132) small, thin, heavily sclerotized, fused to

a wide common base.

Seminal duct (fig. 133) inserted directly into a long canal which

opens by means of a valve-like arrangement into dor so-apical region of

ejaculatory reservoir, latter lying centrally, apically merging into a

sinuous endophallic duct with spout-like apex.

Vulsirea violacea (Fabricius), 1803

Pygophore with dorsal margin (fig. 134) bearing two oval patches

of short heavily sclerotized setae one on either side, similar to patches

of setae foundin Scutellerinae (McDonald 1961), however in this species

setae not arranged in rows. Ventral margin gently concave bearing

laterally on each side a stout finger-like process, further behind these

is a pair of stout bifid processes one on each side; a narrow ridged floor

withadeep median V-shaped cleft running between these inner pygophoral

processes.

Claspers (fig. 135) with stem short, apically produced into three

lobes. Outer surfaces of lobes finely scalloped.

Theca small, somewhat oblong (fig. 136). Two pairs of conjunc-

tival appendages: first membraneous, basally wide, apically produced

into a narrow sclerotized point; second membraneous, cylindrical.

McDonald

21

apically blunt, basally fused to the first. Median penal lobes (fig. 137)

disc-like, medianly fused to one another and to sub-apex of vesica,

basally each lobe produced into a bifid process.

Ejaculatory reservoir (fig. 138) large with a partial septum

dividing it into two chambers. Seminal duct inserted into base of endo-

phallic duct, from this point is a long duct slightly convoluted posteriorly

opening into the apico-dorsal portion of ejaculatory reservoir. Endo-

phallic duct wide and short, fused between median penal lobes.

Acrosternum pennsylvanicum (DeGeer), 1773

Dor sal border of pygophore with a broad superior ridge (fig. 139).

Ventral border shallowly concave (fig. 140), two flat processes found on

margin one on either side, outer margin concave with a number of very

heavily sclerotized teeth.

Claspers (fig. 141) simple, spear-shaped with a very short stout

stem.

Theca small, cylindrical. One pair of small membraneous con-

junctival appendages (fig. 142) attached to a large membraneous base.

Median penal lobes tubular, sclerotized, with a small expanded head

apically, basally fused to a common stem.

Seminal duct (fig. 143) opening ventrally into a simple sac-like

and membraneous ejaculatory reservoir. Endophallic duct passing for-

ward from apex of reservoir, slightly kinked, short, apically terminating

between apices of median penal lobes.

Chlorocoris subrugosus (Stal), 1872

Pygophore with dorsal border (fig. 144) medianly evenly arched,

laterally bearing two small projections one on each side; ventral border

(fig. 145) deeply concave, sinuate, medianly with a bilobed inferior ridge

with a minute spine on either side laterally. Proctiger long, narrow,

bearing apically two flat lobes covered with a mat of fine setae.

Apex of claspers formed into a trilobed umbr ella«like structure

(fig. 146) stem short and slender, upper surface of apex covered with a

dense mat of fine setae.

Theca large, cylindrical with very large basal plates; apically

produced into a short cylindrical thecal shield (fig. 147) surrounding the

sub-apical portion of the vesica. No conjunctival appendages or median

penal lobes.

Seminal duct wide passing medianly into a canal, latter encircles

posterior end of ejaculatory reservoir and opens dorsally. Ejaculatory

reservoir simple sac-like, endophallic duct originating from posterior

end of reservoir as a wide duct, narrowing anteriorly, short, slightly

curved, apically terminating a short distance beyond thecal shield.

Carpocoris remotus (Horvath), 1907

Pygophoral opening wide, triangular (fig. 148); dorsal border

widely arched bearing two large cone-shaped lateral projections one on

either side of mid line; lying beneath these, one on each side, thin

saucer-like genital plates (fig. 149), very lightly sclerotized with a

fringed margin and numerous small spines on upper surface. Ventral

22

North American Pentatomoidea

border (fig. 150) with a small median V-shaped emargination and two

rounded prominences one on either side of median emargination.

Claspers (fig. 151) with short stout stem produced into a flattened

oblong leaf-like apex bearing a sharp tooth on lower angle, dorsal sur-

face deeply cleft.

Theca (fig. 152) small, squat with two small projections one each

side, on the apical margin in a dor so-lateral position. One pair of

membraneous conjunctival appendages (figs. 152, 153), leaf-like, with

a sharp sclerotized ridge along ventral margin.

Ej aculatory reservoir (fig. 154) sac-like with posterior canal

into which seminal duct opens ventrally. Endophallic duct long S- shaped,

basally entering apex of reservoir, apex spout-like.

Nezara viridula (Linnaeus), 1758

Described and figured by Pruthi (1925). Additional descriptions

and corrections given below.

Pygophore, dorsal border concave with a very narrow superior

ridge (fig. 155); ventral border (fig. 156) with a deep emargination.

Claspers (fig. 157) with inner surface finely scalloped.

Theca moderately long, cylindrical. One pair of small mem-

braneous conjunctival appendages (fig. 158), very short, broad (appar-

ently notnoted by Pruthi). Medianpenal lobes (fig. 159) semi-circular,

fused together into a broad U along ventral margins, enclosing apex of

vesica.

Vesica described and figured by Kumar (1964). Seminal duct

(fig. 160) merging into a wide funnel-shaped canal (conducting canal of

Kumar), canal narrowing, encircling posterior margin of ejaculatory

reservoir and opening dorsally. Ejaculatory reservoir oval; endophallic

duct short and slightly sinuous, basally merging with apex of reservoir.

Thyanta perditor (Fabricius), 1794

Pygophore somewhat globular, pygophoral opening small, facing

caudad; dorsal margin with a very narrow superior ridge (fig. 161).

Veatral border with a deep median V-shaped notch. Stout setae on

lateral margins.

Claspers C- shaped (fig. 162), upper margin straight, bearing a

number of fine setae, basal margin broad, stout, forming stem.

Theca balloon-like and acentric, two small knobs (fig. 163) one

on each side on dorsal surface near apex. One pair of conjunctival

appendages, dorsally produced into an oblong membraneous process,

basally appendage wide, apically tapering into a sclerotized horn. Median

penal lobes (fig. 164) cylindrical and curved in a U on either side of apex

of vesica, each with a pointed tooth apically, heavily sclerotized through-

out.

Vesica very similar in construction to Chlorocoris subrugosus , ejacu-

latory reservoir (fig. 165) very large, globular, endophallic duct narrow

and short.

Padaeus viduus ( Vollenhoven), 1868

Dorsal border of pygophore with a narrow superior ridge (fig.

McDonald

23

166); ventral margin flattened, border with a median V-shaped notch,

two ridges, one on each side, forming a wide V, run from outer angles

of ventral margin to centre of pygophore.

Claspers (fig. 167) flattened, basally wide, apically produced

into a blunt point, a line of fine scalloping running from apex a short

distance down inner surface.

Theca (fig. 168) oval in shape. Two pairs of conjunctival appen-

dages (fig. 169): fir st heavily sclerotized, rod-like; second bifid; dorsal

arm membraneous balloon-like; ventral arm apically sclerotized and

oblong in outline. Median penal lobes (fig. 170) fused into a horseshoe-

like shield surrounding apex of vesica (fig. 171).

Vesica consisting internally of a complicated series of ducts.

Seminal duct (fig. 172) passing ventrally into a long canal opening into

dorsal half of ejaculatory reservoir; latter divided into dor sal and ventral

ducts. Endophallic duct short, basally widening and merging into ventral

chamber .

Proxys punctulatus (Palis ot de Beauvois), 1805

Dorsal border of pygophore medianly with a well developed

superior ridge (fig. 173), laterally bearing a small oblong projection on

each side. Ventral margin flattened, border with a wide median U-shaped

emar gination, on either side of which is an oblong callus (fig. 174)

bearing a number of setae; running back from each callus is a pair of

well sclerotized ridges medianly forming between them a U-shaped

trough.

Claspers (fig. 175) with stout wide stem, produced apically into

a blunt cylindrical process, inner margin with a narrow band of scallop-

ing. Five or six small stout setae found on margin of stem.

Aedoeagus and vesica similar in most respects to those of

Padaeus viduus • Second conjunctival appendages (fig. 176) somewhat more

sclerotized than in P. viduus. The general similarity of the genitalia of

this species to that of P- viduus would suggest that these 2 genera are

closely related and that Padaeus should be placed in Proxys .

Neottiglossa trilineata (Kirby), 1837

Pygophore with dorsal border (fig. 177) evenly arched, laterally

bearing on each side a rounded lobe with a fringe of hair s. Ventral border

almost straight with a wide median emar gination, ventral margin below

border almost vertical.

Claspers (fig. 178) with stout oblong stem apically produced on

inner side into a flat plate and on outer side into a short blunt process.

Theca short, stout, bearing laterally two large knobs (fig. 179)

one on either side; apex of theca produced dorsally into a thecal shield

consisting of two pointed lobes with a wide U-shaped depression between

them. One pair of membraneous conjunctival appendages, apically

broadly rounded; dorsal to these is a large and voluminous conjunctival

lobe (not fully expanded in fig.). Median penal lobes sclerotized,

cylindrical, and curved inwards, fused basally along their ventral

margins and connected to sub-apex of vesica by two thickened arms one

on each side.

24

North American Pentatomoidea

Seminal duct (fig. 180) opening ventrally into a wide canal ex-

tending round base of reservoir and opening into a simple sac-like

ejaculatory reservoir . Endophallic duct short, slightly sinuous, entering

ejaculatory reservoir apically.

Murgantia histrionica (Hahn), 1834

Pygophor e with dorsal border (fig. 181) deeply arched and some-

what XL shaped; ventral border sinuous with a wide median U-shaped

concavity, two small processes borne one on either side of the ventral

border on the lateral margins. Proctiger box-like, centrally concave and

produced into two flattened median processes distally.

Claspers (fig. 182) flattened basally, apically narrowing into

blunt curved rods, no differentiation between stem and apex.

Theca oblong, bearing distally a large cylindrical thecal shield

(fig. 183). One pair of long cylindrical, membraneous conjunctival

appendages (not fully expanded in fig. ), apically blunt and moderately

sclerotized. Median penal lobes hook-like, basally fused to a common

stem dor so-medianly, apices connected by a small plate bearing a conical

cap (fig. 184) fitting over apex of vesica.

Ejaculatory reservoir (fig. 185) S- shaped, terminating in a closed

chamber. Seminal duct opening ventrally into base of endophallic duct,

latter proximally connected to apical chamber of ejaculatory reservoir ,

distally narrowing into a slightly curved duct, apically attached to median

penal lobes.

Eysarcoris aeneus (Scopoli), 1763

Pygophor e with dorsal border (fig. 186) sinuous; ventral border

with a deep median U-shaped emar gination, on each side of which is

triangular flattened area bearing a number of setae. Proctiger apically

with a median bilobed sclerotized process.

Claspers divided into two sections (figs. 187, 188), a broad

flattened diamond shaped blade apically acute and a semi-circular plat-

form attached to base of blade, a number of setae around margin; outer

edge finely scalloped.

Theca (fig. 189) conical, flattened laterally. Two pairs of con-

junctival appendages: first (fig. 190) membraneous, elongate, tube-like,

fused together basally, apically bluntly rounded; second bifid, consisting

of two heavily sclerotized, flattened, spatula-like appendages, fused

basally and tapering into a long pointed process.

Ejaculatory reservoir (fig. 191) small globose with a canal round

posterior mar gin into which seminal duct opens ventro-apically. Endo-

phallic duct long, sinuous, connected basally to apex of ejaculatory

reservoir .

Eysarcoris inter gressus (Uhler), 1893

Pygophore with dorsal border (fig. 192) evenly arched bearing

on either side of the mid-line a small triangular genital plate; ventral

border with an inferior ridge, centrally below the ventral border is a

shallow depression.

Claspers chisel-like (fig. 193) bilobed apically, a number of

McDonald

25

stout setae on outer surface.

Theca tubular (fig. 194). One pair of conjunctival appendages

(fig. 195) divided into two arms, a large ventral cylindrical membraneous

appendage, apically tapering to a sclerotized point; at base of this large

arm is a small dorsal cylindrical appendage, apically blunt. A pair of

rounded slightly sclerotized ventral conjunctival lobes present, fused to

common base of conjunctival appendages.

Vesica very similar to Cosmopepla bimaculata , endophallic duct more

loosely S- shaped (fig. 196).

Eysarcoris intergressus shows no similarity with the European species

E. aeneus studied but shows very great similarity to Cosmopepla in posses-

sing genital plates, chisel-like claspers and a very similar aedoeagus

and vesica. It is suggested that Eysarcoris should be placed in Cosmopepla.

The European species Eysarcoris aeneus possess no genital plates, very

peculiar claspers and the shape of the conjunctival appendages is quite

different; the vesica shows some similarity.

Cosmopepla bimaculata (Thomas), 1865

Pygophore (fig. 197) and claspers (fig. 198) described by Baker

(1931).

Theca cylindrical, somewhat curved when viewed laterally, two

small projections (fig. 199) one on each side near base of theca. One

pair of conjunctival appendages each divided into a large membraneous

cylindrical lobe, apically tapering to a blunt sclerotized point, and a

second small rounded sclerotized lobe, borne dor sally. A pair of small

rounded conjunctival lobes ventral to conjunctival appendages may

represent second conjunctival appendages. No median penal lobes.

Seminal duct (fig. 200) opening ventrally into a narrow canal

encircling posterior end of reservoir, and terminating dorsally. Endo—

phallic duct long, broadly S-shaped entering ejaculatory reservoir through

a groove formed by two sclerotized ridges on apex of reservoir.

Rhytidolomia senilis (Say), 1831

R hytidolomia viridicata (Walker), 1867

Rhytidolomia saucia (Say), 1831

Chlorochroa sayi (Stal), 1872

Chlorochroa ligata (Say), 1831

Chlorochroa uhleri (Stal), 1872

From a study of the male genitalia alone it is very clear that

these species are all very closely related to one another and should all

be included in the genus Rhytidolomia . This fact was suspected by Sailer

(1954) who also found that three of the species of Chlorochroa broke down

into a maze of intermediate populations. A thorough study is needed to

elucidate the validity of species included within the genus Rhytidolomia .

Rhytidolomia senilis

Dorsal border of pygophore with a broad median superior ridge

(fig. 201) passing down on each side round the base of the proctiger.

Ventral border gently concave with a trilobed inferior ridge. Ventral

and dorsal margins covered with fine setae.

Claspers (fig. 202) with a stout stem apically produced into three

26

North American Pentatomoidea

blunt lobes. A number of fine setae on outer and inner apical surfaces.

Theca cylindrical, flattened slightly laterally. One pair of

membraneous conjunctival appendages (fig. 203), basally broad, apically

broadly bilobed, a long membraneous blunt dorsal lobe at the base of

which is a ventral lobe with heavily sclerotized apical point. Median

penal lobes (fig. 204) club-like, basally fused along their dor salmargins.

Seminal duct (fig. 2 05), opening ventrally into a heavily sclerotized

canal, latter widening posteriorly and opening mid-dor sally into an oval

ejaculatory reservoir . Endophallic duct short, almost straight, merging

into apex of reservoir.

Rhytidolomia viridicata

Pygophore (fig. 206) and claspers (figs. 207, 208), very similar

to R. senilis , lobes of clasper somewhat more rounded.

Aedoeagus and vesica - similar to R. senilis , conjunctival appen-

dages divided into two completely membraneous lobes, no sclerotized

apical point present.

Rhytidolomia saucia

Pygophore (fig. 211) and claspers (fig. 212) similar to R. senilis a

shape of lobes at apex of claspers differs slightly, outer lobe acute.

Aedoeagus (fig. 213) and vesica (fig. 214) similar inmost respects

to those of R ■ senilis. However differences exist in the basal plates of

the theca, useable at a specific level.

Chlorochroa ligata

Dorsal border of pygophore with superior ridge (fig. 215) exten-

ding laterally on each side in an arc forming a small lateral projection.

Ventral border sinuous with a shallow median emargination.

Claspers apically trilobed (fig. 216) very similar to C. uhleri ,

slight differences in shape exist, however.

Aedoeagus and vesica, similar to those of Rhytidolomia senilis .

Chlorochroa uhleri Chlorochroa sayi

Pygophore (fig. 217), claspers (figs. 218, 219) and aedoeagus

described by Baker (1931). The genitalia of these species are similar

in all respects. Aedoegus and vesica similar to Rhytidolomia senilis,

Banasa dimidiata (Say), 1831

Dorsal border of pygophore with a wide superior ridge (fig. 220)

with a median emargination. Ventral border flattened bearing two double

knobbed processes one on each side of amedian square projection on the

border. Proctiger and margins of pygophore covered with fine setae.

Claspers (fig. 221) flattened, leaf-like, covered with fine setae.

Theca oblong, compressed laterally. One pair of membraneous

conjunctival appendages (fig. 222), broadly rounded apically. Median

penal lobes elongate, spatulate, apically free, broadly rounded, medianly

fused on the ventral surface, not enclosing apex of vesica.

Ejaculatory reservoir (fig. 223) oval, simple; seminal duct

opening into reservoir antero- ventrally; a canal extending from entrance

McDonald

27

of ductus seminis around posterior portion of reservoir to open antero-

dor sally. Endophallic duct short, slightly curved basally, connected to

apex of ejaculatory reservoir, apex of duct enclosed in a broad sclero-

tized sheath.

Loxa flavicollis (Drury), 1773

Pygophore with dorsal border (fig. 224) evenly arched, ventral

border deeply concave and flattened. An unusual pair of pygophoral

appendages (fig. 225) borne medianly, one on each side at the base of

dorsal margin; apex triangular in shape, with a broad concave surface,

produced into a long arm basally truncate, apex with long stout setae.

Claspers unusual (fig. 226), stem short, apex broad bearing

seven processes on outer margin, six blunt and oblong, apical process

produced into an acute point. Fine setae found over surface. Clasper

resembles a drilling bit when viewed apically.

Theca (fig. 277) small, acentric, with very large and well

developed basal plates; basal half of theca oblong becoming constricted

medianly and curving dorsad and produced into a semi-circular plate,

latter bearing a large bowl-like structure (thecal shield) with cr enulated

margins (fig. 228). From centre of thecal sheath a second sheath,

probably fused median penal lobes, surrounds apex of vesica. No con-

junctival appendages present.

Seminal duct (fig. 229) inserted ventrally into a heavily scler otized

canal, latter pas sing round base of ejaculatory reservoir and terminating

apically in a small 180° turn capped by a large sclerotized apodeme.

Ejaculatory reservoir very small, endophallic duct basally continuous

with reservoir.

Loxa flavicollis presents a most aberrant type of genitalia. It is

unique amongst specimens examined in possessing the unusual median

penal lobes, no conjunctival appendages, and the peculiar sheath developed

on the margin of the theca. The vesica is also extremely simple and

possesses an unusual pumping mechanism. On the basis of these distinct

peculiarities Loxa could be placed in a sub-tribe of its own, however this

would probably be better left till further work has been done on other

species of Loxa .

Menecles insertus (Say), 1831

Pygophore (figs. 230, 231) and claspers (fig. 232) described by

Baker (1931).

Theca vasiform with a pair of finger-like thecal processes (fig.

234). Two conjunctival appendages present, both membraneous and

fused to a wide common base: first (fig. 233) long thin cylindrical

structures, apically blunt; second short, broad, bearing five small

lobes. Median penal lobes (fig. 235) flattened and fused together forming

a deep narrow groove between lobes in which apex of vesica is situated.

Seminal duct (fig. 236) connected ventrally to wide base of endo—

phallic duct, latter merging posteriorly into a broad very heavily sclero-

tized ejaculatory reservoir with heavy striae on its lateral margins.

Endophallic duct anteriorly narrow, very long and coiled around itself

in a series of loops on right side of theca.

28

North American Pentatomoidea

Coenus delius (Say), 1831

Pygophore (fig. 237) and claspers (fig. 238) described by Baker

(1931).

Aedoeagus figured and described by Baker (1931), his lateral

penal lobes (= conjunctival appendages) were not fully expanded. Theca

vasiform with an apical overhanging rim, a pair of elongate finger -like

thecal processes (fig. 239) on dor sal mar gin of theca (titillator s of Baker).

One pair of conjunctival appendages (fig. 239), membraneous, apically

tapering and divided into two small blunt processes, one shorter than the

other. Median penal lobes (fig. 240) fused into a semi-circular disc-like

structure with a wide median groove, apex of vesica lying within this

groove.

Seminal duct (fig. 240) heavily sclerotized opening into a small

ventral sinus; latter merges posteriorly into a heavily sclerotized oblong

ejaculatory reservoir, lateral margins scored with a number of striae

running in a ventr o-dor sal direction. Endophallic duct basally united to

anterior sinus, as a fairly wide duct, then narrowing and looping to right

hand side of theca, passing through a wide circle to terminate within apex

of median penal lobes.

Hymenarcys nervosa (Say), 1832

Pygophore (fig. 241) and claspers (fig. 242) described by Baker

(1931).

Theca oblong bearing dor sally a pair of long cylindrical thecal

processes (fig. 244). One pair of conjunctival appendages (fig. 243),

membraneous, very broad and voluminous, apically terminating in a

short blunt point. Median penal lobes fused into a semi-circular flange

bearing a deep median groove.

Seminal duct (fig. 245) heavily sclerotized, connecting ventrally

into wide base of ejaculatory duct which posteriorly communicates with

dorsal ejaculatory reservoir . Latter flattened dor so- ventr ally, trilobed

dor sally and heavily sclerotized, lateral margins with a number of well

marked striae. Anteriorly endophallic duct narrows, twists sharply

twice passing to right side of theca, then looping in a large circle ter-

minates within groove between median penal lobes.

Euschistus tristigmus (Say), 1831

Pygophore (fig. 246) and claspers (fig. 247) described by Baker

(1931). Lower apical margin of clasper finely striated.

Aedoeagus described by Baker (1931). Theca oblong, with two

rounded processes (fig. 249) (titillator s. Baker, 1931) dorsally, one on

each side of the median line. Two pair s of conjunctival appendages (lateral

penal lobes of Baker) fused onto a commonmembraneous base: first (fig.

248) membraneous, wide basally, apically acute, slightly sclerotized;

second small, apically acute slightly sclerotized, possibly only one bifid

appendage represented since these appendages are not sharply divided

from one another. Median penal lobes (fig. 249) present, fused into a

flat semi-circular plate with a median dorsal groove.

Seminal duct (fig. 250) stout, opening ventrally into ejaculatory

reservoir where it is bent through 180°, passing along proximal end of

McDonald

29

reservoir to open into a dorsal chamber, latter connecting with a narrow

ventral chamber by means of a longitudinal slit-like aperture in septum

between dorsal and ventral chambers. Endophallic duct extremely long

arising from middle of ventral chamber in ejaculatory reservoir, exten-

ding forwards as a wide tube, bending through 90° and becoming enclos ed

by bases of median penal lobes for a short distance; then turning through

90° to pass a short distance ventrally and loop round to right hand side

of theca, from here looping in an S to finally pass in a wide circle ter-

minating apically on dorsal groove formed by median penal lobes.

Prionosoma podopioides (Uhler), 1863

Pygophore with dorsal border (fig. 251) rounded with a broad

superior ridge medianly extending down on each side of base of proctiger.

Ventral border laterally diffuse, medianly with a U-shaped groove on

either side of which is a slightly raised and rounded platform. A number

of long fine setae found along dorsal and ventral margins.

Claspers (fig. 252) stout, hook-shaped, stem wide, a number of

fine setae on inner margin of hook.

Theca small, oblong, dorsal margin with two long cylindrical

thecal processes (fig. 254). One pair of conjunctival appendages (fig.

253), basally wide membraneous, apically divided into two blunt lobes,

ventralmost one apically sclerotized. Median penal lobes narrow semi-

circular plates fused medianly forming a groove in which apex of vesica

rests .

Seminal duct (fig. 256) opening ventrally into a sclerotized canal

encircling proximal end of ejaculatory res ervoir , latter large, flattened

dor so- ventrally, heavily sclerotized and with a number of striae along

posterior half. Reservoir with a narrow more membraneous ventral

portion forming a duct merging postero-ventr ally into wide base of

endophallic duct (fig. 255); latter narrowing, looping round in three

small turns and finally coiling in a wide circle to apically terminate

between median penal lobes.

Halyini

Brochymena arborea (Say), 1825

Pygophore with dorsal border evenly arched (fig. 257), a wide

lateral flange found on either side bearing a patch of thick stout setae;

ventral margin concave with a deep median U-shaped emar gination.

Claspers described and figured byRuckes (1946). T-shaped (fig.

258) in outline, stem stout flattened laterally, apically somewhat abruptly

narrowed and bearing a cross-arm, situated in a dor so-ventral plane

when clasper at rest in pygophore; dorsal arm of T produced into a stout

hook, ventral arm blunt and shallowly bilobed apically. Base of stem

bearing a small cylindrical proces s bearing a number of long stout setae.

Theca cylindrical, elongate (fig. 259). One pair of broad mem-

braneous conjunctival appendages, apically produced into a blunt sclero-

tized point; a narrow band of sclerotization running from apex down inner

margin of appendages. Median penal lobes thin rod-like (fig. 260) basally

fused to a common stem not enclosing apex of vesica.

30

North American Pentatomoidea

Seminal duct (fig. 260) inserted ventr o-apically into a long canal

encircling posterior margin of ejaculatory reservoir and opening dorso-

apically into reservoir; latter large sac-like with a sclerotized apical

cap. Endophallic duct basally opening into apex of reservoir, long S-

shaped, apically free.

Brochymena quadripustulata (Fabricius), 1775

Pygophore figured by Crampton (1922). Dorsal border (fig. 262)

widely arched bearing medianly a narrow superior ridge; ventral border

(fig. 263) with a median V-shaped emargination in which the apex of the

pygophore rests. Numerous long fine setae along dorsal and ventral

margins.

Claspers G-shaped (fig. 264), stem stout, produced into a curved

hook, a number of long setae at base of hook.

Theca large cylindrical (fig. 265). One pair of conjunctival ap-

pendages, small membraneous on outer surface, slightly sclerotized on

inner surface, apically acutely pointed. Median penal lobes (fig. 266)

flattened oblong plates, basally fused to a common base not closely

associated with apex of vesica.

Vesica very similar to that of Brochymena arborea , shorter (fig. 267).

Edessini

Edessa bifida (Say), 1832

Pygophore with dorsal border widely arched (fig. 268); ventral

border gently concave, A pair of heavily sclerotized peg-like genital

plates present, one on each side laterally on dorsal border (fig. 269).

Claspers (fig. 270) with a broad stem merging into a triangular

spear-like head set at 45°, inner margin finely scalloped.

Theca (fig. 27 1) heavily sclerotized elongate and cylindrical. One

pair of very small sclerotized conjunctival appendages, broadly hook

shaped and fused to margin of theca.

Ejaculatory duct (fig. 272) consisting of a complicated series of

parallel ducts, their actual connections could not be worked out adequately

in whole mounts even with Kumar* s (1964) technique of introducing air

into these canals. Seminal duct entering vesica apically. Endophallic

duct short, curved. Sections will have to be made to work out the detailed

connections of the seminal duct and canals within the ejaculatory

reservoir .

Discocephatini

Lineostethus clypeatus (Stal), 1862

Dorsal margin of pygophore (fig. 273, 274) with two large rec-

tangular flaps one on either side of a median V-shaped depression.

Ventral border (fig. 275) sinuous produced on either side into two narrow

downwardly projecting flanges, apically acute and separated by a small

U-shaped emargination. Ventral surface below margin with a deep U-

shaped depression, a row of small stout setae found along proximal

margin.

McDonald

31

Claspers (figs. 276, 277) with a narrow stem, cylindrical, apically

expanded into a flat triangular plate terminating in a small heavily sclero-

tized spine; upper surface of blade finely scalloped.

Theca (fig. 278) pyriform with a small rim round apex. No

conjunctival appendages. Median penal lobes fused into an oblong

envelope-like structure enclosing basal two thirds of the endophallic duct.

Seminal duct (fig. 278) opening into a long canal extending round

base of ejaculatory reservoir and opening dorsally. Reservoir flask-

shaped, apically merging into endophallic duct; latter with a heavily

sclerotized collar at base; apex of endophallic duct protruding from slit

in median penal sheath. The heavily sclerotized collar at the base of the

endophallic duct may be some type of pumping device.

Sciocorini

Sciocoris microphthalmus (Flor), 1860

Pygophor e with dorsal border widely concave (fig. 279), laterally

bearing one on each side, a short stout spine; ventral border deeply

emarginate medianly produced into a short square process, ventral

margin flattened into a lip.

Claspers (fig. 280) with a narrow stem bearing apically a broad

spatulate plate bearing a dense mat of long setae.

Theca (fig. 281) elongate narrow and cylindrical, apically

expanded into a fan-like thecal shield. One pair of conjunctival appen-

dages, basally very broad membraneous, apically divided into two lobes,

a small ventral lobe with a small heavily sclerotized circular cap and a

larger dorsal lobe, apically moderately sclerotized, broadly rounded

and bearing a mat of stout spines (not fully expanded in diagram). Median

penal lobes (fig. 282) small curved horn-like, basally fused to a common

stem and enclosing apex of vesica.

Seminal duct inserted ventrally into a small anterior sinus (fig.

283); an S-shaped duct from sinus connects with a sac-like ejaculatory

reservoir. Ejaculatory duct short, broadly S-shaped, apically opening

between median penal lobes, basally continuous with anterior sinus.

Mecidini

Mecidea longula (Stal), 1854

Pygophore and claspers described and figured by Sailer (1952).

Aedoeagus figured by Sailer (1952). Theca large and cylindrical

(fig. 284). One pair of bag-like conjunctival appendages divided into

dorsal and ventral lobes, dorsal lobe apically bluntly pointed; ventral

lobes shorter than dorsal, apically produced into a sclerotized point.

Median penal lobes (fig. 285) stout cylindrical structures, apically broadly

rounded, basally fused together in a U around apex of vesica.

Ejaculatory reservoir (fig. 286) large saccular. Seminal duct

opening into a wide canal ventrally; latter situated on posterior margin

of reservoir and opening dor so-apically. Endophallic duct basally re-

cessed somewhat into apex of ejaculatory reservoir, apically tapering

into a short narrow duct.

32

North American Pentatomoidea

Pentatomidae- Asopinae

Zicrona caerulea (Linnaeus), 1758

The species examined by Baker (1931) under this name has sub-

sequently been found to be a new taxon of either specific or subspecific

rank.

Pygophore with dorsal border (fig. 287) medianly evenly arched

bearing two small processes one on each side dor so-laterally; ventral

border (fig. 288) sinuate, bearing medianly two distinct tufts of setae

borne on slight prominences . Genital plates crescent-shaped and bearing

along inner margin and upper surface a number of stout peg-like teeth.

Claspers (fig. 289) simple, blade-like, apically acute.

Theca small, tubular, produced distally into a large thecal shield

(fig. 290) with a deep V-shaped emargination ventrally. One pair of

membraneous conjunctival appendages, basally wide, apically bluntly

rounded, bearing a dorsal lobe. Median penal lobes (fig. 291) broad,

heavily sclerotized, basally fused.

Seminal duct (fig. 292) ventrally connected to a canal, latter

encircling posterior margin of reservoir and expanding into a small

chamber apically, incompletely separated from remainder of reservoir.

Endophallic duct long, sinuous, basally opening into ejaculatory reservoir

adjacent to seminal duct, apically terminating between median penal

lobes.

Baker (1931) stated that the genital plates were absent from the

species examined by him. They are in fact present, but are completely

smooth.

Oplomus tripustulatus (Fabricius), 1803

Pygophore with dorsal border (fig. 293) gently concave with a

shallow median emargination above base of proctiger. Ventral border

sinuous medianly; on ventral surface is a deep heart-shaped depression.

Genital plates oblong with an emargination on outer edge, surfaces ridged.

Claspers L- shaped (fig. 294) apical half blade-like, joined at

right angles to stout stem.

Theca small conical, anteriorly expanded into a thecal shield (fig.

295) almost twice as large as theca, ventral mar gin with a wide V-shaped

incision. Two pairs of voluminous membraneous conjunctival appen-

dages, both apically broadly rounded. Median penal lobes elongate plate-

like (fig. 296), medianly fused, distally each produced into a long process.

Ejaculatory reservoir (fig. 297) small, globose with a canal

encircling posterior surface into which seminal duct opens ventrally,

canal broadens dor sally into an oval chamber communicating with reser-

voir. Endophallic duct long, thin and sinuous, basally entering ejacu-

latory reservoir adjacent to seminal duct.

O

Heterosceloides lepida (Stal), 1862

Pygophore with dorsal margin (fig. 298) deeply concave; ventral

margin slightly sinuous. A large oval pit found below median section of

ventral border. Genital plates P- shaped, upper surface smooth.

Claspers (fig. 299) flattened, spatulate, apical margin straight.

McDonald

33

basally tapering to a short stout stem. Outer surface finely scalloped.

Theca small, conical, proximally produced into a thecal shield

(fig. 300) with deep V- shaped emargination on ventral surface . Two pairs

of conjunctival appendages: first membraneous, basally broad cylin-

drical, apically tapering to a blunt point; second smaller, cylindrical

and membraneous, attached to a large membraneous bag-like base.

Median penal lobes (fig. 301) small, flattened leaf-like structures,

medianly fused, basally free and produced into two long processes.

Vesica (fig. 30Z) very similar to that of Oplomus tripus tulatus , endo-

phallic duct with a U-shaped loop anteriorly.

Rhacognathus americanus (Stal), 1870

Dorsal and ventral borders of pygophore (fig. 303) evenly and

gently arched; genital plates small, top shaped; dorsal and ventral

margins slightly crenulate.

Claspers (fig. 304) small, stem short widening into a flattened

triangular apex.

Theca oblong (fig. 305), apically expanded into a thecal shield.

One pair of membraneous conjunctival appendages, basally broad, apex

sclerotized and acute. Median penal lobes (fig. 306) laterally oval in

outline, apically free disc-like, medianly fused bymeans of a cross-bar,

basally free and tapering.

Seminal duct (fig. 3 07) entering ventrally into a canal extending

round posterior mar gin of ejaculatory reservoir and opening into a small

chamber apically, latter incompletely divided by means of a septum from

ejaculatory reservoir. Endophallic duct looping in a wide U, opening

apically between median penal lobes, basally joining reservoir adjacent

to seminal duct.

Apateticus bracteatus (Fitch), 1856

Pygophore and claspers (fig. 308) described by Baker (1931).

Theca oblong, produced distally into a large thecal shield (fig.

309), lateral margins sharply pointed, dorsal margin W-shaped, ventral

margin broadly emarginate. One pair of very broad membraneous con-

junctival appendages (fig. 3 09), apically produced into a stout sclerotized

horn. Median penal lobes (fig. 310) oblong in shape medianly fused,

basally produced into two narrow processes connected centrally by a

membrane forming a hollow tube.

Vesica (fig. 311) very similar to that of Rhacognathus americanus but

apical chamber of ejaculatory reservoir somewhat larger.

Apateticus lineolatus (Her r ich-Schaeffer ), 1839

Dorsal border (fig. 31Z) of pygophore with deep V-shaped emar-

gination above proctiger; ventral mar gin sinuous bearing long fine setae,

lying beneath ventral margin is a large oval depr es sion part of a vertical

wall between ventral border and ventral surface of pygophore. Genital

plates oblong with a small pointed process on upper margin, surface

with three longitudinal ridges.

Claspers (fig. 313) C-shaped, blade-like, apex acute, stem

narrow.

34

North American Pentatomoidea

Theca (fig. 314) small, conical, distally with thecal shield,

latter rounded laterally and with a deep V-shaped cleft ventrally. One

pair of membraneous conjunctival appendages, apically slightly sclerotized

and produced into a small sharp point. Median penal lobes (fig. 315)

apically disc-like, flattened, medianly fused and distally each produced

into a long narrow process.

Vesica (fig. 316) very similar to that of Oplomus tripustulatus , ejacu-

latory duct somewhat shorter.

o

Podisus acutissimus (Stal), 1870

Pygophore with dor sal border (fig. 317) broadly arched, medianly

with a superior ridge bearing a small prominence (fig. 319) with a tuft

of long stout setae on each side. Ventral border (fig. 318) sinuous,

thickened medianly on either side of a central emar gination, below this

is a deep median depression. Dorsal and ventral margins covered with

long fine setae. Genital plates flat, triangular, inner margin broadly

scalloped with a row of scalloping behind.

Claspers L-shaped (fig. 320), stem short, stout with a broad blade

attached at right angles, apically acute and finely scalloped on lower

surface.

Theca small, oblong, anteriorly produced into a thecal shield (fig.

321), dor sally with a deep V-shaped emar gination. One pair of mem-

braneous conjunctival appendages, basally broad, apically tapering to a

blunt point. Median penal lobes thin, oblong, fused along ventral mar gins

forming a horseshoe-like structure around the apex of the vesica.

Ejaculatory reservoir (fig. 322) with a canal extending round

proximal end into an anterior chamber cut off from rest of reservoir by

an incomplete septum. Seminal duct and endophallic duct entering

ejaculatory reservoir adjacent to one another; endophallic duct short,

sinuous; seminal duct opening directly into canal.

Podisus maculiventris (Say), 1899

Pygophore and claspers (fig. 323) described by Baker (1931)

Theca small, oblong, with a large thecal shield (fig. 324) evenly

rounded laterally on apical margins, ventral margin V-shaped. One

pair of membraneous conjunctival appendages, apically tapering and

blunt. Median penal lobes (fig. 325) laterally flattened, disc- shaped,

basally fused.

Vesica (fig. 325) very similar in construction to Podisus acutissimus .

Alcaeorrhynchus grandis (Dallas), 1851

Dorsal border (fig. 326) of pygophore medianly evenly arched,

laterally strongly curved and thickened; ventral border with a deep median

U-shaped incision, beneath latter is an oval horizontal depression lined

with short stout setae. Genital plates oblong, emarginate on outer border,

inner margin broadly serrate.

Claspers (fig. 327) short, stout, stem apically broadened into a

flat plate, upper margin emarginate.

Theca short, compact, somewhat globose, heavily sclerotized,

dorsally bearing a thecal shield (fig. 328). Two pairs of conjunctival

McDonald

35

appendages: first membraneous, wide at base, apically tapering to a

heavily sclerotized point; second shorter, basally membraneous, fused

to base of first appendages, apically tapering and sclerotized. Median

penal lobes (fig. 329) heavily sclerotized, laterally flattened, apically

free, basally fused to a common stout stem.

Ejaculatory reservoir (fig. 330) globular; seminal duct and

endophallic duct entering ventrally adjacent to one another; latter long

narrow and sinuous, apically opening between lateral penal lobes.

Seminal duct opening into a canal extending round posterior margin of

reservoir, expanding apically into a small chamber incompletely separ-

ated from rest of reservoir.

Euthyrhynchus floridanus (Linnaeus), 1767

Dorsal border of pygophore (fig. 331) with a shallow median

emar gination, genital plates lying one on either side; latter elongate

narrow structures, inner margins crenulate. Ventral border with two

prominent projections on either side of a median U-shaped emar gination.

Below ventral margin is a deep groove. Stout setae found on lateral

corners of dorsal margin and ventral margin.

Claspers (fig. 332) with triangular stem, distally produced into

a broad flat blade, truncate apically.

Theca small conical bearing a large rounded thecal shield (fig.

333) deeply cleft ventrally, dorsal margin U-shaped. One pair of mem-

braneous bilobed conjunctival appendages, apices blunt; a median ventral

conjunctival lobe present. Median penal lobes flattened, oblong plates,

medianly united, basally each produced into a long process (fig. 334).

Vesica very similar to that of Alcaeorrhynchus grandis . Endophallic

duct long and sinuous.

Stiretrus anchorago (Fabricius), 1781

Described and figured by Pruthi (1925).

Pygophore with dorsal border (fig. 335) evenly arched, ventral

border with a wide median U-shaped emargination belowwhich is a deep

pit. Genital plates oblong, finely scalloped on their upper surfaces. A

number of long fine setae along ventral margin.

Claspers (fig. 336) with apices flattened and hastate, attached at

right angles to a slender stem.

Theca small bearing distally a large thecal shield (fig. 337) with

a wide U- shaped emar gination ventrally. One pair of bifid membraneous

conjunctival appendages, basally wide, apically produced into two small

narrow rounded processes. Median penal lobes (fig. 338) apically flat-

tened, disc-like, centrally united around apex of vesica, basally each

produced into a long process.

Vesica (fig. 339) very similar in construction to

Euthyrhynchus floridanus . Endophallic duct long, narrow and thrown into a

number of loops.

Mineus strigipes (Her rich-Schaeff er ), 1853

Dorsal border of pygophore. (fig. 340) evenly arched; ventral

border with two small projections on either side of shallow median

36

North American Pentatomoidea

emar gination, ventral margin vertical with a wide shallow depression

medianly. Genital plates large, oblong, dorsal and inner margins with

9-12 peg-like processes.

Apex of each clasper triangular (fig. 341), stem stout; apical

portion of clasper bentat approximately 120° to stem, upper surface flat

and bearing a series of minute scallopings.

Theca small oblong, bearing distally a thecal shield (fig. 342)

with rounded lateral margins, ventrally with a deep V-shaped incision

reaching margin of theca. One pair of conjunctival appendages, mem-

braneous, basally very broad, apically tapering to a blunt point. Median

penal lobes elongate, bluntly pointed apically, medianly fused around

apex of vesica, basally each lobe produced into a free process.

Vesica (fig. 343 ) very similar in construction

to that of Euthyrhynchus floridanus , endophallic duct short.

Perillus confluens (Her rich-Schaeffer ), 1839

No essential difference could be noted between the structure of

the pygophore, aedoeagus or vesica of this species and of Mineus strigipes .

It is probable that Mineus should be placed in Perillus , the type material

would have to be examined for final decision.

Andrallus spinidens (Fabricius), 1787

Dorsal border of pygophore (fig. 344) steeply concave and with a

superior ridge covering base of proctiger, at each end of superior ridge,

is a small globose genital plate with a number of small ridges on upper

surface.

Ventral border sinuous, lateral edges somewhat thickened and

bearing numerous long stout setae, latter also found on outer angles and

inner margin of dorsal border.

Claspers claw-shaped (fig. 345), apex acute, stem short and

flattened.

Theca small, narrow with a large thecal shield (fig. 346) one and

a half times as long as theca itself, enclosing conjunctival appendages,

apex of each slightly sclerotized, tapering to a blunt point; dorsal to

conjunctival appendages is a single membraneous conjunctival lobe.

Median penal lobes (figs. 347, 348) in lateral view somewhat oblong in

outline, medianly fused, apically free, flattened, basally produced into

two long tapering processes.

Vesica (fig. 348) very similar to that of Rhacognathus americanus .

Penfatomidae - Podopinae

Podopini

Amaurochrous cinctipes (Say), 1828

Previously described and figured by Barber and Sailer (1953).

Pygophore with dorsal border (fig. 349) evenly arched; laterally

bearing two large flattened appendages (figs. 349, 350). These pygophoral

appendages are the ^.popygeal appendages of Barber and Sailer (1953)

and parandria of Leston (1953). The appendages fit into grooves on

McDonald

37

lateral margins of pygophore, eachis bluntly rounded and bears a small

peg-like tooth on inner dorsal margin. Ventral border with median U-

shaped emar gination, on either side of which is a stout pointed process.

Claspers very characteristic (figs. 351, 35Z), consisting of a

lower platform and an upper curved hook. A number of long stout setae

on upper surface of platform and on outer surface of hook.

Theca very similar to asopine type, short, cylindrical and bearing

a large thecal shield (fig. 353) not developed on ventral margin. One

pair of membraneous bag-like conjunctival appendages. Median penal

lobes present; oblong flattened, heavily sclerotized plates (figs. 353,

354) lying on either side of apex of endophallic duct.

Seminal duct (fig. 355) opening ventrally into a canal extending

round posterior margin of large globose ejaculatory reservoir. Endo-

phallic duct continuous with apex of reservoir, short, apically widening

and opening between median penal lobes.

Amaurochrous dubius (Palisot de Beauvois), 1805

No difference could be noted between the genitalia of this species

and of A cinctipes strengthening the supposition made by Barber and Sailer

(1953) that A. cinctipes is conspecific with A dubius .

I Veda parvula (Van Duzee), 1904

Described and figured by Barber and Sailer (1953).

Pygophore with dorsal border (fig. 356) almost straight, lateral

margins bearing two large flap-like pygophoral appendages. Ventral

border with a shallow median emar gination on either side of which is a

small broadly rounded process.

Claspers (fig. 357) small, very similar to tho s e

of Amaurochrous cinctipes .

Theca small, cylindrical, bearing a large thecal shield (fig. 358).

One pair of membraneous balloon-like conjunctival appendages. Median

penal lobes heavily sclerotized somewhat broadly hook-shaped, flattened

laterally and lying on either side of vesica (fig. 359).

Ejaculatory reservoir (fig. 360) bulb-like, simple, apically

continuous with a short endophallic duct; a shallow canal extends from

ventro-apical entrance of seminal duct round posterior margin todorso-

apical region of ejaculatory reservoir; seminal duct opening into this

canal ventrally.

Oncozygia clavic ornis (Stal), 1872

Claspers and aedoeagus figured by Barber and Sailer (1953).

Pygophore (fig. 361) very similar to that of Weda parvula .

Claspers biramous (fig. 362) one arm forming a blunt process,

the other broadened into a flat platform bearing a fringe of long setae.

Stem very short, almost non-existent.

Theca small, cylindrical bearing distally a thecal shield (fig.

363). One pair of membraneous and balloon-like conjunctival appendages.

(See Sailer (1953) fig. 18, for expanded view of conjunctival appendages).

Median penal lobes (fig. 365) flattened plates fused into a horseshoe-like

structure (fig. 364) round apex of vesica.

38

North American Pentatomoidea

Ejaculatory reservoir (fig. 366) small, globular, with a canal extending

round posterior mar gin to dorso-apical half of reservoir. Seminal duct open-

ing ventrally into canal; endophallic duct short sinuous, apically terminating

between median penal lobes.

Tessaratomidae - Oncomermae

Piezosternum subulatum (Thunberg), 1783

Pygophore with dorsal border deeply concave (fig. 367), ventral mar gin

shallowly concave bearing medianly a heavily sclerotized oblong process.

Ventral surface of pygophore produced into a large scoop-like platform pro-

jecting some way beyond the ventral margin. Outer angles of this platform

with a small patch of short heavily sclerotized setae (fig. 368).

Claspers (fig. 369) simple spatulate, slightly curved. A number of long

stout setae on outer apical surface.

Theca (fig. 370) squat and somewhat lopsided being produced into shield-

like projection on ventral margin. Two conjunctival appendages: first heavily

sclerotized, oblong, thin and flap-like lying laterally at base of conjunctiva;

second lying above first, divided into two broadly rounded lobes, dorsal most

lobe enitrely membraneous, ventral most lobe lightly sclerotized.

Vesica consisting of a long membraneous tubular lobe, apically tapering

to a fine needle-like point. Internally vesica very complex consisting of an

ejaculatory reservoir (fig. 371) divided into dorsal and ventral chambers,

connected anteriorly by means of a short spiral duct (fig. 370) and posteriorly

througha wide canal. Dor sal chamber oval in outline attached directly to bases

of second conjunctival appendages; ventral chamber C- shaped. Seminal duct

continuous with apex of ventral chamber, extr emely long thin and highly coiled

tube, apically becoming straight and tapering into a very fine duct opening at

apex of vesica (fig. 372).

The genitalia of this species resemble very closely those of P. calidum

(Fabricius) described by Leston (1954), he states that there are three pairs of

conjunctival appendages in P. calidum but does not show them on his diagram.

Acanfhosomidae

Meadorus lateralis (Say), 1831

Pygophore (fig. 373) and claspers (fig. 374) described and figured by

Baker (1931). Theca (fig. 375) squat and tub- shaped. Two pair s of conjunctival

appendages: first flattened, leaf-like, slightly sclerotized; second (fig. 376)

acentric, consisting of three flattened leaf-like lobes arranged around vesica,

one lobe considerably longer than other two.

Seminal duct (fig. 377) opening ventrally into a globular ejaculatory

reservoir, latter bearing a pair of processes on apico-dorsal surface to which

bases of first conjunctival appendages are attached. Endophallic duct long,

narrow and looped in a wide S, apically tapering to a very fine thread-like

duct, basally merging with apex of ejaculatory reservoir.

Elasmoslethus cruciatus (Say), 1831

Pygophore (fig. 378) and claspers (fig. 379) described and figured by

Baker (1931).

Theca with a large rounded dorsal diverticulum (fig. 380) described as

ventral by Leston (1953) for Elasmostethus inters tine tus3 squat and tub-shaped. One

McDonald

39

pair of sclerotized, flattened and leaf-like conjunctival appendages, apically

acute.

Vesica consisting of a large membraneous cylindrical lobe bluntly

rounded and bearing a rounded median dorsal process. Opening of ejaculatory

duct diffuse, consisting of a small crenulated lobe about a third of the way up

on ventral surface (fig. 381). Ejaculatory reservoir (fig. 382) found at base of

vesical lobe, generally withdrawn into theca, globular and divided by means of

a septum into two chambers. Seminal duct opening into posterior chamber,

latter connected directly to anterior chamber. Endophallic duct long, looped

basally merging into apex of anterior chamber, apically widening and forming

a diffuse opening on ventral margin of vesica.

The genitalia of this species resemble very closely those of E. interstinctus

described and figured by Leston (1953).

Oydnidae - Corimeiaeninae

Corimelaenini

Corimelaena pulicaria (Germar), 1839

Pygophore with dorsal border diffuse medianly; ventral border almost

straight. Pygophoral opening small surrounded by a wide flange dor sally and

laterally.

Claspers very small chisel-like (fig. 384), a number of very small setae

along apical margin.

Theca (fig. 385) small squat and broad, bearing a pair of spiny processes

one on each side on dorsal surface near base. Laterally apical margin bears

a pair of thin flat wing-like appendages one on each side. Three pairs of con-

junctival appendages (not fully expanded in fig. 386): first moderately scle.ro-

tized, basally wide, tapering apically into a curved horn; second smaller,

moderately sclerotized, flattened, triangular in outline, apex blunt, outer

margins serrate; third chisel-like lying inside second, lightly sclerotized.

Vesica very simple. Seminal duct (fig. 387) connected ventrally to a

simple saccular ejaculatory reservoir . Endophallic duct short, curved, basally

merging with apex of reservoir.

Gydnidae-Gydnsnae

Sehirini

Sehirus cinctus (Palisot de Beauvois), 1805

Genitalia described by Froeschner (I960).

Dor sal border of pygophore (fig. 388) arched medianly, laterally sinuous;

ventral border gently concave . Pygophoral opening surrounded by a wide flange.

Claspers figured by Froeschner (I960, fig. 188). Stem slender, short,

bearing a narrow sickle- shaped blade; a tuft of long setae situated at base of

blade.

Theca long cylindrical basally membraneous, apically becoming lightly

sclerotized; two small elongate heavily sclerotized flanges (fig. 389) found

laterally one on each side. One pair of conjunctival appendages, membraneous

and bilobed.

Vesica very small lying at base of theca, consisting of a simple sac-like

ejaculatory reservoir (fig. 390)which apicallymer ges into a short straight ejacu-

latory duct. Seminal duct attached ventrally to base of endophallic duct, latter

opening at base of a median canal formed from bases of conjunctival appendages.

40

North American Pentatomoidea

The aedoeagus of this species bears no resemblance to that of

Sehirus sp. described by Pruthi (1925).

Cydnini

Pangaeus aethiops (Fabricius), 1787

Pygophore (fig. 391), previously figured and described by

Froeschner (I960).

Claspers peculiar in possessing two distinct sections (fig. 392);

clasper proper (fig. 393) flattened leaf-like, outer mar gin with a number

of long fine setae; attached to this dor sally is a tubular arm (figs. 392,

394) the function of which is unknown.

Theca (fig. 395) long, tubular, heavily sclerotized, dorsal margin

produced into a lip. Three pairs of conjunctival appendages: first very

small cylindrical membraneous; second fused into a membraneous tube

bearing apically a pair of keavily sclerotized pads (fig. 396); third when

fully inflated balloon- like, totally membraneous, apically produced into

a blunt finger -like process.

Vesica very heavily sclerotized. Seminal duct (fig. 397) connec-

ting into base of endophallic duct; a long highly convoluted duct extending

from entrance of seminal duct and merging posteriorly into a saccular

ejaculatory reservoir. Endophallic duct short, sinuous, basally a long

duct running above convoluted duct and opening into ejaculatory reservoir.

This type of vesica resembles closely the type found in the tribe Scutel-

leraria (Scutellerinae) .

Cyrtomenus crassus (Walker), 1867

Dorsal mar gin of pygophore broad, covered with fine setae (fig.

398). Ventralmargin widely U-shaped not connected with dor sal margin.

Claspers figured by Froeschner (I960). Broad flattened with a

small tooth apically (fig. 399); apical mar gin broadly impressed bearing

a large number of long fine setae; inner lateral margin withan oval area

finely scalloped.

Theca cylindrical (fig. 400) very heavily sclerotized. One pair

of short stout heavily sclerotized conjunctival appendages ventral to

vesica.

Vesica bearing a long thin tubular infravesicular process on

ventral surface (fig. 401); seminal duct opening ventrally into base of

endophallic duct, latter moderately long, straight, ensheathed in a stout

tapering tube. Ejaculatory reservoir flattened, tube-like, connected by

means of a short spiral duct to ejaculatory duct.

Melanaethus subglaber (Walker), 1867

Pygophore with dorsal border broadly arched (fig. 402), ventral

border gently concave. Pygophoral opening with a wide flange on dorsal

and lateral margins.

Claspers figured by Froeschner (I960, fig. 213), somewhat

triangular in outline with numerous hairs on thickened apical margin.

Theca (fig. 403) elongate, tubular. Two pairs of conjunctival

appendages: first basally membraneous, apically heavily sclerotized

McDonald

41

and blunt; ventral most appendages (probably third) (fig. 404) basally

fused, apically produced into two small broadly rounded lobes, sclero-

tized throughout.

Vesica small; seminal duct (fig. 405) connected ventr ally to base

of ejaculatory duct; latter a short straight tube surrounded by a stout

sheath, tapering apically; basally endophallic duct merging into an

unusual spiral ejaculatory reservoir.

Amnestini

Amnestus pallidus (Zimmer), 1910

Pygophore with dorsal border evenly arched (fig. 406), ventral

border concave. Pygophoral opening surrounded by a wide flange.

Claspers (fig. 407) with a short narrow stem, widening medianly,

apically produced into an acute point. A number of fine setae scattered

over outer surface of clasper.

Theca (fig. 408) small, membraneous, globose. Two pairs of

conjunctival appendages (fig. 409): first divided into two pairs of small

spikes, apically slightly sclerotized; second, large bag-like, lightly

sclerotized (probably balloon shaped when fully inflated). All appendages

attached to fairly voluminous conjunctiva.

Seminal duct (fig. 410) very fine, opening ventrally into a long

canal at apex of vesica; canal merging into an internal duct opening into

ejaculatory reservoir . Endophallic duct long, basally merging with apex

of central sinus.

DISCUSSION

The major work dealing with the male genitalia of North American

pentatomids is that of Baker (1931). However he dealt with Canadian

species of Pentatominae and Asopinae only. Recently, Lattin (1964) has

examined the male genitalia of all North American Scutellerinae and

thereby filled a large gap in our knowledge. Pruthi (1925) worked with

the world Hemiptera; his findings are of limited value in some taxa,

because only a very small number of species was examined. This gave

an inaccurate view of some groups. Several other workers have dealt

with the male genitalia of a small number of species of various families

within the Pentatomoidea. These papers have been considered in the

present study wherever relevant.

Pentatomidae -Scutellerinae

The male genitalia amongst species of North American scutel-

lerines are very varied and difficult to assess. The tribe Eurygastrini,

as constituted by Leston (1952), included three subtribes: Eurygastraria,

Odontoscelaria and Odontotar saria. Lattin (1964) has separated the

Eurygastraria from the rest of this group on the basis of the male genitalia

and accorded it tribal status.

The European species of Eurygastrini, possess very uniform

characters, Wagner (1963), Vidal (1949) and Piotrowski (1950). Most

42

North American Pentatomoidea

members of this tribe have the following features in common: T-shaped

claspers, two to three pairs of heavily sclerotized horn-like conjunctival

appendages and a cylindrical membraneous vesica. Unfortunately details

of the internal structure of the vesica have not been consider ed by other

workers in this field so far. The internal details of the vesica of

Eurygaster alternatus are definitely pentatomid (fig. 26) and do not resemble

the type found in the Scutelleraria. The ejaculatory reservoir is simple

and is connected directly via an anterior sinus to the seminal duct and

ejaculatory duct.

The remaining members of the tribe Eurygastrini are now in-

cluded in the Odontoscelini. The male genitalia of the four North American

species show remarkably little similarity to one another with the exception

of Euptychodera corrugata and F okkeria producta . This relationship can be seen

in figure 520, which is an analysis of eleven character differences found

inmales and females based on the method of James (1953). Three species

all possess stout spiny conjunctival appendages; the

vesica of Homaemus aeneifrons is, however, quite differ ent from the other two.

The tribe Pachycorini is represented in North America by ten

genera, of remarkably uniform character. They have two patches of

fine striae on abdominal sterna four to six, one on each side of the mid-

line. Outside of the New World only Hotea and Deroplax (Leston, 1952) and

female Tectocoris (Lattin, 1964) possess this character. Hotea and Deroplax

are central African in distribution; Tectocoris is Australian.

The male genitalia of this tribe show a remarkable array of

different types of structure. The analysis of character differences (fig.

52 0) shows that the species in this group are very

variable. Homaemus aeneifrons has been discussed above under Odontoscelini.

The remaining species show two trends as far as the structure of the

vesica is concerned. The ejaculatory reservoir is absent or very small

inmost; the second group generally has a large S- shaped ejaculatory

reservoir and in Stethaulax marmoratus and Camirus moestus a convoluted duct

typical of the Scutellerini. The Australian species Tectocoris diopthalmus

(McDonald, 1961) is quite aberrant in possessing a very small tube-like

ejaculatory reservoir . The conjunctival appendages vary in number from

one to three pairs, however the third when present is never heavily

sclerotized and S-shaped as in the Scutelleraria. Claspers are hook-

shaped, except in Symphylus carribeanus , where they are T-shaped.

Augocoris gomesii has very clear cut characters, possessing a con-

voluted thickened duct, hook-shaped claspers, sclerotized S-shaped third

conjunctival appendages and a short stout endophallic duct and is typical

of other members of the Scutellerini and quite distinct from other species

examined (fig. 520). Augocoris also shows very great similarity to

Australian members of this tribe in the structure of its aedoeagus

(McDonald, 1961, 1963) and vesica (Kumar, 1964).

Pentatomidae - Pentatominae

As stated in the introduction both Baker (1931) and Pruthi (1925)

dealt with this subfamily in some detail. In the tribe Pentatomini,

containing the vast majority of the species in North America, five species

were found to possess an enormously lengthened endophallic duct which

McDonald

43

is coiled like a watch spring. This type of genitalia was described by

Baker (1931) but not commented on. The other group containing all other

species studied has a relatively short endophallic duct.

All species possessing elongate coiled endophallic ducts have

several characters in common. All have a pair of dorsal thecal processes

(titillators of Baker, 1931), one or two membraneous conjunctival

appendages and a pair of heavily sclerotized median penal lobes fused

into a flat circular structure with a dor salmedian groove. The ejaculatory

reservoir is heavily sclerotized, consisting of two chambers divided by

means of an internal septum and with the exception of Euschistus tristigmus

bear a number of transverse striae on the sides.

The other group of species does not present such a uniform

picture. Claspers vary greatly and have many forms. Five species

were found to have a thecal shield (an asopine character) and of these,

Loxa flavicollis , has such peculiarly constructed claspers (fig. 226) and

aedoeagus (fig. 228) that its inclusion in this tribe is suspect. The

remaining four species all have one pair of membraneous conjunctival

appendages, a pair of median penal lobes and a simple sac-like ejaculatory

reservoir with a posterior canal, all characters possessed by the

Asopinae. However, none of these species have genital plates.

Carpocoris remotus , Dendrocoris humeralis , and Pentatoma rufipes all have

genital plates, one pair of membraneous conjunctival appendages and a

simple ejaculatory duct. However, all lack a thecal shield. Both these

groups thus connect the asopines very closely to the Pentatomini.

The other species examined in this grouping generally had one or

two conjunctival appendages varying greatly in shape. The ejaculatory

reservoir is simple, generally with the seminal duct opening into a

posterior canal. The endophallic duct varies greatly in its length and

shape. Median penal lobes, usually present, are absent from five species.

Piotrowski (1950), Kumar (1962) and Pruthi (1925) have all des-

cribed and figured the male genitalia of Pentatominae except for the

vesica. The structure of the aedoeagus varies . Kumar ( 1964) has figured

the vesicae separately of eight species of Pentatominae, their structure

agrees closely with those of North American species (one, Nezara viridula

occurs both in Australia and North America). Leston (1952) states that

the genitalia of Deroplax circumducta (Scutellerinae) are like the typical

Pentatomid genitalia in possessing a long thin vesica surrounded by the

conjunctiva. This is not correct since the type of the

family Pentatoma rufipes has a short stout vesica and only one genus so far

examined, Trichopepla semivittata , has anything like an elongate, thin vesica.

So far the only genera possessing the extraordinary elongated coiled

endophallic duct are North American (Baker 1931).

Investigation of the remaining tribes within the Pentatominae is

limited to single genera, and comments on these are therefore rather

speculative. The genitalia of the two species of Brochymena examined are

so similar to those found generally among the Pentatomini that the validity

of the Halyini is suspect.

The genitalia of the genus Mecidea were studied in detail by Sailer

(1952). The aedoeagus is very similar to that found among the Pentatomini

and is remarkably constant for the group in possessing two pairs of bag-

44

North American Pentatomoidea

like membraneous conjunctival appendages and a pair of median penal

lobes. The vesica is very simple in construction and resembles the

general pattern found among the majority of Pentatomini. Once again on

the basis of genitalia the elevation of this genus to tribal level is unwar-

ranted.

The remaining tribes, Edessini, Discocephelini and Sciocorini

all show certain characteristics peculiar to the species studied. Until

more work has been done, little can be said on the status of these tribes

except that they all share characters with the Pentatomini.

Pentatomidae - Asopinae

Baker (1931) described the Canadian species of this subfamily.

The following characters are common to all the species examined by

myself and to those described in the literature.

1. Pygophor e with a pair of genital plates on the dorsal margin,

one on each side. 2. Theca with apical margin developed into a thecal

shield. 3. Conjunctival appendages variable in number but always mem-

braneous. 4. Median penal lobes present and enclosing the apex of the

vesica. 5. Ejaculatory reservoir simple with seminal duct entering a

posterior canal. Endophallic duct and seminal duct enter reservoir

adjacent to one another.

The male genitalia show remarkable constancy in this group.

The Asopinae are differentiated on minor character differences

externally, but can now, on the basis of the male genitalia, be very clearly

defined. The general structure of the male genitalia is similar to that

found in the Pentatomini discussed above.

Leston (1954a) describes and figures the genitalia of Afrius figuratus

(Germar) a species from Africa which also clearly possesses the charac-

ters set out above. The genital plates are termed dorsal processes by

Leston. The status of the Asopinae will be discussed later.

Pentatomidae - Podopinae

The North American species were revised by Barber and Sailer

(1953). The aedoeagus of four species is figured, but not the internal

structure of the vesica. The genitalia of this subfamily are rather uni-

form; the following characters were found to be common to all species

so far examined. 1. Lateral margins of pygophor e with a pair of

appendages. 2. Theca with a thecal shield. 3. One pair of membraneous

conjunctival appendages. 4. A pair of median penal lobes. 5. Ejacu-

latory reservoir simple, with a posterior canal.

Leston (1953a) described the genitalia of Podops inuncta (Fabricius)

and they fit the general pattern found among North American species.

The Podopinae are very closely related to the Asopinae, the former

subfamily lacking genital plates. Their place seems to have been taken

by the pygophoral appendages.

The Podopinae like the Asopinae are thought to be closely related

to the Pentatominae. Leston (1953a) noted this whenhe raised this group

to subfamily status.

McDonald

45

Tessarafomidae

Leston (1954c, 1954d, 1957) and Pruthi (1925) have both described

the male genitalia of this family. Only one species Piezosternum subulatum ,

is described here. The genitalia of this species and of Piezosternum calidum

(Fabricius) (Leston 1954c), an African species, are very similar. Both

possess very long and highly convoluted endophallic ducts (the vesica

does not appear to be fully expanded in Leston's diagram), and one pair

of heavily sclerotized conjunctival appendages. Elizabetha courteauxi

Schouteden (Leston, 1954c) and Phyllocoris acuta Jeannel also have very long

endophallic ducts. However the Australian species Musgravea sulciventris

(Stal) and Rhoecocoris australasiae ( Westwood) do not have elongate endophallic

ducts (Leston, 1957). Kumar (1964) studied the vesicae of four Australian

t e s s a r a t o m i d s . None have the elongate endophallic

duct of Piezosternum subulatum • all, however, including Piezosternum , have a

complicated series of canals within the ejaculatory reservoir (conducting

chamber of Kumar). It would appear on the basis of the male genitalia

that the subfamily Oncomerinae should be split into two or more sub-

families. Leston (1955) suggested that Piezosternum might have to be

removed from the Oncomerinae.

Leston (1954d) described the genitalia of Tessaratoma papiZZosa(Drury).

These agree closely with Tessaratoma sp. figured by Pruthi (1925). The

endophallic duct is short and two pairs of membraneous conjunctival

appendages are present relating this tes saratomine to the Australian

species of Oncomerinae. Other species figured and described by Pruthi

(1925) from the Eustheninae show close similarities to the Tessaratominae

but not to Piezosternum ,

Acanthosomidae

Very little work has been done on the male genitalia. Leston

(1953b) describes and figures Cyphostethus tristriatus (Fabricius)and

Elasmostethus interstinctus (Linnaeus). Only two species were examined in the

present work, Meadorus lateralis and Elasmostethus cruciatus . All species with

the exception of Meadorus lateralis have a peculiar dorsal diverticulum on

the theca and all have at least one pair of flattened sclerotized conjunctival

appendages. However Cyphostethus and Meadorus have elongate whip-like

endophallic ducts whereas both species of Elasmostethus have rather apically

diffuse endophallic ducts. The ejaculatory reservoir in both species

examined by me were pentatmoid in construction, being simple sacs

with a dorsal canal, the endophallic duct passing out apically. Leston

did not, unfortunately, examine the internal structure of the vesica of

the two specimens he describes.

Cydnidae

A very extensive study was made on the European species of this

group by Wagner (1963). The present study is rather cursory and will

indicate certain trends among species of the North American fauna.

Wagner did not consider the vesica, so no comparisons of this structure

can be made.

46

North American Pentatomoidea

Cydnidae-Corimelaeninae

The pygophore of Corimelaena pulicaria } the only species of this sub~

family examined, resembles that described by Wagner (1963) for

Corimelaena scarabaeoides . The aedoeagus of this latter species differs from

that of C. pulicaria in possessing only two pairs of heavily sclerotized

horn-like conjunctival appendages (Wagner's spicula). Three pairs of

conjunctival appendages were found in C. pulicaria (fig. 386) and were quite

different in shape from those of C. scarabaeoides . McAtee and Malloch (1933)

figured the aedoeagus of twelve corimelaenines . All possessed two to

three pairs of stout sclerotized appendages, five possessed the wing-like

appendages on the margin of the theca.

Cydnidae-Gydninae

Wagner (1963) recognizes two major types of genitalia in this

subfamily, the Geotomus type and the Sehirus type. One species only,

Sehirus cinctus , of the tribe Sehirini exists in North America. On examin-

ation, the male genitalia of this species proved to be quite different from

any of the genitalia described by Wagner for species in this tribe. The

European species all possessed at least one pair of heavily sclerotized

conjunctival appendages, generally rod-like. The clasper s show striking

similarity being somewhat Y-shaped. Sehirus cinctus has only one pair of

membraneous conjunctival appendages (fig. 389) and the clasper s are

large and sickle-shaped. I note here that thevesicaeof Sehirus cinctus and

Corimelaena pulicaria are very similar.

Froeschner (I960) recognized Amnestus as a separate subfamily.

One species, Amnestus pallidus , Zimmer was examined. The vesica is

unusual in possessing a very long canal into which the seminal duct opens

apically (fig. 410); the canal passes back into the ejaculatory reservoir .

Wagner's (1963) Geotomus type genitalia characterized by the

possession of two pair s of conjunctival appendages, and a moderately long

endophallic duct, although he notes that the genus Cydnus is aberrant. The

three species of North American Cydnini studied, show great variation

in the aedoeagus and vesica. The number of conjunctival appendages

varied from one pair in Cyrtomenus (fig. 400) to three in Melanaethus (fig.

403). The vesica of Pangaeus aethiops is very similar to that found in the

Scutellerini, in possessing a long convoluted duct. Aethus indicus and

Geotomus apicalis (described by Kumar 1962) have an infra-vesicular process

also found in Cyrtomenus crassus (fig. 401). The structure of the ejaculatory

reservoir and associated ducts is very similar both in Geotomus apicalisa.n&

Cyrtomenus crassus . Aethus indicus was shown tohave three pairs of conjunc-

tival appendages (Kumar 1962) and a convoluted duct, characters shared

in common with Pangaeus aethiops. However, the latter species lacks the

infra-vesicular proces s. Wagner (1963) figures an extremely long coiled

endophallic duct for Chilocoris spp. and Cydnus aterrimus Forst. In the latter

species the duct is coiled at the base of the theca. This condition was

not observed in any of the North American species examined. It is

unfortunate that Wagner (1963) did not deal with the structure of the

ejaculatory reservoir. From his work itwould appear that the Sehirini,

except for the North American species, is a good grouping and resembles

the Eurygastrini (Scutellerinae) in the structure of the aedoeagus . Leston

McDonald

47

(1954b) describes the genitalia of Sehirus bicolor and in another paper Leston

(1956) describes two species of Dismegistus . All these species possess

three pairs of conjunctival appendages, the third lightly sclerotized, and

an endophallic duct projecting well beyond the mar gin of the theca. Wagner

(1963) apparently only found two conjunctival appendages (spicula) in

members of this tribe. It is clear that further workis required on species

included in this tribe. Sehirus cinctus (fig. 389) has only one pair of mem-

braneous bifid appendages and a very short vesica not projecting beyond

the margin of the theca and the aedoeagus in no way resembles that of

Sehirus bicolor . It would appear that on the basis of the male genitalia

Sehirus cinctus is wrongly placed with the Old World species.

MORPHOLOGY OF FEMALE GENITALIA

The female genitalia are not as complex as those of the male.

Detailed diagrams and descriptions are not given for each species since

the genitalia generally vary in broad characteristics only. Scudder (1959)

has described the genitalia of this group fully and any divergence from

his general descriptions has beennoted. The spermatheca of each species

was studied in more detail and provides some useful characters which

give some good clues to the relationships of the various groups.

The female genitalia are situated on abdominal segments eight and

nine and are of the plate-shaped type with a posterior or postero-ventral

aspect. The paratergites of segments eight and nine, together with the

first gonocoxae (segment 8), form the major part of the external genitalia.

The second gonocoxae (segment 9) generally forma bridge-like sclerite

beneath sternum 10. The gonapophyses attached to the gonocoxae are

generally membraneous. The gonangulum is fused posteriorly to ter gum

9. In some species the dorsal edge of the first gonapophysis is heavily

sclerotized and forms the grooved outer ramus. The ventral edge of the

second gonapophysis is in some species also heavily sclerotized and forms

the inner ramus.

Pentatomidae- Scute! lerinae

Odontoscelini

Genitalia externally plate-like, very similar in all species.

Descriptions are given by several authors, detailed descriptions will

not be included here.

Fokkeria pro due ta [Y an Duzee), 1904. (figs. 411, 412)

Euptychodera corrugata (Van Duzee), 1904

Genitalia of these two species almost identical. Genital chamber

with a deep median sclerotized groove (fig. 413) at dorsal end of which

is a small membraneous pouch, into which spermatheca opens. Sperma-

thecal duct long, leading into a pumping region, poorly defined from

spermathecal bulb, proximal flange of pump developed (fig. 414), lightly

sclerotized. The shape of the spermathecal bulb differs in the two

species, being somewhat more elongate in Fokkeria than in Euptychodera .

48

North American Pentatomoidea

Vanduzeeina balli (Van Duzee), 1904

Very similar (fig. 415) to Fokkeria producta 9 spermathecal duct long

(fig. 416), membraneous, spermathecal bulb elongate cylindrical.

Phimodera binotata (Say), 1824

Entrance of spermatheca into genital chamber surrounded by a

circular sclerite (fig. 418); a short groove extending along base of

chamber from this sclerite.

Spermathecal duct short, opening into a large tough sac-like

dilation (fig. 417); from latter a short duct connects to pumping region

with proximal flange only developed. Spermathecal bulb dumb-bell shaped.

Eurygastrini

Eurygaster alternata (Say), 1828

External genitalia flattened and facing ventrad, similar to Penta-

tomine type. Internally a pair of sclerotized interlocking rami present,

similar to those found in Scuteller ini (McDonald 1963). Second gonocoxae

lightly sclerotized elongate plates, not fused centrally. Genital chamber

with a long sclerotized groove (fig. 419).

Spermathecal duct short; pumping region with flanges indicated

only by slight swelling for muscle attachment; spermathecal bulb

spherical, separated from pump by a short duct. This species is quite

distinct in possessing sclerotized rami.

Pachycorini

The external genitalia are essentially very similar. Minor

differences exist among species and these are described.

Pachycoris torridus (Scopoli), 1722

Visible portion of first gonocoxae reduced, bases hidden beneath

seventh sternum. Opening of spermatheca (fig. 420) into genital chamber

surrounded by a heart shaped sclerite; a deep median sclerotized groove

extending along length of genital chamber from this sclerite. Sperma-

thecal duct with a large spherical dilation (fig. 421) pumping region small

with distal and proximal flanges developed, spermathecal bulb elongate

cylindrical.

Diolcus irroratus (Fabricius), 1775

Genital chamber with a narrow, heavily sclerotized groove (fig.

423), anteriorly opening into a heavily sclerotized pouch (fig. 442), into

which spermatheca opens; spermathecal duct short, stout, dilating into

a thick walled chamber (fig. 424) from which apically a short membraneous

duct leading to a pump, with proximal flange only developed; spermathecal

bulb elongate cylindrical.

Tetyra antillarum (Kirkaldy), 1909

First gonocoxae (fig. 425) each with a large sclerotized base

projecting internally beneath sternum seven. A large anchor-shaped

sclerite (fig. 426) present around opening of spermatheca into genital

McDonald

49

chamber. Spermathecal duct narrow, membraneous, with a large mem-

braneous sac-like diverticulum, pumping region small, proximal and

distal flanges (fig. 427) developed; spermathecal bulb globose connected

by a short duct to pump.

Symphylus carribeanus (Kirkaldy), 1909

A long, heavily sclerotized plate-like sclerite extending along

base of genital chamber (fig. 428) from entrance of spermatheca. Sper-

mathecal duct basally broad expanding into a globular dilation, from which

a narrow duct connects to pumping region; latter with both flanges

developed and connected by means of a moderately long, stout duct to a

spherical spermathecal bulb.

Sphyrocoris obliquus (Germar), 1839

Very similar to Homaemus aeneifrons .

Dilation smaller, pumping region not clearly differentiated (fig.

429), proximal flange only developed. Spermathecal bulb continuous with

pump, elongate cylindrical (fig. 430).

Homaemus aeneifrons (Say), 1824

Sclerotized groove (fig. 431) presentin base of genital chamber.

Spermathecal duct marked by numerous annulations; pumping region

(fig. 432) poorly defined, proximal flange present, membraneous; distal

flange missing; spermathecal bulb elongate, S- shaped.

Acantholomidea porosa (Germar), 1839

Eighth paratergites absent, ninth narrow and elongate (fig. 433).

Spermatheca opening into base of heavily sclerotized groove (fig. 434)

lying in base of genital chamber; a sac-like spermathecal diverticulum

also opening into this groove adjacent to spermathecal entrance. Sperma-

thecal duct medianlywith a sac-like dilation, pumping region small, with

proximal flange only developed; spermathecal bulb elongate and rod-like

(fig. 435).

C he lysomidea guttata (Her r ich-Schaeffer ), 1839

First gonocoxae (fig. 436) triangular, smaller than other species

examined in this tribe; internally a pair of sclerotized outer rami (fig.

436) present. Spermatheca opening into a pouch (fig. 437) with a heavily

sclerotized groove. Spermatheca minute, duct long, narrow passing to

a small pumping region (fig. 438), proximal flange reduced, distal flange

present, spermathecal bulb elongate sausage-like.

This species is distinct from other members of this tribe in

possessing one pair of rami.

Stethaulax marmoratus (Say), 1831

A long heavily sclerotized groove extending along base of genital

chamber (fig. 440) from spermathecal opening. Spermathecal duct long,

with a large saccular diverticulum (fig. 441) attached mid-way; pumping

region small (fig. 442), proximal flange well developed, distal flange

very small; spermathecal bulb oval connected to pump by a short duct.

50

North American Pentatomoidea

Scutellerini

Augocoris gomesii (Burmeister), 1835

External genitalia plate-like, typically Scutellerine (Scudder,

1959). Sclerotized and interlocking rami present.

Spermatheca typical for members of this tribe. Spermathecal

duct (fig. 443) medianly expanded into a heavily sclerotized globular

chamber with a series of fine markings externally; pumping region well

developed connected to spermathecal dilation by a short duct; sperma-

thecal bulb elongate apically expanded into a spherical bulb. This species

is very similar to other members of this tribe in possessing sclerotized

rami (Scudder 1959, McDonald 1963) and the large sclerotized median

dilation of the spermathecal duct (Pender grast 1957).

Pentatomidae - Penfatominae

The external genitalia are all very similar in this sub-family and

are described by Scudder (1959). The presence or absence of spiracles

on the eighth paratergites varies from species to species. Sclerotized

rami are lacking, ring sclerites were found in the two European species

studied, P entatoma rufipes , and Eysarcoris aeneus .

The spermatheca has been described by Pender grast for several

species and is remarkably constant. The spermathecal duct is expanded

into a large elongate balloon-like dilation (fig. 474) down the centre of

which is a sclerotized rod varying in thicknes s fr om species to species.

The apeK of this rod is free and a narrow channel extends down the centre

and basally emerges from the diverticulum as a narrow duct connecting

with the pumping region and spermathecal bulb. The pumping region has

well defined proximal and distal flanges for the insertion of muscles and

is attached directly to the spermathecal bulb. The shape of the latter

varies somewhat but in the majority of species is spherical or oval. One

exception only to this general pattern was found, in Trichopepla semivittata

the spermatheca consists of a long duct terminating in a membraneous

sac with no differentiation of pumping region or bulb (fig. 476).

Any variation from the general pattern described above will be

noted under each specific description.

Pentatomini

Rhytidolomia senilis ( Say), 1831. (figs. 444, 445)

Rhytidolomia viridicata (Walker) , 1867. (fig. 446)

Rhytidolomia saucia (Say), 1831

Chlorochroa ligata (Say), 1831

Eighth paratergites with spiracles. Second gonocoxae fused into

a single plate. Second gonapophyses found above the spermathecal en-

trance (fig. 444) and two small sclerites surround the opening of the

spermatheca into the genital chamber. Spermatheca as described above.

Banasa dimidiata (Say), 1831. (fig. 447)

Carpocoris remotus (Horvath), 1907. (fig. 448)

Murgantia histrionica (Hahn), 1834. (figs. 449, 450)

McDonald

51

Padaeus viduus ( Vollenhoven) , 1868

.Eighth paratergites with spiracles. Two small sclerites surroun-

ding opening of spermatheca. Incase of Mur gantia histrionic a , these sclerites

are somewhat larger (fig. 450), the ventralmost sclerite forming a plat-

form and the smaller dorsal sclerite forming a spout round the sperma-

thecal opening. Spermatheca normal, shape of spermathecal bulb varies

from species to species.

Mormidea lugens (Fabricius), 1775

Euschistus trisligmus ( Say), 1831. (fig. 452)

Hymenarcys nervosa (Say), 1832

Cosmopepla bimaculata (Thomas), 1865. (figs. 453, 454)

Menecles insertus (Say), 1831

Brepholoxa heidemanni (Van Duzee), 1904. (figs. 455, 456)

Dendrocoris humeralis (Uhler), 1877. (fig. 457)

Coenus delius (Say) , 1831

Eysarcoris intergressus (Uhler), 1893

Prionosoma podopioides (Uhler), 1863. (figs. 458, 459)

Solubea pugnax (Fabricius), 1775. (figs. 460, 461)

Eighth paratergites without spiracles. Entrance of spermatheca

surrounded by one or two small sclerites. Spermatheca as described

under general heading, shape of spermathecal bulb varies in each species

as does the size and shape of the flanges of the pumping region.

Neottiglossa trilineata (Kirby), 1837. (fig. 462)

Loxa flavicollis { Drury), 1773. (fig. 463)

Nezara viridula (Linnaeus), 1758

Arvelius albopunctatus (DeGeer) , 1773. (figs. 464, 465)

Aelia americana (Dallas), 1851. (fig. 466)

Acrosternum pennsylvanicum (DeGeer), 1773. (figs. 467, 468)

Peribalus limbolarius (Mulsant and Rey), 1866. (figs. 469, 470)

Vulsirea violacea (Fabricius), 1803. (figs. 471, 472)

Pentatoma rufipes ( Linnaeus^), 1758. (figs. 473, 474)

Chlorocoris subrugosus (Stal), 1872. (fig. 475)

All the above species are characterized by the fact that the sper-

mathecal bulb has from 2-4 hollow horn-like processes (fig. 474) on

it, these vary in size and shape being long and slender in Chlorocoris subrugosus }

small and squat in Peribalus limbolarius . Nezara viridula has been described

and figuredby Pendergrast (1957). The spermatheca is otherwise normal

in possessing a median dilation with central sclerotized rod, spermathecal

opening surrounded by one or two sclerites. Pentatoma rufipes has in

addition ring sclerites , this species is palaearctic in distribution.

Spiracles may be present or absent.

P eribalus limbolarius (fig. 461), Nezara viridula , Aelia americana (fig. 466),

Acrosternum pennsylvanicum (fig. 468) and Pentatoma rufipes all possess two pro-

cesses on the spermathecal bulb.

N eottiglossa trilineata ) Chlorocoris subrugosus 3 Loxa flavicollis (fig. 463) and

Arvelius albopunctatus possess three appendages. Vulsirea violacea has four

appendages .

The function of these processes is unknown.

52

North American Pentatomoidea

Trichopepla semivittata (Say), 1832

Eighth paratergites with spiracles. First gonapophyses sclero-

tized. Spermatheca simple consisting of a narrow duct terminating in

a simple membraneous sac (fig. 476), no pumping region present.

Proxys punctulatus (Palisot de Beauvois), 1805

Eighth paratergites without spiracles.

Spermathecal dilation constricted anteriorly (fig. 477) giving it

a bottle shape, otherwise spermatheca similar to standard description.

Thyanta perditor (Fabricius), 1794

Eighth paratergites with spiracles. A small circular sclerite

surrounding opening of spermatheca. Spermathecal dilation (fig. 478)

elongate bearing proximally a bulbous cap within which sclerotized rod

expanded into a bell shaped apex. Proximal to pumping region, duct

swollen into a sclerotized bulb (fig. 479) with a number of transverse

ridges, spermathecal bulb very elongate rod-like.

Eysarcoris aeneus (Scopoli), 1763

Eighth paratergites without spiracles. Two ring sclerites (fig.

480) present one on either side of a V-shaped sclerite surrounding

spermathecal opening. Spermathecal dilation constricted proodlmally

forming a large distal chamber and a smaller more elongate proximal

chamber, spermatheca otherwise normal.

This is a European species and shows marked differences from

the American species Eysarcoris intergressus as noted in the description of

the male genitalia.

Halyini

Brochymena quadripustulata (Fabricius), 1775

Brochymena arborea (Say), 1825

Eighth paratergites with spiracles. First gonapophyses sclero-

tized; second gonocoxae fused plate-like. Opening of spermatheca

surrounded by two sclerites (fig. 481). Spermathecal bulb with two pro-

cesses in Brochymena quadripustulata ; B. arborea with an additional small third

appendage. Spermatheca in other respects similar to general description

under Pentatomini.

Edessini

Edessa bifida (Say), 1832

Eighth paratergites with spiracles; second gonocoxae fused.

Spermatheca similar to Brochymena quadripustulata spermathecal bulb with

three processes of equal size (fig. 482)

Sciocorini

Sciocoris microphthalmus (Flor), I860

Eighth paratergites without spiracles, external genitalia typically

McDonald

53

Pentatomine in character. Base of spermathecal duct surrounded by a

small horseshoe shaped sclerite (fig. 483) with a second crescent shaped

sclerite in front of it, spermatheca as described under Pentatomini.

Dis c oc ephalini

Lineostethus clyveatus (Stal), 1862

Eighth paratergites with spiracle s* Ninth paratergites small

oval structures; second gonocoxae fused, narrow. Entrance of sperma-

theca surrounded by a small circular sclerite, otherwise similar to

Sciocoris microphthalmus , duct from spermathecal dilation to pumping region

wider, longer and convoluted (fig. 484).

Mecidini

Mecidea longula (Stal), 1854

Eighth paratergites with spiracles, ninth vertical, projecting

beyond posterior margin (fig. 485); spermatheca similar

to Sciocoris microphthalmus .

Pentatomidae - Asopinae

Mineus strigipes (Her rich-Schaeffer ), 1853. (figs. 486, 487)

Rhacognathus americanus (Stal), 1870. (fig. 488)

Oplomus tripus tulatus (Fabricius) , 1803

AndralLus spinidens (Fabricius), 1787

Podisus acutissimus (Stal), 1870. (figs. 489, 490)

Podisus maculventris (Say), 1899. (fig* 491)

Apateticus lineolatus (Her rich-Schaeffer ) , 1839. (fig. 492)

Stiretrus anchorago (Fabricius), 1781

H eterosceloides lepida (Stal), 1862

Perillus confluens (Herrich-Schaeffer), 1839. (fig. 493)

Alcaeorrhyncus grandis (Dallas), 1851. (fig. 494)

Euthyrhynchus floridanus (Linnaeus) , 1767, (fig. 495)

Zicrona caerulea (Linnaeus), 1758. (figs. 496, 497)

All the above species were examined and present a remarkably

uniform picture in the structure of the female genitalia and agree with

the general description given by Scudder (1959) for Pentatomidae.

The spermathecae were also extremely uniform and resemble

Hoploxys coeruleus Dallas figured by Pendergrast (1956). Minor variations

in the size and shape of the spermathecal bulb were found. Eighth para-

tergites with spiracles, second gonocoxae fused (figs. 486, 496), heavily

sclerotized and visible externally as a trapezoidal plate; no rami present.

Spermatheca of typical Pentatomine construction. One or two

small sclerites found round the entrance of the spermatheca into the

genital chamber (fig. 492); medianly spermatheca dilated into an elongate

chamber down the centre of which runs a heavily sclerotized rod, a narrow

duct passing along centre of rod and out of dilation to a well developed

pumping region with proximal and distal flanges, spermathecal bulb

attached directly to pump, varying in shape from species to species (see

54

North American Pentatomoidea

figures) .

Euthyrhynchus floridanus is unique in pos sessing a pair of ring sclerites

one on either side of the spermathecal opening, also the distal flange is

absent in the pumping region, otherwise similar to previous species.

Pentatomidae - Podopinae

Podopini

Amaurochrous dubius (Palisot de Beauvois), 1805

Amaurochrous cinctipes (Say), 1828

Genitalia typically Pentatomine in construction. Eighth para-

tergites ventrally not fused, without spiracles (fig. 498). Spermathecal

bulb with three processes (fig. 499) spermatheca otherwise similar to

that described under Pentatomini.

Weda parvula (Van Duzee), 1904

Genitalia and spermatheca very similar to preceding species,

spermathecal bulb with two processes only (fig. 500).

Tessaratomidae - Oncamerinae

Piezosternum subulatum (Thunberg), 1783

Eighth and ninth paratergites long apically acute sclerites (fig.

501), eighth with spiracles. Sclerotized and pair ed rami present, second

gonocoxae fused plate-like; second gonapophyses partially sclerotized.

Ring sclerites present (fig. 502), also noted by Scudder (1959) in

Piezosternum calidum ,

Spermathecal duct wide on entrance into genital chamber, slightly

sclerotized at base, long, coiled, terminating in a pumping region with

proximal and distal flanges; spermathecal bulb heavily sclerotized oval

in shape attached directly to pump. The spermatheca of this species

differs from Musgravea (Rhoecocoris) sulciventris figured by Pender grast(1956),

also a member of the Ocomerini, in not possessing a spermathecal

diverticulum but does resemble the other three species figured.

Acanthosomidae

Elasmostethus cruciatus (Say), 1831

External genitalia similar to Acanthosoma haemorrhoidale described by

Scudder (1959), tenth sternum divided (fig. 503); paired and sclerotized

rami present.

A small sclerotized groove found in floor of genital chamber

extending between entrance of spermatheca and that of oviduct. Sper-

matheca (fig. 504) consisting of a narrow duct terminating in a pumping

region with proximal and distal flanges; spermathecal bulb cylindrical.

Meadorus lateralis (Say), 1831

Genitalia and spermatheca (fig. 505) very similar

to Elasmostethus cruciatus ; eighth paratergites divided; distal and proximal

McDonald

55

flanges of pump well developed.

Cydnidae - Corimelaeninae

Corimelaenini

Corimelaena pulicaria (Germar)j 1839

Paratergites eight fused centrally (fig. 506); two pair s of elongate

sclerites visible above the large flap-like fir st gonocoxae, tenth sternite

being dorsalmost, ninth paratergites lying beneath. Second gonocoxae

not visible externally. No sclerotized rami or ring sclerites present.

Spermatheca (fig. 507) consisting of a simple duct connecting to

pumping region with proximal flange only developed; spermathecal bulb

mushroom shaped, attached directly to pump.

Galgupha nitiduloides (Wolff), 1802

External genitalia very similar to Corimelaena pulicaria . No sclero-

tized rami.

Spermatheca very similar to that figured and described by

Pender grast (1957) for Galgupha ovalis Huss, shape of spermathecal bulb

differs somewhat (fig. 509). Two accessory sacs present one on either

side of spermathecal entrance (fig. 508), their function unknown; sper-

mathecal opening into a narrow sclerotized groove.

Cydnidae -Cydninae

Cydnini

Dallasiellus discrepans (Uhler), 1877

Ovipositor facing caudad. Eighth paratergites (fig. 510) notfused

medianly. An elongate narrow sclerite found dor sally lying between

eighth paratergites, probably representing remains of the bridge between

them. Tenth sternum divided; ninth paratergites small oblong lying on

either side of fused second gonocoxae, latter clearly visible as a crescent

shaped sclerite almost divided into two by a deep ventral cleft; bases of

second gonapophyses visible. First gonocoxae large plate-like. Sclero-

tized rami present; ring sclerites present.

Spermatheca quite unlike any figured by Pendergrast (1957) for

Cydnidae. Spermathecal duct broad becoming somewhat dilated distally

and bearing internally a short stout sclerotized rod, a narrow duct passing

down the centre of this rod and into a broader coiled duct, passing to

pumping region (fig. 511) from dilation, pump with well developed

proximal and distal flanges; spermathecal bulb pear shaped.

Cyrtomenus crassus (Walker), 1867

External genitalia (fig. 512) very similar to Dallasiellus discrepans ;

eighth paratergites joined medianly by a very narrow bridge; bases of

second gonocoxae not visible externally. Sclerotized rami present; no

ring sclerites.

Spermatheca differing somewhat from that of Dallasiellus discrepans

although built along same lines. Spermathecal duct (fig. 513) basally

56

North American Pentatomoidea

wide, narrow medianly, and distally expanding into a globular chamber

within which is a second globular chamber, longitudinally ridged; a stout

sclerotized duct originating from inner chamber and linking external

chamber to pump; latter with well developed distal and proximal flanges;

spermathecal bulb oval, attached directly to pump.

Pangaeus aethiops (Fabricius), 1787

External, internal genitalia, and spermatheca (fig. 514) similar

to Dallasiellus discrepans. Internal rod of spermathecal diverticulum much

less heavily sclerotized; spermathecal bulb globular.

Amnestini

Amnestus pallidus (Zimmer), 1910

External genitalia most unusual, described and figured by

Froeschner (I960). Eighth paratergites (fig. 515) small V-shaped

structures lying one on either side laterally. Greater part of external

genitalia consisting of a large triangular sclerite surrounding an oval anal

aperture, probably representing fused ninth paratergites and tenth

sternum. First gonocoxae laterally placed, moveable, partly hidden by

the margin of the seventh sternum.

Base of spermathecal duct narrow opening into a small mound or

evagination of genital chamber; a small accessory spermathecal diver-

ticulum (fig. 516) opening into base of spermathecal duct. Medianly

spermathecal duct widening and thrown into two or three tight coils ter-

minating in an oval spermathecal bulb. Pumping region not clearly

evident although a small flange is present at base of spermathecal bulb.

Amnestus pusio (Stal), I860

External, internal genitalia, and spermatheca (fig. 517) similar

to Amnestus pallidus. Eighth paratergites not clearly deliminated, sperma-

thecal bulb spherical.

Sehirini

Sehirus cinctus (Palisot de Beauvois), 1805

Wagner (1963) gives a general description of the Sehirus type of

genitalia using Tritomegas sexmaculatus Rambur as an example. Scudder

(1959) gives a more complete general description for Sehirinae.

In Sehirus cinctus , eighth paratergites (fig. 518) continuous above

the anus, external genitalia otherwise similar to Tritomegas sexmaculatus .

Internally no sclerotized rami or ring sclerites present, differing in this

respect from Scudder rs general description for this group.

Spermatheca very similar to that of Sehirus bicolor (Linnaeus)

figured by Pender grast (19-57). Basally spermatheca wide (fig. 519) and

with numerous annulations apically narrowing and attached to a large

spermathecal bulb; pumping region part of basal portion of spermathecal

bulb, clearly marked by proximal and distal flanges.

McDonald

57

DISCUSSION

The female genitalia present a remarkably uniform picture

throughout this superfamily. Scudder (1959) made a detailed study of the

female genitalia of Heteroptera and Pender grast (1957) of the sperma-

thecae. Dupuis (1955) deals with the morphology of the genitalia in very

general terms. Several other workers have dealt with various genera

and families within the Pentatomoidea, their work has been incorporated

where relevant.

Pentatomidae-Scutellerinae

Odontoscelini

The external genitalia are very uniform in this group. The

spermathecal bulbs of the four species placed in this tribe tend to be

elongate. One species, Phimodera binotata , possesses a spermathecal

diverticulum and lacks the sclerotized canal running from the sperma-

thecal entrance in the base of the genital chamber, foundin the remaining

three species. The female genitalia and spermathecae are very similar

to those of the Pachycorini.

Eurygas trini

Eurygaster alternatus is well placed in a tribe of its own since it

possesses sclerotized and interlocking rami, a character found only

among members of the tribe Scutellerini so far. However, the sperma-

thecais much more similar to species in the Odontotar sini and Pachycorini

because it lacks the sclerotized spermathecal dilation and heavily sclero-

tized spermathecal bulb of the Scutellerini.

Pachycorini

All species examined in this tribe possess a sclerotized groove

or sclerite running along the genital chamber from the spermathecal

entrance. The spermatheca itself varies somewhat. All but two species

have either a spermathecal diverticulum or a dilation. The spermathecal

diverticulum is membraneous and sac-like and is either attached mid-way

to the spermathecal duct by means of a branch duct ( Stethaulax rnarmoratus ,

fig. 441) or is entirely separate ( Acantholomidea pprosa , fig. 434). The

spermathecal dilation is generally membraneous but is tough and sclero-

tized in Diolcus irroratiis (fig. 424). Chelysomide a guttata is very unusual in

possessing only one pair, (the outer) of sclerotized rami suggesting a

relationship with the Scutellerini. However, its spermatheca is much

more typical of the Pachycorini in possessing a weakly defined pumping

region, and elongate sausage-like spermathecal bulb (fig. 438).

Scutellerini

Augocoris gomesii was the only North American species studied.

Pendergrast (1957) studied five species of scutellerines. The sperma-

theca is characterized by the development in this group only of a very

tough sclerotized globose spermathecal diverticulum and a well defined

pumping region with proximal and distal flanges.

58

North American Pentatomoidea

Pentatomidae - Pentatominae

P entatomini

As pointed out in the introductory remarks, the genitalia and

spermathecae of this subfamily are remarkably homogeneous. The

presence or absence of spiracles on the eighth paratergites appear s to be

a random character of specific value only. However, in nine species a

very distinct character was noted, the spermathecal bulb had a series

of hollow horn-like projections varying in number from two to four. The

significance of these, structures is unknown. This character does not

occur in any of the species possessing an elongate endophallic duct and

hence does not reinforce in any way the division into two groups found

among the males of the North American genera*

Other characters included ring sclerites, found otherwise in only

two European species Pentatoma rufipes and Eysarcoris aeneus . It would be

interesting to know if this was common to all palaearctic genera, how-

ever, little work has been done at this level on the palaearctic fauna.

Scudder (1959) notes that ring sclerites may be present, also a tendency

for the development of additional sclerotizations around the opening of

the spermathecal duct. Most species examined in the present study

possessed one or two small sclerites around the spermathecal opening.

Thyanta perditor was slightly unusual in possessing an elongate

spermathecal bulb (fig. 479) with a peculiar pumping region but was

otherwise normal. The most aberrant species examined

was Trichopepla semivittata in which the spermatheca was a simple sac

resembling that of the Cryptostemmatidae. However further work will

have to be done on this genus to elucidate its homologies.

Halyini, Edessini, Discocephalini, Sciocorini and Mecidiini

Specimens examined from all these tribes all proved to have

genitalia similar to those of the Pentatomini. Brochymena (Halyini) and

Edessa (Edessini) have processes on the spermathecal bulb.

Pentatomidae - Asopinae

The female genitalia and spermathecae of species in this sub-

family are very similar to those of the Pentatominae, a fact noted by

Pender grast (1957). The female genitalia and spermatheca show a

remarkable uniformity throughout the group, paralleling that found in the

male genitalia. Based on the female genitalia and spermatheca the

Asopinae are very closely related to the Pentatominae.

Pentatomidae - Podopinae

Genitalia and spermatheca are similar to those of the Pentatominae

and the spermathecal bulb has two to three processes. This subfamily,

as in the case of the Asopinae, is very closely related to the Pentatominae

on the basis of the spermatheca and female genitalia.

Acanthosomidae

The genitalia of the two species examined are like those of the

Pentatominae externally; internally, sclerotized rami are present.

McDonald

59

agreeing with Scudder's (1959) general description. The spermatheca

has no diverticulum, differing in this respect from the general pentatomid

type. Acanthosoma haemorrhoidale (Linnaeus) figured by Pendergrast (1957)

also lacks a spermathecal diverticulum.

Unfortunately very few acanthosomids have been studied so far.

A total of six species (in four genera) of the world's fauna have been

described including the two species in this paper. On the basis of the

female genitalia the Acanthosomidae appear to be distinct from the

Pentatomidae in possessing sclerotized rami and in the form of the

spermatheca. More work needs to be done on this family before a de-

finitive statement can be made about its relationships. The status of this

family is considered below.

Tessaratomidae

Pendergrast (1957) figures the spermatheca of four species of

tessaratomids, Kumar (1962) describes and figures the genitalia of four

species and the spermatheca of one; Scudder (1959) examined three

species. Sclerotized and i nt e r 1 o c ki n g rami were found in

Piezosternum subulatum , consistent with Scudder *s description. Kumar (1962)

noted that in two species of Oncomerini the rami were absent, but were

present in Stilidia sp. The spermatheca of Lyromorpha rosea (Westwood) is

very similar (Kumar 1962) to that found in the Pachycorini (Scutellerinae)

as is the spermatheca of Musgravea sulciventris (Pendergrast 1957). The

spermatheca of Piezo sternum subulatum does not r esemble that of Musgravea

very greatly, consisting of a wide long spermathecal duct terminating in

a pumping region and bulb (fig. 502). This tends to reinforce the obser-

vation made in the discussion of the male genitalia that the subfamily

Oncomerinae is taxonomically heterogeneous. Other species described

all show great similarity to one another (Pendergrast 1957). They have

in common an ovoid spherical spermathecal dilation which is lacking in

Piezosternum m

Cydnidae

The major works on Cydnidae (Froeschner I960, Wagner 1963),

deal only with the external female genitalia and these give very little clue

to the relationships within this complex group. Scudder (1959) has

examined the genitalia of nine species; Pendergrast (1957) has figured

the spermatheca of four species; and Kumar (1962) the female genitalia

of four species and the spermatheca of two. The present study deals

with seven species, intended only to give a general idea of the relation-

ships of the Cydnidae. However, such a diver sity of form was discovered

especially in the type of spermatheca, that much more work will have to

be done to elucidate the systematic s of this family. Some tentative ideas

are presented, based on this and other work mentioned above.

Cydnidae -Corimelaeninae

Two species, Corimelaena pulicaria and Galgupha nitiduloides , were

examined. The genitalia were similar in both species agreeing with the

general description given by Scudder (1959). The spermathecae differ,

however, quite radically. Corimelaena pulicaria has a simple spermathecal

60

North American Pentatomoidea

duct with no diverticulum or dilation, terminating in a pump and sper-

mathecal bulb, resembling very closely the spermatheca of acanthosomids

and plataspids . Gatgupha nitiduloides has a sclerotized groove extending from

the entrance of the spermathecal duct in the genital chamber and a large

sac-like spermathecal diverticulum resembling very closely the type of

spermatheca found in the Pachycorini (Scutellerinae). An identical type

of spermatheca was found in Galgupha ovalis by Pendergrast (1957). How-

ever Thyrecoris scarabaeoides (Pendergrast 1957) has a third type of sper-

matheca which is similar to the one found in several species of Cydnini

(see below). This suggests that the subfamily Corimelaeninae is a

composite grouping. Pendergrast (1957) states that further species

should be examined to show whether Gulgupha ovalis is aberrant in its type

of spermatheca or whether there is diversity of form in this subfamily.

There is indeed diversity of form and further work needs to be done in

this group.

Oydnsdae-Oydnsnai

Cydnini

Three species were examined and all showedmarked similarities.

The female genitalia are somewhat more complicated in Dallasiellus discrepans9

two series of sclerites being found above the anus, the dorsal-most

probably representing the median section of the eighth parater gites.

Sclerotized rami are present in this group, these are not found in the

Cor emelaeninae. Ring sclerites were found in Dallasiellus discrepans and

Pangaeus aethiops , although Scudder (1959) stated that these are absent.

The spermatheca is very similar in all forms posses sing a small

spermathecal dilation within which is a stout sclerotized rod, globular

in Cyrtomenus crassus . The same type of spermatheca was found in the

species studied by Pender grast (1957) and in Stibaropus callidus (Kumar 1962),

The latter author however found a completely different type of sperma-

theca in Geotomus apicalis , In this species the spermathecal duct is very

long and highly coiled and the dilation is lacking. Pendergrast (1957)

notes the similarity of the type of spermatheca with a dilation and internal

rod to that found in the Pentatomidae. The cydnid dilation is, however,

much modified, the whole structure being smaller and stouter than the

structure found in the Pentatominae. Wagner (1963) has created a new

tribe Geotominiand this division may be further confirmed on the basis

of the spermatheca.

Amnestini

Two species were examined and these showed a highly aberrant

type of genitalia and spermathecae. Froeschner (I960) describes the

peculiar triangular plate which surrounds the anal opening. Its exact

homology is difficult to determine but probably represents the fused

ninth paratergites and tenth sternum. The spermatheca is unique, con-

sisting of a wide highly coiled spermathecal duct terminating in a sper-

mathecal bulb, no pumping region was apparent. Adjacent to the sperma-

thecal opening into the vulva is a small sac-like diverticulum.

It would appear that on the basis of the female genitalia and

McDonald

61

spermatheca, the Amnestini probably deserve at least subfamily status .

Sehirini

The genitalia of Sehirus’ cinctus do not res emble those of Sehirus bicolor

figured by Scudder (1959). Contrary to his general description of this

group, sclerotized rami and ring sclerites were not found in Sehirus cinctus.

The spermatheca of S. bicolor , (Pender grast 1957) is not the same as that

of S. cinctus , the latter species possesses a basal dilation connected by a

short duct to the pump and bulb (fig. 519). The dilation does not appear

to have the sclerotized rod found in the Cydnini. 'The position

of Sehirus cinctus is doubtful and is discussed below.

INTERRELATIONSHIPS AND CLASSIFICATION

OF THE PENTATOMOIDEA

A great deal of work has been done on the systematic s of the

Heteroptera of which the Pentatomoidea are a part. However, I feel

that too much emphasis has been laid on results obtained from a very

small number of species examined in the various families. It became

clear after detailed examination of the North American Pentatomoidea

that great variation of structure occurs in all families and that workers

choosing but a few random species would get and have got quite an

erroneous impression of the group as a whole. This is particularly so

among the male genitalia of the Scutellerinae where the only work pre-

viously done was by Leston (1952) on the tribe Pachycorini. The results

proved to be quite startlingly different from those obtained by other

workers examining species only from the Scutellerini.

Pruthi (1925), Pendergrast (1957), Scudder (1959), Leston (1958),

Manna (1958), and Miyamoto (1961), have dealt with the clas sification of

the Pentatomoidea from various points of view. Leston ei al. (1954)

proposed a clas sification of terrestrial Heteroptera based on a synthesis

of all previous morphological work on this group. Its weakness, as

China (1955) points out, is in the fact that most of the previous work on

the Heteroptera had been rather fragmentary and dealt with rather small

samples in the various groups each worker had had under consideration.

The present work will help fill in some gaps in our knowledge, but it

cannot be regarded as being complete in any way and any conclusions

reached must be regarded with some reserve.

I shall consider first the phylogeny and relationships of the

Pentatomoidea. This superfamily together with the Pyrrhocoroidea,

Lygaeoidea, Coreoidea, Piesmatoidea and Aradoidea forms the group

Pentatomorpha (Leston et al. 1954). The following features are charac-

teristic for the Pentatomoidea. The males have the ninth segment

developed in the pygophore in which are found a pair of claspers and the

aedoeagus. The latter consists of a toughened theca, the conjunctiva

which generally bears one to three pairs of conjunctival appendages and

the vesica. The seminal duct generally enters a sclerotized ejaculatory

reservoir, variously modified, and this in turn connects with the endo-

phallic duct, which opens at the secondary gonopor e. The female genitalia

62

North American Pentatomoidea

are of the plate-like type (Scudder 1959). Ring sclerites and rami may

be present. The spermatheca is characterized by awellmarked pumping

region, generally with proximal and distal flanges, terminating in a

spermathecal bulb of varied shape.

The Coreoidea and Pyrrhocoridea are, on the basis of the female

genitalia, the closest to the Pentatomoidea in possessing a plate-like

ovipositor (Schaefer 1964). The Lygaeoidea, on the other hand, have a

laciniate type of ovipositor, with some exceptions. The male genitalia

of the coreoid complex are probably the closest to the pentatomoid type

in possessing a distinct ejaculatory reservoir and membraneous conjun-

ctival appendages (Scudder 1957), but the vesica is different in that the

endophallic duct in the coreoids is generally very elongate (Pruthi 1925,

Scudder 1957). Piezosternum (Tes seratomidae) has, however, a very highly

endophallic duct within the apex of the vesica and if this were extrusible

it would produce a very long apical duct. The latter would resemble the

very long coiled endophallic ducts found in the Lygaeidae (Ashlock 1957) ,

The remainder of the families in the Pentatomoidea have a relatively

short vesica with exception of a few genera in the Pentatominae. The

latter group is probably a highly specialized development of the normal

pentatomine type.

Miyamoto (1961) found that the coreoids showed resemblances to

the pentatomoids on the basis of the gastric caeca but that the structure

of the salivary gland resembled that of the pyrrhocorids.

The cytogenetics of the pentatomoid group is complex and evidence

for relationship with other groups is not clear cut. Leston (1958) found

two distinct groups within the Pentatomoidea: the Acanthosomidae,

Tessaratomidae and Scutelleridae with 2n = 12 chromosomes, and the

Pentatomidae with 2n = 14. However, in the latter family chromosome

numbers range from 2n = 6 to 2n = 27. Neither the Coreoidea nor the

Lygaeoidea show close relationship to the Pentatomoidea cytologically.

The coreids have varying chromosome number s with a mode of 2n = 21.

The Lygaeidae on the other hand have also variable chromosome number s

but have a diploid number of 2n = 14. The coreids have an XO sex mec-

hanism, the lygaeids an XY sex mechanism resembling the Pentatomidae.

However, the latter family does not possess m-chr omosomes found in a

great majority of the species of lygaeids.

Manna (1958) derives the Lygaeidae from the Pentatomidae on the

basis of cytological evidence. This I doubt on the basis of evidence from

the male and female genitalia, the lygaeids generally possessing a

primitive laciniate type ovipositor.

Schaefer (1964), on the basis of a vast amount of data, derived

both the Coreoidea and Pyrrhocoroidea from the Lygaeoidea. The

Pentatomoidea show some relationship to the above superfamilies but

this is not very close. The Pentatomoidea probably were derived indepen-

dently of the lygaeoids from some common ancestor, as suggested by

China's (1955) diagram of the relationships of the heteropterous families.

Leston (1958) and China and Miller (1959), on the other hand proposed

independent origins for the Lygaeoidea, Coreoidea and Pentatomoidea

from a common ancestor. The evidence obtained so far indicates that

the latter theory is closer to reality.

McDonald

63

The relationship of the four groups is shown below.

COREOIDEA PYRRHOCOROIDEA

LYGAEOIDEA PENTATOMOIDEA

Relationships within the Pentatomoidea

Ranking of the Scutellerinae

The status of the Scutellerinae has posed quite a problem in the

past. Now that all representative tribes have been examined, I am

inclined to agree with Pendergrast (1957) and Kumar (1962), and raise

the Scutellerinae to family rank. The group is, however, difficult to

define on the basis of the male and female genitalia. The Scutellerini

forma distinct group possessing in the males three pairs of conjunctival

appendages, with the third generally heavily sclerotized and S- shaped.

The vesica has a long convoluted duct and the endophallic duct is short.

The females have paired sclerotized rami and the spermatheca has a

heavily sclerotized dilation and a distinct pumping region. The Eury-

gastrini show characters intermediate between the Scutellerini and

Pachycorini. They have sclerotized and interlocking rami in the females.

The spermatheca is, however, much simpler, lacks a dilation and flanges

in the pumping region but there is a sclerotized groove in the genital

chamber running from the spermathecal opening, a characteristic of the

Pachycorini.

The genus Eurygaster has been raised to tribal status by Lattin

(1964) and this is supported by my own work. However, Wagner (1963)

raised this group to family level, and this I think is hardly warranted on

the basis of the morphology of the genitalia. The tribe shows very great

similarities to the other tribes within the Scutellerinae, especially the

Pachycorini. The latter tribe and the Odontotar sini* are very similar.

Both groups possess elongate sclerotized grooves in the floor of the

genital chamber, a feature not found in the Pentatominae. Rami are

lacking except in one species Chelysomidea guttata (Pachycorini) in which only

the outer rami are present. The spermatheca has either a simple duct

or a membraneous diverticulum attached half way along the spermathecal

duct, or separatelyat the base of the duct (sclerotized in DioLcus irroratus ).

All these types of spermatheca do not resemble in any way the elongate

dilation with central rod found in the Pentatominae. More work will have

to be done on the Palaearctic species of the Odontoscelini before an

adequate definition of this tribe can be made.

Relationships within the Pentatomoidea

The Pentatominae and Podopinae are remarkably constant in

genitalic characters. The male and female genitalia of the Asopinae show

remarkable similarity to one another and to the Pentatominae. I think

on this basis the subfamily should be downgraded and given tribal status

64

North American Pentatomoidea

within the Pentatominae. The characters possessed in common by all

genera in the Asopinae such as the genital plates and thecal shield are

also found in species of the Pentatominae but never in combin-

ation. The internal structure of the vesica is typically pentatomine and

the structure of the spermatheca is identical to that found in that sub-

family. The similarity of the Asopinae to the Pentatominae was noted by

Leston (1954a).

The Podopinae and Asopinae are very closely related. The

Podopinae lack genital plates but have in their place a pair of pygophoral

appendages. The structure of the aedoeagus and vesica is identical in

both subfamilies. Leston (1953a) raised the podopines to subfamily

status but felt that further research might lead to a drop in its rank. On

the basis of the work done by Barber and Sailer (1953) and the present

study this group should be given tribal status within the subfamily

Pentatominae. Pendergrast (1957) states that the Podopinae and Asopinae

are so close to the Pentatominae that they should either be lowered in

status or that the other subfamilies should be raised in status. I think

the former course more desirable because of the very close affinities

this group shows to the Pentatominae.

Tribal status in the P entatominae

Within the Pentatominae the tribes Halyini and Mecidiini on the

basis of the male and female genitalia are so similar to the Pentatomini

that these two tribes should be given subtribal status or incorporated

into the Pentatomini as genera. However, other genera within the Halyini

may warrant tribal status. The vesicae of five species of Australian

Halyini have been described by Kumar (1964) and these all resemble the

typical plan found among Pentatomini. Ruckes (1946, 1958) who has made

a major study of this group, has not described the internal details of the

male genitalia. The genitalia of the Mecidiini studied by Sailer (1952)

were all remarkably uniform in character and are similar to the Penta-

tominae.

The Discocephalini have recently been raised to subfamily status

by Ruckes (I960, 1963). However, I hesitate to follow such a step until

further work has been done on the male and female genitalia of the species

in this tribe. Ruckes (personal communication) informs me that he has

several excellent characters which distinguish this tribe from others in

the Pentatominae. The male genitalia of Lineostethus clypeatus differ but

slightly from the pentatomid type in having a thickened sclerotized ring

at the base of the endophallic duct. The female genitalia are typically

pentatomine in construction.

A single species of both the Edessini and Sciocorini was studied.

In both the female genitalia were typically pentatomine. The male

genitalia, however, showed slight differences from the general pattern

especially in Edessa bifida. More work will have to be done on these groups

before any definitive statement can be made. It would seem, however,

that both these tribes are fairly closely related to the Pentatomini.

McDonald

65

Status of the Acanthosomidae

The Acanthosomidae have been accorded family status by Leston

(1953b). China (1959) retains the group as a subfamily of the Penta-

tomidae. On the basis of the male genitalia I agree with China. The

acanthosomids are undoubtedly older and less specialized than the pen-

tatomines in many respects. The spermatheca lacks the highly

specialized dilation it has in the pentatomines. Leston (1958) found the

chromosome number tobe2n = 12 with an XY sex determining mechanism,

characters common to the Scutellerinae. Both Dupuis (1948) and

Southwood (1956) consider the Acanthosomidae to be a primitive family.

The Cydnidae

A preliminary study of a few species in this family has revealed

a considerable diversity in the structure of the genitalia and the family

is one of great interest. Froeschner (I960) was dubious regarding the

phylogenetic relationships indicated by the presence of a fringe of close

set bristles on the apices of the middle and posterior coxae, external

morphological characters used to distinguish members of this family

from other families in the Pentatomoidea. He goes on to question the

value of characters used to define groups within the Pentatomoidea as

indicators of phylogeny within this superfamily as a whole. From a study

of the male and female genitalia it becomes clehr that the Cydnidae is a

somewhat heterogeneous assemblage. This view was also held by

Pendergrast (1957).

Very little can be said regarding the Corimelaeninae at this stage.

The male genitalia of Corimelaena pulicaria do not resemble the general pattern

found in the Pentatominae, as three pairs of conjunctival appendages

were found, and the theca possessed a peculiar pair of thecal appendages 5

the vesica is simple. The male genitalia of species figured by McAtee

and Malloch (1933) appear toresemble Corimelaena pulicaria in many details.

The two spermathecae examined were quite different; one was similar

to the acanthosomid type and the other to the pachycorine (Scutellerinae).

The genitalia show very little similarity to those of the Cydninae studied

so far. It may be that on further investigation the present recognition

of Corimelaeninae as a family by specialists in North America will

achieve wider acceptance.

The Cydninae, on the basis of male and female genitalia so far

examined show relationship with the Pentatominae. The eggs (Southwood,

1956) show similarity to those of the Pyrrhocoridae and some Lygaeidae.

Southwood (1956) stated that the family as a whole was rather ancient and

closer to the Tessaratomidae than to the Pentatomidae. The ovariole

number (Miyamoto 1957, Woodward 1950) for the majority of species

examined was seven which is also the most frequent number found among

Pentatominae. Little work has been done on the chromosomes of this

subfamily (Leston 1958, Manna 1958).

The relationships of the Cydnidae will not be discussed further.

I feel at the moment that too little is known about the basic morphology

of the species in this family and further research needs to be done.

66

North American Pentatomoidea

Phylogenetic considerations

The ancestral pentatomoid probably had very simple male

genitalia. The vesica was long, the seminal duct probably entered into

a simple sac-like ejaculatory reservoir. Conjunctival appendages, if

present, were small and membraneous. The female spermatheca was

a simple duct terminating in a pumping region and spermathecal bulb.

The ovipositor was of the plate- shaped type. The Tessaratomidae are

probably the most primitive family (Leston 1954d) with Piezosternum being

the most primitive genus so far examined in the family. Piezosternum both

in the structure of the aedoeagus and spermatheca shows definite cor eoid

affinities and is radically different from other members of this family,

e.g. Musgravea sulciventris . Piezosternum is probably very close to the ancestral

pentatomoid which gave rise to the variously modified groups in this

superfamily.

The ancestral pentatomoid stock appears to have evolved into

two distinct lines, the Scutelleridae and the Pentatomidae. In theScutel-

leridae the male genitalia became slightly more complex with the seminal

duct opening into an internal canal within the ejaculatory reservoir.

Conjunctival appendages became more complex but were still generally

membraneous. The spermatheca was still simple but in some cases

possessed a diverticulum or dilation. These characters are typical of

Pachycorinae and Odontoscelinae. The Scutellerinae are the most highly

evolved and specialized group. The third conjunctival appendages have

become sclerotized and S- shaped. The vesicahas a specialized convoluted

duct and the endophallic duct has become very much shortened. The

females possess interlocking and sclerotized rami, a distinctly non-

pentatomine character, and the spermathecal duct has a sclerotized

dilation.

Within the Pentatomidae, the Acanthosominae must be considered

a very early offshoot of the pentatomid stock but still closely related to

the Pentatominae. The females retain the simple type of spermatheca

but have developed sclerotized rami in the ovipositor, parallelling the

Scutellerinae. The male genitalia, in the structure of the vesica, re-

semble the Pentatominae closely in that the seminal duct either opens

into an internal canal within the ejaculatory reservoir, or directly into

the reservoir. The conjunctival appendages tend to be more specialized

and are sclerotized. As pointed out previously the Acanthosominae retain

the more primitive chromosome number 2n = 12 (Leston 1958) and very

likely represent an early group in the development of the more highly

specialized Pentatominae.

The Pentatominae have generally retained a simple vesica with

the seminal duct opening into the ejaculatory reservoir via an internal

canal. The North American fauna have evolved a small group of very

specialized species in which the endophallic duct has become enormously

lengthened and coiled; the ejaculatory reservoir has a complex series

of internal ducts. The Pentatominae have developed two specialized

features, the sclerotized medianpenal lobes and the dilation with internal

rod of the spermatheca. Sclerotized rami have not been developed in

this group. The type of spermatheca is quite constant throughout this

subfamily with the sole exception of Trichopepla semivittata in which the

McDonald

67

spermatheca is sac-like. The median penal lobes are not found in all

species. TheAsopini, Podopini, Edessini, Discocephalini and Sciocorini

are all very recent specializations of the main pentatominae stock and

r etain many character s in common with the latter group. The phylogenetic

sequence of the Pentatomoidea excluding the Cydnidae follows:

Ancestor

68

North American Pentatomoidea

The paired sclerotized rami found in the female genitalia have

apparently evolved independently three times, in the Tessaratomidae

( Piezosternum) 9 the Scutellerinae, and the Acanthosominae. Spermathecal

dilations have evolved twice. The Pentatominae have developed the

specialized membraneous dilation with internal rod and the Scutellerinae

a heavily sclerotized and less specialized dilation. In the male genitalia

specialized structures have evolved in great profusion in each group.

Median penal lobes have apparently evolved in two groups, the Penta-

tominae and in the Scutelleridae where they are found in a single species

Symphylus carribeanus . The conjunctival appendages are subject to great

change and have become variously modified in each subfamily.

Proposed C Eassif leaf ion of Families, Subfamilies and Tribes of North American Pentatomoidea

SCUTELLERIDAE

Odontoscelinae

Eurygastrinae

Pachycor inae

Scutellerinae

PENTATOMIDAE

Pentatominae

Pentatomini

Edes sini

Discocephalini

Sciocorini

Asopini

Podopini

Acanthosominae

CYDNIDAE

Corimelaeninae

Cydninae

Cydnini

Amnestini

Sehirini

TESSARATOMIDAE

Piezosterninae

McDonald

69

KEY TO GENERA OF NORTH AMERICAN SCUTELLERIDAE

BASED ON MALE GENITALIA

4.

5.

1. Vesica with long convoluted duct* extending from anterior end into

ejaculatory reservoir (fig. 30) 2

Vesica without such duct 4

2. Endophallic duct with a broad oblong dorsal sheath (fig. 30£; con-

junctival appendages membraneous Camirus Stal 1862

Endophallic duct without sheath; at least one pair of conjunctival

appendages heavily sclerotized apically 3

3. Ejaculatory reservoir globose (fig. 61); apex of vesica flattened

Stethaaiax Bergroth 1891

Ejaculatory reservoir elongate (fig. 83); apex of vesica tubular

Augocoris Burmeister 1835

Three pairs of sclerotized horn- like conjunctival appendages; theca

with cylindrical ventral process (fig. 25) Eurygaster Laporte 1832

Never with all characters above . 5

Ejaculatory reservoir simple, sac-like composed of a single cham-

ber, or absent 6

Ejaculatory reservoir complex; if apparently simple, large spiny

third conjunctival appendages present (fig. 11) or apex of vesica

with spiny processes one on either side (fig. 17) 11

6. Ejaculatory reservoir absent; seminal duct opening directly into

endophallic duct (fig. 36) 7

Ejaculatory reservoir present as a small diverticulum (fig. 56) . . 8

7. Pygophore with dorsal margin produced into an elongate process

(fig. 32); endophallic duct very short, not projecting beyond margin

of theca Pachycoris Burmeister 1835

Pygophore with smooth dor sal. mar gin; vesica with complex pum-

ping apparatus (fig. 72); endophallic duct projecting well beyond

margin of theca Diolcus Mayr 1864

8. Apex of vesica covered with a number of stout spines and with a

stout spiny dorsal process (fig. 55) Sphyrocoris Mayr 1864

Vesica without spines 9

9. Three pairs of conjunctival appendages present, third spiny; apex

of vesica very broad, covered with spines, basally with a number

of partitions giving a coiled appearance (fig. 46)

. Homaemus Dallas 1851

Only two pairs of conjunctival appendages present 10

10. Dor sal margin of proctiger produced into a number of spiny processes

(fig. 37); apex of vesica membraneous . . . Chelysomidea Lattin 1965

Dorsal margin of pygophore smoothly arched; apex of vesica sclero-

tized Tetyra Fabricius 1803

11. Third conjunctival appendages present, broad and spiny (fig. 11) .

12

Third conjunctival appendages absent 13

Second conjunctival appendages bifid, bearing two sclerotized horns

(fig. 6) Fokkeria Schouteden 1904

Second conjunctival appendages bearing a large single horn ....

Euptychodera Bergroth 1908

12.

70

North American Pentatomoidea

13. Vesica with a pair of spiny lobes, one on each side near apex (fig.

17) Vanduzeeina Schouteden 1904

Vesica without such lobes 14

14. Endophallic duct basally with a very short convoluted section (fig.

22); only one pair of membraneous conjunctival appendages present.

Phimodera Germar 1839

Endophallic duct without convolutions; two pairs of conjunctival

appendages present ... 15

15. Apex of vesica very short projecting slightly beyond mar gin of theca

(fig. 77); claspers broadly hook- shaped . . A cantholomidea Sailer 1945

Apex of vesica long enclosed between median penal lobes (fig. 65);

claspers T-shaped Symphylus Dallas 1851

KEY TO GENERA OF NORTH AMERICAN PENTATOMINI

BASED ON MALE GENITALIA

1.

2.

3.

4.

5.

6 .

7.

8.

Endophallic duct very long, coiled (fig. 236), dorsal margin of theca

with thecal processes (fig. 234), ejaculatory reservoir complex

(fig. 236) 2

Endophallic duct short, theca with or without dorsal processes;

ejaculatory reservoir generally simple with posterior canal (fig.

89) 6

Ejaculatory reservoir moderately sclerotized, not possessing a

number of lateral striae (fig. 250) Euschistus Dallas 1851

Ejaculatory reservoir very heavily sclerotized with a number of

well marked striae laterally 3

Two distinct pairs of membraneous conjunctival appendages, second

with five lobes (fig. 233) Menecles Stal 1867

One pair of conjunctival appendages generally only, shallowly divided

4

Thecal proces ses with a distinct projection between them from mar-

gin of theca, conjunctival appendages elongate distinctly bifid (fig.

239) Coenus Dallas 1851

Combination of characters not as above, conjunctival appendages if

bifid, broadly so 5

Ventral border of pygophore with a deep median U-shaped emar-

gination (fig. 241) conjunctival appendages broad, undivided (fig.

243) Hymenarcys Amyot and Serville 1843

Ventral border without emargination, conjunctival appendages broadly

bifid (fig. 253) Prionosoma Uhler 1863

Conjunctival appendages absent 7

Conjunctival appendages present 8

Theca shield-like, enclosing a further sheath-like structure (fig.

228), claspers very complex (fig. 226)

Loxa Amyot and Serville 1843

Theca not as above, claspers trilobed (fig. 146)

Chlorocoris Spinola 1837

Thecal shield present (fig. 183) 9

Thecal shield absent 11

McDonald

71

9.

10.

11.

12.

13.

14.

15.

16.

17.

18.

19.

20.

21.

22.

Seminal duct extended into dorsal canal, ejaculatory reservoir

simple • 10

Seminal duct extended into base of endophallic duct; ejaculatory

reservoir divided (fig. 185 Murgantia Stal 1862

Apex of vesica projecting well beyond the margins of the median

penal lobes, not enclosed by them (fig. 108)

Peribalus Mulsant and Rey 1866

Apex of vesica not or only slightly projecting beyond margins of

median penal lobes which otherwise enclose apex 11

Conjunctival lobe present, very large (fig. 179): ventral surface

of pygophore vertical (fig. 177) Neotiglossa Kirby 1837

Conjunctival lobe small (fig. 131): ventral surface of pygophore

horizontal Aelia Fabricius 1803

Genital plates present 13

GenitaL plates absent 14

Ejaculatory reservoir simple, with posterior canal; vesica S-shaped

(fig. 154) «... Carpocoris Kolenati 1846

Ejaculatory reservoir with internal duct, vesica short

Dendrocoris Bergroth 1891

Median penal lobes absent 15

Median penal lobes present (fig. 17 0) 18

Ejaculatory reservoir without posterior *canal (fig. 114) ......

. . Trichopepla Stal 1867

Ejaculatory reservoir with posterior canal (fig. 123) 16

Conjunctival appendages divided into three distinct broad lobes (fig.

122). Endophallic duct short, curved (fig. 123)

Brepholoxa Van Duzee 1904

Conjunctival appendages elongate; endophallic duct S- shaped (fig.

196) 17

Theca with a small pair of projections one on each side, near base

(fig. 199). Second conjunctival appendages present

Cosmopepla Stal 1867

Theca without projections; second conjunctival appendage absent

(fig. 194) Eysarcorls Hahn 1834

Ejaculatory reservoir divided into two ducts by means of an internal

septum 19

Ejaculatory reservoir simple, with posterior canal (fig. 128) . . 20

Claspers flattened, leaf-like; lateral borders of pygophoral opening

smooth Padaeus Stal 186 2

Claspers, stout, wide, apically produced into cylindrical processes

(fig. 175); lateral borders of pygophoral opening with a small oblong

process, one on each side (fig. 173) ...... Proxys Spinola 1837

Ventral margin of pygophore flattened with two longitudinal ridges

medianly (fig. 124) . . . \ Arvelius Spinola 1837

Ventral margin without ridges 21

Claspers apically trilobed (fig. 216) 22

Claspers not lobed 23

Conjunctival appendages voluminous bag-like structures (fig. 204).

Dorsal margin of pygophore with a few setae

Rhytidolomia Stal 1872

Chlorochroa Stal 1872

72

North American Pentatomoidea

Conjunctival appendages somewhat C-shaped. Dorsal margin of

pygophore with two distinct patches of stout setae, one on either

side of the mid-line (fig. 132) , Vulsirea Spinola 1837

23. Ejaculatory reservoir with no internal canal, simple sac-like (fig.

143) 24

Ejaculatory reservoir with posterior canal 26

24. Median penal lobes produced into a sheath around apex of vesica . . .

25

Median penal lobes produced into two tubular processes one on either

side of apex of vesica (fig. 142) Acrosternum Fieber 1861

25. Endophallic duct with a well developed broad curved dorsal flange

(fig. 118). Clasper small, knob-like

Mormidea Amyot and Serville 1843

Endophallic duct without flange, slightly curved at apex. Clasper

C-shaped with teeth on inner margin (fig. 102)

Solubea Bergroth 1891

26. Conjunctival appendages basally with four distinct lobes (fig. 98),

second conjunctival appendages with well sclerotized apices ....

. Piezodorus Fieber 1861

First conjunctival appendages without sclerotized apices; second

absent 27

27. Claspers oblong, flattened, leaf-like (fig. 221). Ventral margin of

pygophore with two double knobbed processes, one on each side

(fig. 220) J Banasa Stal I860

Claspers generally C-shaped or T-shaped when viewed laterally.

No processes on ventral margin of pygophore 28

28. First conjunctival appendages distinctly bilobed, apex of one lobe

sclerotized 29

Fir st conjunctival appendages bag-like structures, not divided . . .

30

29. Ejaculatory reservoir globose (fig. 165); median penal lobes curved

around apex of vesica Thyantq Stal 1862

Ejaculatory reservoir elongate; median penal lobes not curved

around apex of vesica Mecidea Dallas 1851

30. Apex of vesica short, not projecting beyond mar gins of median penal

lobes (fig. 159). Ejaculatory reservoir elongate, dorsal margin

entire Nezara Amyot and Serville 1843

Apex of vesica projecting well beyondmar gins of medianpenal lobes.

Ejaculatory reservoir oblong, dorsal margin with a deep groove

(fig. 260) Brochymena Amyot and Serville 1843

McDonald

73

ACKNOWLEDGEMENTS

I wish to thank the following persons for the loan of material:

Dr. Jon Herring and Dr. Richard Froeschner of the United States

National Museum; Dr. Herbert Ruckes, American Museum of Natural

History; Mr. Hugh B. Leech, California Academy of Sciences and Dr.

John D. Lattin, Department of Entomology, Oregon State University.

I am especially grateful to Dr. John D. Lattin for his help and

many comments on my work while working with him at Corvallis during

part of the summer of 1964.

I wish to express my gratitude to Dr. G. E. Ball for his help and

guidance throughout this study. I also wish to thank the following persons

for advice given: Dr. B. Hocking, Dr. W. G. Evans, Professor J. G.

Packer and Dr. Janet Sharplin.

REFERENCES

Ashlock, P.D. 1957. An investigation of the taxonomic value of the

phallus in the Lygaeidae (Hemiptera-Heteroptera) . Ann. ent. Soc.

Amer. 50 : 407-426.

Baker, A.D. 1931. A study of the male genitalia of Canadian species

of Pentatomidae. Canad. J. Res. 4 : 148-220.

Barber, H. G. and R.I. Sailer. 1953. A revision of the turtle bugs of

North America (Hemiptera, Pentatomidae). J. Wash. Acad. »Sci.

45 : 150-162.

China, W.E. 1955. A reconsideration of the systematic position of the

family Joppeicidae Reuter (Hemiptera-Heteroptera), with notes on

the phylogeny of the suborder. Ann. Mag. nat. Hist. ser. 12, 8 :

353-370.

China, W. E. and N. C.E. Miller. 1959. Checklist and keys to the

families and subfamilies of the Hemiptera-Heteroptera. Bull. Brit.

Mus. (Nat. Hist.), Ent. 8(1) : 1-45.

Crampton, G. C. 1922. The genitalia of the males of certain Hemiptera

(Heteroptera) and Homoptera. Bull. Brooklyn ent. Soc. 17 : 46-55.

Dupuis, C, 1948. Nouvelles donnes biologiques et morphologiques sur

les dipteres Phasinae, parasites d-Hemipteres Heteropter es . Ann,

Parasit. hum. comp. 22 : 201-232.

Dupuis, C. 1955. Les genitalia des Hemipter es -Heteropter es (genitalia

externes des deux sexes; voies ectodermiques femelles). Revue de

la morphologie, lexique de la nomenclature. Index bibliographique

analytique. Mem. Mus. Hist. nat. Paris (A) 6 : 183-278.

Froeschner, R.C. I960. Cydnidae of the western hemisphere. Proc.

U.S. Nat. Mus. Ill : 337-680.

James, M. T. 1953. Determining generic limits. Syst. Zool. 2 : 136-

137.

Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera). 1

Cimicidae. Felix L. Dames, Berlin. 392 pp.

Kumar, R . 1962. Morpho-taxonomic studies on the genitalia and salivary

glands of some Pentatomoidea. Ent. Tidskr. 83 : 44-84.

74

North American Pentatomoidea

Kumar, R. 1964. On the structure and function of the so-called ejacu-

latory reservoir in the Pentatomoidea (Hemiptera, Heter optera).

Proc. R. Soc. Qd. 75(8) : 51-65.

Kumar, R. 1965. Contributions to the morphology and relationships of

Pentatomoidea (Hemiptera : Heter optera). Part I. Family Scutel-

leridae. J. ent. Soc. Qd. 4 : 41-55.

Lattin, J. D. 1964. The Scutellerinae of America north of Mexico

(Hemiptera : Heteroptera : Pentatomidae) . Ph.D. Thesis, Univ. of

California, Berkeley.

Leston, D. 1952. Notes on the Ethiopian Pentatomoidea (Hemiptera) :

VIII, Scutellerinae Leach of Angola, with remarks upon the male

genitalia and classification of the sub-family. Publcoes cult. Co.

Diam. Angola 16 : 9-26.

Leston, D. 1953a. On the wing venation, male genitalia and spermatheca

of Podops inuncta (F. ), with a note on the diagnosis of the sub-family

Podopinae Dallas (Hemiptera, Pentatomidae). J. Soc. Brit. Ent.

4 : 129-135.

Leston, D. 1953b. Notes on the Ethiopian Pentatomoidea (Hemiptera):

XVI, an Acanthosomid from Angola with remarks upon the status and

morphology of the Acanthosomidae Stal. Publcoes cult. Co. Diam.

Angola 16 : 123-132.

Leston, D. 1953c. The suprageneric nomenclature of the British

Pentatomoidea (Hemiptera). Entomologist's Gaz. 4 : 13-25.

Leston, D. 1954a Notes on the Ethiopian Pentatomoidea (Hemiptera) :

XII, On some specimens from Southern Rhodesia, with an inves-

tigation of certain features in the morphology of Afrius figuratus (Ger-

mar) and remarks upon the male genitalia in Amyotinae. Occ. Pap.

Nat. Mus. S. Rhodesia 19 : 678-686.

Leston, 1954b. The male genitalia of Sehirus bicolor (L. ) (Heteroptera,

Cydnidae). J. Soc. Brit. Ent. 5 : 75-78.

Leston, D. 1954c. Notes on the Ethiopian Pentatomoidea (Hemiptera):

XVII, Tessaratominae, Dinidorinae and Phyllocephalinea of Angola.

Publcoes cult. Co. Diam. Angola 24 : 13-22.

Leston, D. 1954d. Wing venation and male genitalia of Tessaratoma

Berthold, with remarks on Tessaratominae Stal (Hemiptera,

Pentatomidae). Proc. R. ent. Soc. Lond. (A) 29 : 9-16.

Leston, D. 1955. A key to the genera of Oncomerini Stal (Heteroptera :

Pentatomoidae : Tessaratominae), with the description of a new

genus and species from Australia and new synonomy. Proc. R. ent.

Soc. Lond. (B) 24 : 62-68.

Leston, D. 1956. The Ethiopian Pentatomoidea Hemiptera : XXII, on

Dismegistus Amyot and S er ville (Cydnidae). Proc. R. ent. Soc. Lond.

(A) 31 : 87-94.

Leston, D. 1958. Chromosome number and the systematic s of Pentato-

morpha (Hemiptera). Int. Congr. Ent. 10. Montreal, 1956, 2 :

911-8.

Leston, D., J. G. Pender grast and T. R . Southwood. 1954. Classification

of the terrestrial Heteroptera (Geocorisae) . Nature, Lond. 174 :

91-92.

Leston, D. and G. G. E. Scudder. 1957. The taxonomy of the bronze

McDonald

75

orange-bug and related Australian Oncomerinae (Hemiptera : Tes-

saratomidae). Ann. Mag. nat. Hist ser. 12, 10 : 439-448.

McAtee, W. L. and J. R. Malloch, 1933. Revision of the subfamily

Thyr eocorinae of the Pentatomidae (Hemiptera-Heteroptera). Ann.

Carnegie Mus. 21 : 191-411.

McDonald, F. J.D. 1961. A comparative study of the male genitalia of

Queensland Scutellerinae Leach (Hemiptera : Pentatomidae). Pap.

Dep. Ent. Univ. Qd. 1 : 173-186.

McDonald, F. J. D. 1963. Morphology of the male genitalia of five

Queensland Scutellerinae (Hemiptera : Pentatomidae). J. ent. Soc.

Qd. 2 : 24-30.

Mann, G. K. 1958. Cytology and inter-relationships between various

groups of Heteroptera. Proc. Congr. Ent. 10. Montreal, 1956,

2 : 919-34.

Miyamoto, S. 1957. List of ovariole numbers in Japanese Heteroptera

(in Japanese) Sieboldia, 2 : 69-82.

Miyamoto, S. 1961. Comparative morphology of alimentary organs of

Heteroptera, with the phylogenetic consideration. Sieboldia, 2(4) :

197-258.

Pendergrast, J. G. 1957. Studies on the reproductive organs of the

Heteroptera with a consideration of their bearing on classification.

Trans. R. ent. Soc. Lond. 109(1) : 1-6.3.

Piotrowski, F. 1950. Sur la morphologie de Pappareil copulateur male

des Hemipteres-Heteropter es, avec consideration speciale du groupe

Pentatomoidaria Borner 1934 (In Polish, French summary). Soc.

Amis. Sci. Lettr. Poznan (B) 12 : 237-274.

Pruthi, H.S. 1925. The morphology of the male genitalia in Rhynchota.

Trans, ent. Soc. Lond. 1925 : 127-254.

Ruckes, H. 1946. Notes and keys on the genus Brochymena (Pentatomidae,

Heteroptera). Ent. Amer. 26(4) : 143-238.

Ruckes, H. 1958. New genera and species of neotropical discocephaline

and halyine pentatomids (Heteroptera, Pentatomidae). Amer. Mus.

Novit. 1868 : 1-27.

Ruckes, H. I960. New or little known Neotropical Pentatomids (Heter-

optera, Pentatomidae). Amer. Mus. Novit. 1996 : 1-27.

Ruckes, H. 1964. The genus Antiteuchus Dallas with descriptions of new

species (Heteroptera, Pentatomidae, Discocephalinae) . Bull. Amer.

Mus. nat. Hist. 127 : 49-102.

Sailer, R. I. 1952. A review of the stink bugs of the genus Mecidea .

Proc. U.S. Nat. Mus. 102 : 47-505.

Sailer, R.I. 1954. Interspecific hybridization among insects with a

report on cross breeding experiments with stink-bugs. J. econ.

Ent. 47 : 377-383.

Schaefer, C. W. 1964. The morphology and higher classification of the

Coreoidea (Hemiptera - Heteroptera) : Parts I and II. Ann. ent. Soc.

Amer. 57 : 670-684.

Scudder, G. G. E. 1957. The systematic position of Dicranocephalus Hahn,

1826 and its allies (Hemiptera : Heteroptera). Proc.R. ent. Soc.

Lond. (A) 32 : 147-158.

Scudder, G. G. E. 1959. The female genitalia of the Heteroptera:

76

North American Pentatomoidea

morphology and bearing on classification. Trans. R. ent. Soc.

Lond. 111(14) : 405-467.

Southwood, T.R.E. 1956. The structure of the eggs of the terrestrial

Heteroptera and its relationship to the classification of the group.

Trans. R. ent. Soc. Lond. 108(6) : 163-221.

Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north

of Mexico. Univ. Calif. Publ. Ent. 2 : 1-902.

Vidal, J. 1949. Hemipteres de L'Afrique du Nord et des pays bays

cir cum-mediterraneens. Mem. Soc. Sci. nat. Maroc 48 : 1-238.

Wagner, E. 1963. Unter suchungen uber den Taxonomischen Wert des

Baues der Genitalien bei den Cydnidae (Hem. Het. ) Acta ent. Mus.

Nat. Pragae 35 : 73-115.

Woodward, T. E. 1950. Ovariole and testis follicle numbers in the

Heteroptera. Entomologist's mon. Mag. 86 : 82-84.

Illustrations continued as separate.

CL

Publication of Qujestiones Entomologicae was started in 1965 as part

of a memorial project for Professor E. H. Strickland, the founder of the

Department of Entomology at the University of Alberta in Edmonton

in 1922.

if

It is intended to provide prompt low-cost publication for accounts of

entomological research of greater than average length, with priority given

to work in Professor Strickland’s special fields of interest including ento-

mology in Alberta, systematic work, and other papers based on work

done at the University of Alberta.

Copy should conform to the Style Manual for Biological Journals

published by the American Institute of Biological Sciences, Second

Edition, 1964, except as regards the abbreviations of titles of periodicals

which should be those given in the World List of Scientific Periodicals,

1964 Edition. The appropriate abbreviation for this journal is Quaest. ent.

An abstract of not more than 500 words is required. All manuscripts will

be reviewed by referees.

Illustrations and tables must be suitable for reproduction on a page

size of 9%X63/4 inches, text and tables not more than 7%X4% inches,

plates and figures not more than 8^X5 inches. Reprints must be ordered

when proofs are returned, and will be supplied at cost. Subscription rates

are the same for institutions, libraries, and individuals, $4.00 per

volume of 4 issues, normally appearing at quarterly intervals; single

issues $1.00. An abstract edition is available, printed on one side of

paper or on one or both sides (according to length) of 3X5 inch index

cards (at $1.00 per volume) or on 5X8 inch standard single row punched

cards ($1.50 per volume).

Communications regarding subscriptions and exchanges should be

addressed to the Subscription Manager and regarding manuscripts to:

The Editor, Quaestiones Entomologicae,

Department of Entomology,

University of Alberta, Edmonton, Canada.

Ql

W ILLUSTRATIONS

£ /

(IjAjXjL/d^Zisr^s^

McDonald - The genitalia of North America Pentatomoidea

(Hemiptera - Heteroptera) Quaest. Ent. 2: 77 — 150

LIST OF FIGURES

Figs, occur on page

Male genitalia no. in brackets.

Fig. 1. Generalized diagram of aedoeagus. (87)

Fig. 2. Cross section of vesica of Lampromicra senator showing convo-

luted duct. (87)

Fig. 3. Same enlarged. (87)

Figs. 4-7. Fohheria producta ; 4, pygophore, dorsal view; 5, clasper;

6, aedoeagus, right view; 7, vesica, dorsal view. (88)

Figs. 8-11. Eupychodera corrugata ; 8, pygophore, dorsal view; 9,

clasper; 10, aedoeagus, right view; 11, conjunctival appendages,

apical view. (88)

Fig. 12. Euptychodera corrugata ; vesica, lateral view. (89)

Figs. 13-17. Vanduzeeina balli ; 13, pygophore, dorsal view; 14,

clasper; 15, aedoeagus (unexpanded) left view; 16, conjunctival

appendages, left view; 17, vesica, ventral view. (89)

Figs. 18-20. Phimodera binotata ; 18, pygophore, dorsal view; 19,

clasper; 20, theca, right view. (89)

Figs. 21-22. Phimodera binotata ; 21, aedoeagus, apical view; 22,

vesica, left view. (90)

Figs. 23-26 . Eurygaster altemata ; 23, pygophore, dorsal view; 24,

clasper; 25, aedoeagus, left view; 26, vesica, right view. (90)

Figs. 27-29. Camirus moestus ; 27, pygophore, dorsal view28, clasper;

29, aedoeagus, right view. (90)

Figs. 30-31. Camirus moestus ; 30, vesica, left view; 31, convoluted

duct, ventral view. (91)

Figs. 32-36. Pachycoris torridus ; 32, pygophore, dorsal view; 33,

apex of pygophoral appendage; 34, clasper; 35, aedoeagus,

left view; 36, vesica, left view. (91)

Figs. 37-40 . Chelysomidea guttata ; 37, pygophore, dorsal view; 38,

clasper; 39, aedoeagus, right view; 40, vesica, left view. (91)

Figs. 41-46. Homaemus aeneifrons ; 41, pygophore, dorsal view; 42,

clasper; 43, aedoeagus, right view; 44, conjunctival appen-

dages, right view; 45, apex of third conjunctival appendage; 46,

vesica, lateral view. (92)

Figs. 47-51. Tetyra antillarum ; 47, pygophore, dorsal view; 48,

clasper; 49, aedoeagus, right view; 50, conjunctival appen-

dages, ventral view; 51, vesica, left view. (93)

Figs. 52-54. Sphyrocoris obliquus ; 52, pygophore, dorsal view; 53,

clasper; 54, aedoeagus, left view. (93)

Figs. 55-56. Sphyrocoris obliquus ; 55, apex of vesica; 56, vesica,

right view. (94)

Figs. 57-62. Stethaulax marmoratus ; 57, pygophore, dorsal view; 58,

clasper; 59, aedoeagus, left view; 60, vesica, dorsal view;

61, vesica, left view. (94)

Figs. 63-67 . Symphylus carribeanus ; 63, pygophore, dorsal view; 64,

clasper; 65, aedoeagus, left view; 66, median penal lobes,

ventral view; 67, vesica, left view. (95)

Figs. 68-70. Diolcus irroratus ; 68, pygophore, dorsal view;

69,

78

North American Pentatomoidea

clasper; 70, aedoeagus, right view. (95)

Figs. 71-72. Diolcus irroratus ; 71, aedoeagus, dorsal view; 72,

vesica, right view. (96)

Figs. 73-78. Acantholomidea porosa ; 73, pygophore, dorsal view;74,

clasper; 75, aedoeagus, left view; 76, conjunctival appendages,

ventral view; 77, apex of vesica; 78, vesica, left view. (96)

Figs. 79-83. Augocoris gomesii ; 79, pygophore, dorsal view; 80,

clasper; 81, theca, right view; 82, conjunctival appendages,

right view; 83, vesica, left view. (97)

Figs. 84-89 . Pentatoma rufipes ; 84, pygophore, dorsal view; 85,

ventral margin; 86, clasper; 87, aedoeagus, right view; 88,

median penal lobes, ventral view; 89, vesica, left view. (97)

Figs. 90-95. Dendrocoris humeralis ; 90, pygophore, dorsal view; 91,

genital plate; 92, pygophore, ventral margin; 93, clasper; 94,

aedoeagus, left view; 95, vesica, right view. (98)

Figs. 96-98. Piezodorus lituratus ; 96, pygophore, dorsal view; 97,

clasper; 98, aedoeagus, right view. (98)

Figs. 99-100. Piezodorus lituratus ; 99, median penal lobes, ventral

view; 100, vesica, left view. (99)

Figs. 101-105. Solubea pugnax ; 101, pygophore, dorsal view; 102,

clasper; 103, aedoeagus, right view; 104, median penal lobes,

ventral view; 105, vesica, right view. (99)

Figs. 106- 107. Peribalus limbolarius ; 106, pygophore, dorsal view;

107, clasper. (99)

Figs. 108-110. Peribcdus limbolarius ; 108, aedoeagus, rightview; 109,

median penal lobes, ventral view; 110, vesica, right view. (1(D)

Figs. 111-114. Trichopepla semivittata ; 111, pygophore, dorsal view;

112, ventral border; 113, clasper; 114, vesica, left view. (100)

Figs. 115-116. Mormidea lugens ; 115, pygophore, dorsal view; 116,

clasper. (100)

Figs. 117-118. Mormidea lugens ; 117, aedoeagus, ventral view; 118,

vesica, left view. (101)

Figs. 119-123 . Brepholoxa heidemanni ; 119, pygophore, dorsal view;

120, clasper, inner view; 121, clasper, lateral view; 122,

aedeagus, rightview; 123, vesica, left view. (101)

Figs. 124-126 . Arvelius albopunctatus ; 124, pygophore, dorsal view;

125, lateral margin, inner view of left side. (101)

Figs. 127-128. Arvelius albopunctatus ; 127, aedoeagus, left view; 128,

vesica, left view. (102)

Figs. 129-133. Aelia americana ; 129, pygophore, dorsal view; 130,

clasper; 131, aedoeagus, left view; 132, median penal lobes,

dorsal view; 133, vesica, left view. (102)

Figs. 134-135. Vulsirea violacea • 134, pygophore, dorsal view; 135,

clasper. (102)

Figs. 136-138. Vusirea violacea ; 136, aedoeagus, left view; 137,

median penal lobes, dorsal view; 138, vesica, left view. (103)

Figs. 139-143 . Acrosternum pennsylvanicum ; 139, pygophore, dorsal view;

140, ventral border; 141, clasper; 142, aedoeagus, dorsal view;

143, vesica, left view. (103)

Figs. 144-145. Chlorocoris subrugosus ; 144, pygophore, dorsal view ;

McDonald

79

145, dorsal margin. (103)

Figs. 146-147. Chlorocoris subrugosus ; 146, clasper; 147, vesica,

right view. (104)

Figs. 148-153. Carpocoris remotus ; 148, pygophore, dorsal view; 149,

genital plate; 150, ventral border ; 151, clasper; 152, aedoeagus,

left view; 153, conjunctival a pendages, dorsal view. (104)

Fig. 154. Carpocoris remotus ; vesica left view. (105)

Figs. 155-160. Nezara viridula ; 155, pygophore, dorsal view; 156,

ventral border; 157, clasper; 158, aedoeagus, right view; 159,

median penal lobes, ventral view; 160, vesica, right view. (105)

Figs. 161-163. Thyanta perditor ; 161, pygophore, dorsal view; 162,

clasper; 163, aedoeagus, right view. (105)

Figs. 164-165. Thyanta perditor ; 164, median penal lobes, dorsal

view; 165, vesica, right view. (106)

Figs. 166-172. Padaeus viduus ; 166, pygophore, dorsal view; 167,

clasper; 168, theca, left view; 169, aedoeagus, left view; 170,

median penal lobes, dorsal view; 171, median penal lobes,

ventral view; 172, vesica, right view. (106)

Figs. 173-174. Proxys punctulatus ; 173, pygophore, dorsal view; 174,

ventral border. (106)

Figs. 175-176. Proxys punctulatus ; 175, clasper; 176, aedoeagus,

apical view. (107)

Figs. 177- 180. Neottiglossa trilineata ; 177, pygophore, dor sal view; 178,

clasper; 179, aedoeagus, ventral view; 180, vesica, right view.

Figs. 181-184. Mur gantia histrionic a ; 181, pygophore, dorsal view;

182, clasper; 183, aedoeagus, left view; 184, median penal

lobes, ventral view. (107)

Fig. 185 . Mur gantia histrionic a ; vesica, right view. (108)

Figs. 186-191. Eysarcoris aeneus ; 186, pygophore, dorsal view; 187,

clasper, lateral view; 188, clasper, inner view; 189, aedoeagus,

right view; 190, first conjunctival appendages, ventral view;

191, vesica, left view. (108)

Figs. 192-195. Eysarcoris intergressus ; 192, pygophore', dorsal view;

193, clasper; 194, aedoeagus, right view; 195, conjunctival

appendages, dorsal view. (108)

Fig. 196 . Eysarcoris intergressus ; vesica, left view. (109)

Figs. 197-200 . Cosmopepla bimaculata ; 197, pygophore, dorsal view;

198, clasper; 199, aedoeagus, left view; 200, vesica, right

view. (109)

Figs. 201-203. Rhytidolomia senilis ; 201, pygophore, dorsal view;

202, clasper; 203, aedoeagus, right view. (109)

Figs. 204-205. Rhytidolomia senilis ; 204, aedoeagus, lateral view;

205, vesica, right view. (110)

Figs. 206-209. Rhytidolomia viridicata ; 206, pygophore, dorsal view;

207, clasper, inner view; 208, clasper, lateral view; 209,

aedoeagus, left view. (110)

Figs. 210-214 . Rhytidolomia saucia ; 210, ventral border of pygophore;

211, pygophore, dorsal view; 212, clasper; 213, aedoeagus,

left view; 214, median penal lobes, ventral view. (110)

Figs. 215-216. Chlorochroa ligata ; 215, pygophore, dorsal view; 216,

80

North American Pentatomoidea

clasper. (Ill)

Figs. 217-218. Chlorochroa uhleri ; 217, pygophore, dorsal view; 218,

clasper. (Ill)

Fig. 219 . Chlorochroa sayi ; clasper. (Ill)

Figs. 220-223. Banasa dimidiata • 220, pygophore, dorsal view; 221,

clasper; 222, aedoeagus, left view; 223, vesica, left view.

Figs. 224-229. Loxa flavicollis ; 224, pygophore, dorsal view; 225,

ventral border; 226, clasper; 227, aedoeagus, right view; 228,

aedoeagus, ventral view; 229, vesica, left view. (112)

Figs. 230-231. Menecles insertus ; 230, pygophore, dorsal view; 231,

ventral border. (112)

Figs. 232-235. Menecles insertus ; 232, clasper; 233, aedoeagus,

left view; 234, thecal processes, dorsal view; 235, median

penal lobes, ventral view; 236, vesica, right view. (113)

Figs. 237-239. Coenus delius ; 237, pygophore, dorsal view; 238,

clasper; 239, aedoeagus, left view. (113)

Fig. 240. Coenus delius ; vesica left view. (114)

Figs. 241-245 . Jiymenarcys nervosa ; 241, pygophore, dorsal view;

242, clasper; 243, aedoeagus, right view; 244, median penal

lobes, dorsal view; 245, vesica, right view. (114)

Figs. 246-247. Euschistus tristigmus ; 246, pygophore, dorsal view;

247, clasper. (114)

Figs. 248-250. Euschistus tristigmus ; 248, aedoeagus, right view; 249,

median penal lobes, dorsal view; 250, vesica, left view. (115)

Figs. 251-255. Prionosoma podopioides ; 251, pygophore, dorsal view ;

252, clasper; 253, aedoeagus, left view; 254, median penal

lobes, dorsal view; 255, aedoeagus, right view. (115)

Fig. 256. Prionosoma podopioides ; vesica, left view. (116)

Figs. 257 -261. Brochymena arborea ; 257, pygophore, dorsal view;

268, clasper; 259, aedoeagus, left view; 260, median penal

lobes, dorsal view; 261, vesica, right view. (116)

Figs. 262-263. Brochymena quadripustulata ; 262, pygophore, dorsal

view; 263, ventral margin. (116)

Figs. 264-267. Brochymena quadripustulata ; 264, clasper; 265, aedoeagus,

right view; 266, median penal lobes, dorsal view; 267, vesica,

left view. (117^

Figs. 268-212. Edessa bifida ; 268, pygophore, dorsal view; 269,

genital plate; 270, clasper; 271, aedoeagus, right view; 272,

vesica, left view. (117)

Fig. 273. Lineostethus clypeatus ; pygophore, dorsal view. (117)

Figs. 274-278. Lineostethus clypeatus ; 274, pygophore, dorsal view;

275, ventral margin; 276, clasper, inner view; 277, • clasper,

latteral view; 278, vesica, right view. (118)

Figs. 279-281. Sciocoris microphthalmus ; 279, pygophore, dorsal view;

280, clasper; 281, aedoeagus, dorsal view. (118)

Figs. 282-283. Sciocoris microphthalmus , ; 282, median penal lobes, left

view; 283, vesica, left view. (119)

Figs. 284-286. Mecidea longula ; 284, aedoeagus, left view; 285,

median penal lobes, dorsal view; 286, vesica, left view. (119)

Figs. 287-290. Zicrona caerulea ; 287, pygophore, dorsal view; 288,

McDonald

81

ventral border; 289, clasper; 290, aedoeagus, right view.

Figs. 291-292. Zicrona caerulea ; 291, median penal lobes, dorsal

view; 292, vesica, right view. (120)

Figs. 293-297. Oplomus tripustulatus ; 293, pygophore, dorsal view;

294, clasper; 295, aedoeagus, right view; 296, median penal

lobes, dorsal view; 297, vesica, left view. (120)

Figs. 298-299. Heterosceloides lepida ; 298, pygophore, dorsal view;

299, clasper. (120)

Figs. 300-302. Heterosceloides lepida ; 300, aedoeagus, left view; 301,

median penal lobes, ventral view; 302, vgsica, right view. (121)

Figs. 303-306. Rhacognathus americanus ; 303, pygophore, dorsal view;

304, clasper; 305, aedoeagus, left view. (121)

Fig. 307. Rhacognathus americanus j vesica, left view. (122)

Fig. 308-311. Apateticus hracteatus ; 308, clasper; 309, aedoeagus,

right view; 310, median penal lobes, dorsal view; 311, vesica,

right view. (122)

Figs. 312-313. Apateticus lineolatus • 312, pygophore, dorsal view;

313, clasper. (122)

Figs. 314-316. Apateticus lineolatus ; 314, aedoeagus, right view; 315,

median penal lobes, ventral view; 316, vesica, left view. (123)

Figs. 317-322. Podisus acutissimus ; 317, pygophore, dorsal view;

318, ventral margin; 319, dorsal margin; 320, clasper; 321,

aedoeagus, left view; 322, vesica, right view. (123)

Figs. 323-325. Podisus maculiventris ; 323, clasper; 324, aedoeagus,

right view; 325, vesica, right view. (124)

Figs. 326-329. Alcaeorrhyncus grandis ; 326, pygophore, dorsal view;

327, clasper; 328, aedoeagus, left view; 329, median penal

lobes, ventral view. (124)

Fig. 330. Alcaeorrhynchus grandis ; vesica, left view. (125)

Figs. 331-334. Euthyrhynchus floridanus ; 331, pygophore, dorsal view;

332, clasper; 333, aedoeagus, ventral view; 334, vesica, left

view. (125)

Figs. 335-336. Stiretrus anchorago ; 335, pygophore, dor sal view; 336,

clasper. (125^

Figs. 337-339. Stiretrus anchorago • 337, aedoeagus, right view; 338,

median penal lobes, dorsal view; 339, vesica, left view. (126)

Figs. 340-343. Mineus strigipes ; 340, pygophore, dorsal view; 341,

clasper; 342, aedoeagus, left view; 343, vesica, left view. (426)

Figs. 344-348. Andrallus spinidens ; 344, pygophore, dorsal view; 345,

clasper; 346, aedoeagus, left view; 347, aedoeagus, anterior

view; 348, vesica, lateral view. (127)

Figs. 349-352. Amaurochrous cinctipes ; 349, pygophore, dorsal view;

350, pygophoral appendage; 351, clasper, inner view; 352,

clasper, lateral view. (127)

Figs. 353-355. Amaurochrous cinctipes ; 353, aedoeagus, left view; 354,

median penal lobes, posterior view; 355, vesica, left view.

Figs. 356-360. Weda parvula ; 356, pygophore, dorsal view; 357,

clasper; 358, aedoeagus, left view; 359, median penal lobes,

dorsal view; 360, vesica, left view. (128)

Figs. 361-366. Oncozygia clavicomis ; 361, pygophore, dorsal view;

82

North American Pentatomoidea

362, clasper; 363, aedoeagus, right view; 364, aedoeagus,

dor sal view; 365, median penal lobes, lateral view; 366, vesica,

right view. ^129)

Figs. 367-369. Piezosternum subulatum, * 367, pygophore, dorsal view;

368, ventral border; 369, clasper. (129)

Figs. 370-372. Piezosternum subulatum ; 370, aedoeagus, leftview; 371

vesica, left view; 372, apex of vesica. (120)

Figs. 373-377. Meadorus lateralis ; 370, pygophore, dorsal view;

374, clasper; 375, aedoeagus, right view; 376, second conjunc-

tival appendages, ventral view; 377, vesica, lateral view. (130)

Figs. 378-380. Elasmostethus cruciatus ; 378, pygophore, dorsal view;

379, clasper; 380, aedoeagus, right view. (130)

Figs. 381-382 .Elasmostethus cruciatus ; 381, base of vesica, left view;

382, vesica, leftview. (131)

Figs. 383-387. Corimelaena pulicaria ; 383, pygophore, dorsal view;

384, clasper; 385, theca, dorsal view; 386, conjunctival appen-

dages, left view; 387, vesica, left view. (131)

Fig. 388. Sehirus cinctus ; pygophore, dorsal view. (131)

Figs. 389-390. Sehirus cinctus ; 389, aedoeagus, right view; 390,

vesica, left view. (132)

Figs. 391-397. Pangaeus aethiops • 391, pygophore, dorsal view; 392,

clasper; 393, clasper; 394, dorsal arm of clasper; 395, aedoe-

agus, right view; 396, apex of second conjunctival appendage,

ventral view; 397, vesica, left view. (132)

Figs. 398-401. Cyrtomenus crassus ; 398, pygophore, dorsal view;

399, clasper; 400, aedoeagus, left view; 401, vesica, left view.

Figs. 402-404. Melanaethus subglaber ; 402, pygophore, dorsal view;

403, aedoeagus, right view; 404, third conjunctival appendages,

ventral view. (133)

Fig. 405. Melanaethus subglaber ; vesica, right view. (134)

Figs. 40 6-410. Amnestus pallidus ; 406, pygophore, dorsal view; 407,

clasper; 408, aedoeagus (unexpanded), left view; 409, conjunc-

tival appendages, dorsal view; 410, vesica, left view. (134)

Female genitalia

Figs. 411-412. Fokheria producta • 4Ha genital chamber, dorsal view;

412, spermathecal bulb. (135)

Figs. 413-414. Euptychodera corrugata ; 413, genital chamber, dorsal

view; 414, spermathecal bulb. (135)

Figs. 415-416. Vanduzeeina balli ; 415, genital chamber, ventral view;

416, spermatheca. (135)

Figs. 417-418. Phimodera binotata ; 417a spermatheca; 418, sclerite

surrounding spermathecal opening. (135)

Fig. 419. Eurygaster alternata ; spermatheca. (136)

Fig. 420-421. Pachycoris torndus ; 420, groove in genital chamber,

dorsal view; 421, spermatheca. (136)

Figs. 422-423. Diolcus irroratus ; 422, pouch in genital chamber,

dorsal view; 423, genital chamber, ventral view. (136)

Fig. 424. Diolcus irroratus ; spermatheca. (137)

McDonald

83

Figs. 425-427. Tetyra antillarum. ; 425, ventral view of female genitalia;

426, spermatheca; 427, spermathecal bulb. (137)

Fig. 428. Symphylus carribeanus ; spermatheca. (137)

Figs. 429-430. Sphyrocoris obliquus • 429, spermatheca; 430, sperma-

thecal bulb. (137)

Figs. 431-432. Homaemus aeneifrons ; 431^ spermatheca; 432, sperma-

thecal bulb. (138)

Figs. 433-435. Acantholomidea porosa • 433a genital chamber, ventral

view; 434, spermatheca; 435, spermathecal bulb. (138)

Figs. 436-437. Chelysomidea guttata ; 436, genital chamber, ventral

view; 437, pouch of genital chamber, dorsal view. (138)

Fig. 438. Chelysomidea guttata ; spermatheca. (139)

Figs. 439-442. Stethaulax marmoratus ; 439, female genitalia, ventral

view; 440, groove in genital chamber, dorsal view; 441, sper-

matheca; 442, spermathecal bulb. (139)

Fig. 443. Augocoris gomesii ; spermatheca. (139)

Figs. 444-445 . Rhytidolomia senilis ; 444, sclerites surrounding open-

ing of spermathecal duct; 445, spermathecal bulb. (139)

Fig. 446 . Rhytidolomia viridicata • spermathecal bulb. (140)

Fig. 447. Banasa dimidiata ; spermathecal bulb. (140)

Fig. 448. Carpocoris remotus ; spermatheca. (140)

Figs. 449-450. Mur gantia histrionic a ; 449, female genitalia, ventral

view; 450, sclerites surrounding opening of spermathecal duct.

Fig. 451. Solubea pugnax • spermatheca. (140)

Fig. 452. Euschistus tristigmus ; spermatheca. (140)

Figs. 453-454. Cosmopepla bimaculata ; 453, female genitalia, ventral

view; 454, spermatheca. (141)

Figs. 455-ASG.Brepholoxa heidemanni ; 455, spermatheca; 456, sperma-

thecal bulb. (141)

Fig. 457. Dendrocoris humeralis ; female genitalia, ventral view. (141)

Figs. 458-459. Brionosoma podopioides ; 458, spermatheca; 459, sper-

mathecal bulb. (141)

Fig. 460. Solubea pugnax ; female genitalia, ventral view. (141)

Fig. 461. Solubea pugnax ; spermathecal bulb. (142)

Fig. 462. Neottiglossa trilineata ; sclerites around spermathecal opening.

Fig. 463. Loxa flavi colli s ; spermathecal bulb. (142)

Figs. 464-465. Arvelius- albopunctatus ; 464, spermatheca; 465, sper-

mathecal bulb. (142)

Fig. 466. Aelia americana ’ spermathecal bulb. (142)

Figs. 467-468. Acroslernum pennsylvanicum ) 467, female genitalia, ventral

view; 468, spermatheca. (142)

Figs. 469-470. Peribalus limbolarius ; 469, spermatheca; 470, sper-

mathecal bulb. (143)

Figs. 471-472. Vulsirea violacea ; 471, female genitalia, ventral view;

472, spermatheca. (143)

Figs. 473-474. Pentatoma rufipes- ; 473, female genitalia, ventral view;

474, spermatheca. (143)

Fig. 475. Chlorocoris subrugosus ; spermathecal bulb. (143)

Fig. 476. Trichopepla semivittata ; spermatheca. (144)

Fig. 477. Proxys punctulatus ; spermatheca. (144)

84

North American Pentatomoidea

Figs. 478-479. Thyanta perditor ; 478, spermatheca; 479, sperma-

thecal bulb. (144)

Fig. 480. Eysarcons aeneus ; spermatheca. (144)

Fig. 481. Brochymena quadripustulata ; spermatheca. (144)

Fig. 482. Edessa bifida ; spermatheca. (145)

Fig. 483. Sciocoris microphthalmus ; spermatheca. (145)

Fig. 484. Lineostethus clypeatus ; spermatheca. (145)

Fig. 485. Mecidea longula ; female genitalia, ventral view. (145)

Figs. 486-487. Mineus strigipes ; 486, female genitalia, ventral view;

487, spermathecal bulb. (145)

Fig. 488. Rhacognathus americanus ; spermatfteca. (145)

Figs. 489-490. Podisus acutissimus ; 489, spermatheca; 490, sper-

mathecal bulb. (146)

Fig. 491. Podisus maculiventris ; spermathecal bulb. (146)

Fig. 492. Apateticus lineolatus • spermatheca. (146)

Fig. 493. Perillus confluens ; spermatheca. (146)

Fig. 494. Alcaeorrhynchus grandis ; spermathecal bulb. (146)

Fig. 495. Euthyrhynchus floridanus ; spermatheca. (146)

Fig. 496. Zicrona caerulea ; female genitalia, ventral view. (146)

Fig. 497. Zicrona caerulea • sclerites around spermathecal opening.

Figs. 498-499. Amaurochrous dubius ; 498, female genitalia, ventral

view; 499, spermatheca. (147)

Fig. 500. Wedaparvula ; spermathecal bulb. (147)

Figs. 501-502. P iezo sternum subulatum- 501, female genitalia, ventral

view; 502, spermatheca. (147)

Fig. 503. Elasmostethus cruciatus ; female genitalia, ventral view. (147)

Fig. 504. Elasmostethus cruciatus • spermatheca. (148)

Fig. 505. Meadorus lateralis ; spermatheca. (148)

Figs. 506-507. Corimelaena pulicaria ; 506, female genitalia, ventral

view; 507, spermatheca. (148)

Figs. 508-509. Galgupha nitiduloides ; 508, spermatheca; 509, sperma-

thecal bulb. (148)

Fig. 510. Dallasiellus discrepans ; female genitalia, ventral view. (148)

Fig. 511. Dallasiellus discrepans ; spermatheca. (149)

Fig. 512-513. Cyrtomenus crassus ; 512, female genitalia, ventral view;

513, spermatheca. (149)

Fig. 514. Pangaeus aethiops; spermatheca. (149)

Figs. 515-516. Amnestus pallidus; 515, female genitalia, ventral view ;

516, spermatheca. (149)

Fig. 517. Amnestus pusio ; spermatheca. (149)

Figs. 518-519. Sehirus cinctus ; 518, female genitalia, ventral view;

519, spermatheca. (150)

Fig. 520. Graphical presentation of the degree of difference exis-

ting between genera of North American Scutellerinae using 7 male

and 4 female genitalic characters. (150)

McDonald

85

KEY TO LETTERING OF FIGURES

Figures 1—4 1 9 (male genitalia)

A. ch. , anterior chamber of

ejaculatory reservoir

A. s . , anterior sinus

A. th. pr., anterior thecal process

Ap. , apodeme

Ar. cl., arm of clasper

B. p. , basal plate

C. ap. , conjunctival appendage

1 C. ap. , first conjunctival

appendage

2 C. ap. , second conjunctival

appendage

3 C. ap. , third conjunctival

appendage

C. ap. 2 3 branch of second

conjunctival appendage

C. du. , convoluted duct

C. lo. , conjunctival lobe

Ca. , canal

Cal. , callus

Ce. s., central sinus

Cl., clasper

D. b. , dorsal border of

pygophore

D. dv. , dorsal diverticulum of

theca

D. c. ap. , dorsal lobe of

conjunctival appendage

D. c. lo., dorsal conjunctival

lobe

D. ch. , dorsal chamber of

ejaculatory reservoir

D. m. , dorsal margin of pygophore

D. pr., dorsal process of vesica

D. r., dorsal reservoir

Du. , duct

E. res., ejaculatory reservoir

En. d. , endophallic duct

En. f. , flange of endophallic duct

F. , flange around pygophoral

opening

G. pi. , genital plate

Gp. , secondary gonopore

In. r., inferior ridge

Kn. , knob

L. a. , lower arm of clasper

M. pr., median process

Me. p. , median penal lobe

Mu., muscle fibre

P. , proctiger

P. ch. , posterior chamber

of ejaculatory reservoir

P. th. pr., posterior thecal

process

Pi., pit

Pr., projection

Pro,, process

Py. ap. , pygophoral appen-

dage

R. , sclerotized ring at base

of endophallic duct

Ri., ridge

S. ve. pr., supravesical

process

Se. , septum

Se. d. , seminal duct

Sh. , sheath of vesica

Si., sinus

Sp. , spine

St. , setae

Su. r., superior ridge

Th. , theca

Th. ap. , thecal appendage

Th. f. , thecal flange

Th. pr., thecal process

Th. s., thecal shield

U. a., upper arm of clasper

V. b. , ventral border of

pygophore

V. c. ap. , ventral lobe of

conjunctival appendage

V. c. lo. , ventral conjunc-

tival lobe

V. ch. , ventral chamber of

ejaculatory reservoir

V. m. , ventral margin of

pygophore

Va., valve

Ve., vesica

Ve. f. , vesical flange

Ve. pr. , vesical process

86

North American Pentatomoidea

Figures 4! 1-5 19 (female genitalia)

A. s., accessory sac

An. , anal opening

B. , spermathecal bulb

B. d. , bulb of spermathecal duct

D. f. , distal flange of pump

Dl. , dilation of spermathecal duct

Dl. proximal chamber of

spermathecal dilation

Dl. 2j distal chamber of

spermathecal dilation

Dt. , diverticulum

FI. , flange

Gr., sclerotized groove in floor

of genital chamber

1 Gp. , first gonapophysis

2 Gp. , second gonapophysis

1 Gx. , first gonocoxa

2 Gx. , second gonocoxa

1. r., inner ramus

O. , opening of spermathecal

duct into genital chamber

O. r., outer rami

P. , spermathecal pump

P. f. 3 proximal flange of

pump

Pch. , pouch in genital

chamber

Pr., process of spermathecal

bulb

Pt. g3 paratergite eight

Pt. 9, paratergite nine

R.a sclerotized rod

R. sc., ring sclerite

S. ^q, sternum ten

S. du., spermathecal duct

Sc., sclerite

T. g, tergum eight

T r . , tr iangulum

87

I

0 05 mm

10 mm

90

D. b.

92

D. m.

4 I

CL

0-2 mm

44

1C. ap.

93

D. b.

0 '5 mm

Ve.

96

C. ap.

0-2 mm

76

R

E. res.

En.d.

78

1-0

Se. d.

97

O- 3mm

0 • 5 mm

100

Ve.

0*1 mm

UJUJ p.Q

102

V. m.

135

Se.d.

E. res.

0 • 2 mm

0-2

m m

E. res.

160

0 *5 mm

159

161

163

0 - 3 m m

106

I 65

0 • 4 mm

1 >Omm

0 • 3 m m

V. m.

0-2 mm

D. b.

0*3 mm

E. res.

1 • 0 mm

203

110

E. res.

0 • 5 m m

112

D.b.

0 *5 mm

113

1 • 0 mm 2 37

lC.ap.

0*5 mm

253

255

0 • 3 mm

0 • 2 mm

0 -2 m m

117

269

D. b.

118

0 *5 mm

E. res.

0’1 mm

283

119

Ve.

282

285

297

299

121

122

0* 5 mm

E . res.

0 • 4 m m

0 *2 mm

Me. p.

E . res.

339

Se.d.

34 3

0*2 mm

128

0'2 mm

360

0 • 1 mm

En. d.

129

130

386

387

Se.d.-^

E. res.

En.d.

D- m.

388

0*5 mm

Cl.

0-1 m m

132

0*2 mm

390

0*25 mm

134

O.i mm

•2 mm

136

0*5 mm

Dl.

137

Sc.

138

0*5 m m

0*3 mm

439

139

140

D.f.

0 • 2 mm

0 mm

0 -2 mm

4 70

143

— P. f.

0 • 5 m m

472

475

144

480

1 • 0 mm

488

146

n . i mm

147

0-1 mm

149

01 mm

150

Vamduzeeina balli

P himodera binotata

AugmmU gomesii

Number of character differences

6-8

□o-2 g3-5 3

9-11

1 2 and over

Quaest

lones

entomologicae

A periodical record of entomological investigations,

published at the Department of Entomology, Uni-

versity of Alberta, Edmonton, Canada.

VOLUME II

NUMBER 2

APRIL 1966

QUAESTIONES ENTOMOLOGICAE

151

A periodical record of entomological investigations, published at

the Department of Entomology, University of Alberta, Edmonton, Alberta.

Volume 2 Number 2 6 April 1966

CONTENTS

Editorial 151

Leech - The spiders (Araneida) of Hazen Camp

81°49'N, 7 1°18 1 W 153

Index to spiders 209

Book review 213

Corrigenda 214

Editorial - An eye for an I

Eye, not "the author", "the writer", or any other circumlocutory

description of myself, but eye, Brian Hocking, am publicly and without

shame pleading for the return of the first person, in all cases, singular

and plural, to its modest but respectable position in scientific literature.

This group of words provides the very brevity and precision that the pre-

sent-day world needs. To outlaw the whole of the first person simply

because some people have found difficulty in the discreetuse of the nom-

inative singular is, like total abstinence, an admission of weakness.

More than that, this false modesty must have cost the printing presses

of the world many millions of extra words over the last few years. Can

they or the reading public spare the time?

How delightfully simple this is, this eye that we shun so scrupu-

lously. Just one letter, and the simplest one at that. Just a line. Yet

surely this is the most precise line in the language; no pos sible ambiguity

here. It can mean only one thing. What about our alternative "the

author"? It is ten times as long for a start. In some scientific papers

many author s are referred to so that it becomes necessary to define him

further as "the present author". So many times does "the present author "

appear in some papers that eye come to regard him as the ever-present

author. Although his personality is rarely present, he is far more con-

spicuous and demanding than if he were just an eye.

English has got along for some time now without its second person

singular, although eye for one would welcome the return of the outspoken

"Where art thou?" and "Thou shalt not". The use of the plural form is

usually, after all, a false politeness, just as avoiding the eye is a false

modesty. If we are not to have all of our first persons restored, where

is it to end? And what have we left? Already nothing but second- and

third-rate personal pronouns. And there appears to be no authority for

all this; eye suspect an editorial conspiracy. This is what the author-

ities say:-

152

Perrin: "/ can be used wherever it is needed. People with only-

average concern for themselves need not worry; the conceited will give

themselves away anyway. Circumlocutions to get around the natural use

of 1 are usually awkward and likely to attract attention to themselves. "

(p. 599)

Quiller - Couch: "... when man asks questions about his fortune

or destiny he asks them most effectively in the first person. " (p. 141)

Gowers: "Official prose is made unnecessarily ugly by a shyness

of pronouns." (p. 71)

The only support eye can find for the outlawing of the first person

is in the Royal Society publication "General Notes on the Preparation of

Scientific Papers" . It says, referring only to a synopsis (p. 24): "Itis

preferable touse the third person", but elsewhere (p. 2): "It may seem

superfluous to state that the paper should be clear, precise, logical and

brief. . . Experience shows that clarity and precision are best achieved

by the use of short words and simple sentences. " Eye can find no com-

ments by Ander son and Thistle, or in The Canadian Government Editorial

Style Manual.

The last stronghold of the first person was for some time the ack-

nowledgment section of papers; here the occasional l and we still linger

on, tolerated - or could it be overlooked? - by our painstaking editors.

Perhaps this would be the best route of re-entry for the first person into

scientific paper s; surely it is here that clumsy circumlocutions are most

inappropriate. There are two other constructions in which an author may

still be able to sneak in a first per s on pronoun and get away with it: one

is the reference to one of a number of joint authors as "one of us" -

followed by the initials of the author referred to. The other is in the

explanation of italics in a quotation from another author; with the best

will in the world most editors boggle at the awful ambiguity of "author's

italics".

In view of the editor's privileged use of the first person plural it

is most unfitting that authors should be denied the right to use the sin-

gular. Editors, please, give us back our eyes.

Brian Hocking

Reprinted with permission from Entomology Division Newsletter,

Canada Department of Agriculture 32(6) : 1-2, 1954, withminor changes.

153

THE SPIDERS (ARANEIDA) OF HAZEN CAMP 81°49’N, 71°18’W*

R.E. LEECH

Department of Entomology Quaestiones entomologicae

University of Alberta , Edmonton 2:153—212, 196b

About 20,600 spiders (Araneida) from Hazen Camp (81°49’N, 71°18’W), Ellesmere Is-

land, Northwest Territories, Canada, were examined. These represent four families and thirteen

species: Dictynidae, Dictyna borealis Pickard-Cambridge; Lycosidae, Pardosa glacialis (Tho-

rell), Tarentula exasperans Pickard-Cambridge; Linyphidae, Collinsia spetsbergensis (Thorell),

Collinsia thulensis (Jackson), Cornicularia karpinskii ( Pickard-Cambridge ), Erigone psychrophila

Thorell, Hilaira vexatrix (Pickard-Cambridge), Meioneta nigripes (Simon), Minyriolus pampia

Chamberlin, Savignya barbata (Koch), Typhochraestus latithorax (Strand), and Thomisidae, Xysti-

cus deichmanni Soerensen. The known distribution of each species is listed. Detailed descrip-

tions of the rare species and drawings of the structures useful for identification of each species

and sex are given. An analysis of the seasonal occurrence of the adults of each species in 1964

and partial results from 1963, showed that all the species were active during the first three weeks

following the first day of spring melt, namely June 10 to June 30. Nine species inhabited the

humid terrestrial environment, one was present in all environments, and three only in arid envir-

onments.

Escape orientation of Pardosa glacialis was analyzed in relation to the sun and the

observer; reasons for the escape directions are given. The species Pardosa glacialis and Xysti-

cus deichmanni were found to be parasitized by Hexamermis sp. (Nematoda, Mermithidae).

The zoo geo graphical data indicate that there were one or more refugia at or near the

northern end of Ellesmere Island during the Wisconsin glaciation and perhaps for the entire Plie-

stocene epoch. Some of the extant insects and spiders were present in these refugia though most,

especially spiders restricted to night shadow areas, have probably immigrated since from more

southern localities.

The spiders (Araneida) from the high Nearctic Region are poorly

known in every respect - distribution, number of species, natural history,

and past history. Previous to this study, the largest single collection of

spider s from such a northern location as Hazen Camp (81°49tN, 71°18tW),

Ellesmere Island, Northwest Territories, Canada, was the collection

made by the Danish Peary Land Expedition of 1947-50, which collected

103 specimens of eight endemic species and one obviously introduced

species (Braendegaard I960). In the winter of 1898-99, the Second

Expedition of the "Fram" overwinter ed at R ice Strait, between Ellesmere

Island and Pirn Island (78°34‘N, 74°45,W) (Bryce 1910), and during the

warmer season of 1899, the ship's doctor (my infer ence) collected some

15 specimens of spiders of seven, possibly eight, species (Strand 1905,

and Braendegaard 1936). Collections made by Oliver and others in 1961

and 1962 at Hazen Camp yielded a possible ten species of spiders with

only five positive identifications (Oliver 1963).

The purpose of this paper is to give a detailed account of the spiders

from Hazen Camp. The account includes data on the taxonomy and natural

history, and theories (based on an analysis of the evidence) of the zoo-

geographic history of the spiders and other Arthropoda at Hazen Camp.

* An investigation associated with the programme ‘Studies on Arctic

Insects', Entomology Research Institute, Canada Department of Agri-

culture (Paper No. 24).

154

Spiders of Hazen Camp

THE STUDY AREA

The general biology and taxonomy of spider s was studied during the

summers of 1963 and 1964 at Hazen Camp, Ellesmere Island, Northwest

Territories, Canada (81°49tN, 71°18tW). This is approximately 150 km

southwest of Alert, and is on Lake Hazen, the largest fresh water body

(78 x 11 km) in the Queen Elizabeth Islands . The study area is on the mid

northwest shor e of Lake Hazen and extends along the shore for about 5 km

and away from the shor e for about 3 km. The confines are the Snow Goose

River delta, Blister Creek delta, and Mt. McGill. The range of altitudes

in the study area is from 158 to 1050m and includes the following ecological

areas: clay plains and slopes, sand, gravel, alkaline clay, Dryas hum-

mocks, Dryas-Kobresia tundra, marshes, muddy delta, gravel delta, boulder

talus slopes, and springy slopes (based on Savile1 s personal notes of 1962,

and Savile 1964, see fig. 1).

Biological work was started at Hazen Camp in 1957 and 1958 (Powell

1961), but entomological studies were not started until 1961 when Donald

R. Oliver (Entomology R esearch Institute, Ottawa) did general collecting

and studied pond ecology.

Hazen Camp was opened as an International Geophysical Year

stationin the late summer of 1957, and since then much information has

been published about the Camp and the Lake. Christie (1962) has des-

cribed the geology; Savile (1964) the ecology and vascular plants; Day

(1964) and Yong (1961, et al. 1962) have discussed and analyzed the soil

characteristics; Oliver (1963) has discussed the Insecta and Arachnida

collected to the end of 1962; Jackson (1959 , I960) has analyzed the

meteorological conditions ; Powell (1961) has di scus sed the vegetation and

microclimate; Harington (I960) reported on snow and ice conditions for

the winter of 1957-58; and Hatter sley-Smith (1964) published a biblio-

graphy of "Operation Hazen", covering the years 1957-63.

Soil and Soil Conditions

Soil samples were collected in 1963 from the major habitats as

defined by Savile (1964). The depths of permafrost at 12 sites in mid

August varied from 40. 6 to 99. 0 cm with a mean of 73 . 5 cm. Yong et al.

(1962) at a comparable period in 1962 reported a mean depth for perma-

frost of 51. 0 cm.

Early in the season, just after the snow has melted, the ground

surface becomes a quagmire, but with the sun in view continuously, it

becomes firm in two to three weeks except in mar sh or pond depressions.

The period when the snow and ice leaves the surface of the ground will be

referred to as "spring melt".

Very little of the surface soil is without frost-heave cracks. The

cracks vary from 0. 1 to 5 cm wide, from 1. 5 to 40 cm deep, and from

10 cm to several metres long. Organic content of the soils varied from

0. 3 to 19. 6% with a mean of 4. 1%. The pH was usually between 7. 4 and

8.6. Soil temperature maxima at the surface were generally 7 . 5 to 16. 5°C

higher than air temperature maxima on sunny days, and minima were

2. 5 to 5. 5°C warmer than air minima on overcast days (Powell 1961).

On June 6, 1964, one day befor e the spring melt, I recorded the . following

155

156

Spiders of Hazen Camp

temperatures in bright afternoon sun on a 20° south-facing slope: 2. 5

cm above soil surface. ... 2. 7°C, at surface. ... 6. 3°C, 2. 5 cm below

soil surface. ... 9. 9°C.

In appearance, the soil is mostly sandy to sandy-clayey with

moderate to sparse vegetation cover.

Vegetation

There are about 115 species of vascular plants recorded from the

Lake Hazen region, but there are only a few abundant ones. These are

Salix arctica Pall. , Dryas integrifolia . M. Vahl. , Kobresia myosuroides (Vill. ) Fiori

and Paol. , Carex aquatilis Wahlenb. var . stans (Drej) Boott. , Cassiope tetragona

(L. ) D. Don. , and in restricted areas, Eriophorum Scheuchzeri Hoppe, and

E. triste (Th. Fries) Hadac and Love. The remaining plant species are

sparsely distributed throughout the study area.

Climate and Weather

Detailed records of the weather havebeenmade for the years 1958,

1961, 1962, 1963, and 1964. The lowest winter temperature recorded

by the minimum thermometer at Hazen Camp is -70. 6°F (-57.0°C) during

the winter of 1963-64 (personal record). Figure 2 is a graph of the

cumulative day-means of Stevenson screen temperatures to date since

June 1 at Hazen Camp, 100 metres from the lake shore, 1.8 m above

the ground, and 161 m above sea level.

For the latitude, the summer temperatures are exceptionally high,

often higher for long periods than at many nearby coastal stations. The

inland position of Lake Hazen accounts for the higher temperatures. The

lake isina very stable high pressure trough region, andis in the shadow

of the Garfield Range to the north and northwest.

Precipitation at Lake Hazen is very light. Jackson (1959 , p. 95)

recorded 0.98 inches (2.48 cm) water equivalent during the period August

1957 to August 1958, and the stationat Alert recorded 4.52 inches (12. 8

cm) during the same period. Snowfall between September and May

accounted for 91% of the precipitation at Lake Hazen.

Despite the low precipitation at Lake Hazen, considerable moisture

is available for the plants and animals. During most summer s (1964 was

an exception) the melting of snow and ice from the surface is sufficiently

slow that the ground surface is dry for only two weeks before the water

from permafrost melting comes to the surface. As the permafrost water

percolates to the surface, depressions that were full of water during the

spring melt and which had dried up during the course of the summer,

refill.

The wind speed averages at Lake Hazen are very low with over 76%

of the yearly observations being 5 mph or less, 16% from 6-10 mph, and

8% 11 mph or over. The majority of the winds come from the northeast,

along the Lake Hazen trough (Jackson 1959 , p. 125).

Relative humidities in 1962 and 1963 were read every 6 hours at

1.6 m above the ground, and give dailymean figures of 80-88% for June,

and 76-80% for July and early August (Savile 1964, p. 240), but were

much lower during the summer of 1964 because of continuous high winds

from the southwest. The prevailing low- speed winds permitted conditions

157

3Nnr I 3DNIS 3iva Ol SNV3W-AVCJ 3AllVinWnD

20 30 IO 20 30

158

Spiders of Hazen Camp

of almost 100% relative humidity to exist at or near the surface where

the shearing effect of the ground on the wind caused little disturbance of

the air at one foot (30cm) or less (composite data, Jackson 1959 , pp..

127-130, and my notes).

During the summer Lake Hazen enjoys cloudless periods for two

to three weeks without interruption.

Night Shadow Areas

In 1963 and 1964 I observed that several species of the spiders were

restricted to the night shadow areas. These areas are in the mountain

shadow for a minimum of about four to six hours during each 24 hour

period at the warmest part of the season, and longer earlier and later in

the season. The shadow period is synchronous with night-time in more

temperate regions at this longitude. The shadow area extends from the

shoulders of Mt. McGill to the line marked off as map coordinates K3 to

K6, J7 to J9 to B9 and all the area encompassed (see fig. 1).

MATERIALS AND METHODS

Previous to 1963, possibly ten species of spider s were known from

the Hazen Camp area from fewer than two hundred specimens. The

program "Studies on Arctic Insects" was instigated by D.R, Oliver of

the Entomology Research Institute, Ottawa, and has dealt so far with

insects from Isachsen, Ellef R ingnes Island, (McAlpine 196 5), and Hazen

Camp, Ellesmere Island, (Downes 1964; Oliver 1963). I was

given permis sion to study the spider s. from the Hazen Camp area. Studies

were begun at the end of June, 1963, and continued until the end of August,

1964, with an interruption during the winter of 1963-1964.

Materials

The structure, identification, and distribution of 20, 534 spiders

comprising 13 species collected during two summers within the study

area were examined. All were collected in the Hazen Camp area. Iden-

tified also were 751 spider s from Melville Island (collected by Larry Law),

36 spiders from Tanquary Fjord, Ellesmere Island (collected by Guy

Brassard), 18 spiders from Bathurst and Cornwallis Islands (collected

by Leonard Hills), and 54 spiders from Thule, Greenland (collected by

me). About 522 individuals of two species of Lycosidae were studied in

an attempt to determine the length of life cycle.

The field equipment included 50 aluminum cake pans, 23 x 23 x

6.5 cm, used in 1963 and 37 pans used again in 1964. Each pan contained

the following fluid mixture to a depth of 2 cm: 600 ml water, 400 ml

ethylene glycol, 5 ml formalin, and 1-2 ml of any liquid detergent,

Identifications in the field laboratory were made with the aid of a

Wild M5 binocular dissecting microscope with a maximum power of 50

diameters. Identifications, drawings, and analyses in the laboratory

were made with a Leitz binocular dissecting microscope with a maximum

power of 150 diameters and a 100 watt zircon arc lamp ("Mikrark Illum-

inator", made by the Boone Instrument Corporation of New York). A

Leech

159

Leitz eyepiece grid 10 mm square divided into 0. 5 mm squares and a

10 mm eyepiece micrometre scale with 100 divisions were used in con-

junction with millimetre graph paper in preparing the drawings.

The meteorological equipment of Hazen Camp was set up and used

throughout the study periods. Equipment included corrected maximum

and minimum thermometers, a Feuss corrected millibar barometer, a

hygr ©thermograph, and an anemometer.

A pair, of each species will be sent to the following institutions or

persons: American Museum of Natural History, New York; Zoologisk

Museum, Kystalgade, Copenhagen; Zoological Institute, Uppsala Univer-

sity, Uppsala; Laboratoire de Zoologie, University of Toulouse, Toulouse;

Dr. Hermann Wiehle, Dessau; Museum of Comparative Zoology, Har-

vard University; Department of Entomology, University of Alberta,

Edmonton. The remaining specimens will be deposited in the Canadian

National Collection, Ottawa. Ten males and 10 females of Tarentula

exasperans Pickar d- Cambridge, 1877, will be sent to the Zoologisk

Museum, Copenhagen.

Methods

The study area was examined for the principal ecological zones

(based on Savile^ notes of 1962). In 1963, a total of 50 traps was placed

at carefully selected sites and in 1964,37 traps were used. Eight of the

sites in the habitat of some of the les s-frequently collected species were

used in both years. The new traps of 1964 were in areas not examined

in 1963.

The traps were examined once every four days. This interval was

selected in 1963 in order to fit into a previously established work pattern,

and retained in 1964 for purposes of continuity. Each trap was emptied

of spiders and insects and the fluid was replaced or added to.

The traps were set so that the lip of the pan was flush with the

ground level. There was usually very little sand drift except in some

sites because wind speeds were low. Traps placed in low regions near

streams were often flooded by water and the specimens lost. The biggest

problem was caused by foxes and wolves which would urinate and defecate

into the traps, then scratch sand and any loose vegetation into them. I

have interpreted this to mean that they dislike either the pans or their

contents. Oliver (pers. comm.) and B. Hocking (pers. comm.), after

observing similar behaviour in these animals, have interpreted these

actions in the same way. There did not seem to be anyway to solve this

problem!

Spiders from each trap were preserved and kept in separate vials

by trap and by day. Individual spiders were identified to species in the

field laboratory. The results of examination of this material are recorded

by species on graphs in the text. Identifications were checked, and num-

bers of individuals per sex per day per trap were also recorded at this

time. My analysis of a species habitat is based on where the immatures

and females of a species were collected, asmales wander. Overwinter-

ing sites were used as further evidence of the usual habitat of a species.

Except for four species, the immature stages were not identified as

identification of immature stages can never be positive. Individuals

160

Spiders of Hazen Camp

with anomalies in epigyna and pedipalp organs were examined for mer-

mithid (Nematoda) or other parasites.

All measurements and drawings were made from the microscope.

The pertinent sexual parts of large spiders were drawnat 54 diameters,

and for the smaller species, at 150 diameters. Measurements of all

parts were recorded with each drawing.

Measurements of carapaces were made from directly above the

spider. Length is the distance from the base line between the posterior

median eyes posterior ly along the midline to the incurve of the hind edge

of the carapace. Carapace width was measured at its widest part. The

opisthosoma was measured from the dorsal aspect. Total length of the

spider was measured from the dorsal aspect from the base line between

the anterior median eyes to the end of the opisthosoma.

Measurements of legs and leg parts were always made on the actual

dorsal side of the leg. Measurements were made from the proximal to

the distal part of a leg segment and did not include the membranes at the

joints. Trichobothrium is abbreviated in the text as Tm. The position

of a trichobothrium on the metatarsus is expressed as the ratio of its

distance from the proximal joint to the total length of the metatarsus.

This is expressed as a decimal fraction.

Dictynidae

Dictyna borealis Pickard-Cambridge, 1877 p ■ 273 (Figs. 38, 39)

D. borealis: Bonnet 1956 p. 1431; Holm 1958b p. 534; Braendegaard

I960 p. 7; Chamberlin and Gertsch 1958 p. 136 (in part). Dictyna sp.

Oliver 1963 p. 176.

Notes on taxonomy - Chamberlin and Gertsch (1958, p. 137) place a

few specimens collected by H.W. Levi in the high mountains of Colorado

in this species, but the tips of the emboli of those specimens differ in

shape from the emboli of specimens collected at Lake Hazen. The latter

are definitely members of the species borealis. Further, the Colorado

specimens are considerably larger than any of the Lake Hazen specimens .

Gertsch (pers. comm., 1965) has confirmed my opinion that the Colorado

specimens are not borealis Pickard-Cambridge, even though they are

similar and probably related to the species.

Natural, history - This species is a member of the arid arctic faunal

element (Braendegaard 1946) and is heliophilic. Specimens are most

frequently found on dry, south-facing slopes exposed to the sun, with a

vegetation consisting mainly of Dryas integrifolia , but often of Cassiope tetragona .

It prefers hummocked, almost wind-free areas. It overwinters in the

vegetation on the surface.

Figure 3 shows the main period of activity of the males of this

species. The females are rarely wanderers. The main activity of the

males is dir ectly correlated with the courtship and mating periods of the

species. The adults are not known to overwinter.

Courtship was not observed in this species, but it cannot be long

nor involved, as virgin males and females introduced to one another wer e

found mating within a 45 min lapse of observation. Previous to mating,

the males assumed small territories which they defended against all

numbers of individuals

Leech

161

other males, but which females seemed almost coaxed to enter. Defen-

ding a territory consisted of actively fighting all male intruders.

Four pairs of spiders were observed mating and both of the male

pedipalpi wer e inserted into the female at the same time. The male was

positioned ventral to the female, with the carapace of the male almost

touching the prosomatic sternum of the female. There did not seem to

be any specified angle as two pairs were lying on their side, the third

pair positioned with the female upside down, and the fourth pair with the

female rightside up. Mating continued for about 30 min, then each of the

four pairs began separating. They did not recouple. Males did not mate

more than once.

162

Spiders of Hazen Camp

In 1963, I observed that females of Dic.tyna borealis almost invariably

deposited their egg sacs and built their webs on the south- and southwest-

facing sides of Dryas integrifolia hummocks and in the vegetation, but never

on the ground. About 30 egg sacs were seen. Laboratory specimens in

1964 also laid eggs in the vegetation. Females remained near the eggs

until they hatched. Ten egg sacs were examined and found to contain

from eight to thirteen eggs, with a mean of 8. 7 eggs per sac.

This species appears to be able to overwinter at any stage except

the adult.

When offered a choice of over 30 species of Diptera, this species

prefer red small ones. Chir onomidae and Ceratopogonidae were the main

preferences . Cyclor rhaphous Diptera were always refused. No parasites

or predators of this species were found.

Material examined - About 625 adults of this species were examined,

and all were from Hazen Camp area.

Distribution - Greenland (Peary Land; E. Greenland, 68-77°; W.

Greenland, 60-79°N). Ell esmere Island (Hazen Camp area). Bernard

Harbour, (58°48'N, 114'W)3 Mackenzie District, N. W. T.

This species appears to have a relict distribution (fig. 4), though

as it is able to balloon, it should be found in more of the Near ctic R egion.

Fig. 4. Distribution map of Dirtyna borealis .

Leech

163

Lycosidae

Pardosa glacialis (Thorell) 1872b,p ■ 159 (Figs. 32-34)

Lycosa glacialis Thor ell 1872b, p. 159 . P. glacialis: Bonnet 1958 p. 3371;

Oliver 1963 p. 176. L. glacialis : Holm 1958b, p. 529; Braendegaard I960

p. 8.

Notes on taxonomy - This species is a member of the genus Pardosa , and

not of the genus Lycosa . The characters of both generaare summarized

in Kaston (1948, pp. 321, 331). The egg sacs of Pardosa glacialis are a pale

green-blue and are lenticular. Those of all known Lycosa are white and

spherical.

Natural history - Habitat - This species belongs to the euryoequous (eur-

yecious) arctic faunal element (Braendegaard 1946). Specimens are

found everywhere except in windy places. P. glacialis overwinters in the

soil in cracks and under stones on the surface. It has not been found in

the overwintering sites at depths greater than 2. 5 cm. It does not bur-

row. Individuals of this species drown easily, hence they will not be

found overwintering alive in areas that are inundated or soaked by spring

melting of snow and ice. Apparent migrations or mass movements of

individuals in the spring are really the successful overwintering indi-

viduals radiating from winter quarters.

Figure 5 shows the main period of activity of the males of this

species to be between June 29 and July 6. The activity period of males

is directly correlated with the courtship and mating period of the species .

The adults are not known to overwinter.

Natural history - Courtship - The courtship and courtship preliminaries

were observed from the beginning of the season. On June 29, 1964, the

eighth day after moulting from the penultimate instar, captured adult

males and females suddenly became active in courting. It was noted that

the males would court under natural conditions only if heat and light were

sufficient. In the laboratory, it was noted that courting started or stopped

if a 100 watt bulb were brought to 25 cms or taken away to 75 cms.

When courting was first observed, the males began holding a small

territory, and would defend this against all intruders except females.

In all, 63 males were observed courting females, 46 in the laboratory

and 17 under natural conditions. No variation in courtship was observed.

The males were often seen rubbing the substrate with the venter of

the opisthosoma, but sperm webs were never seen. Evidently some

species of lycosids spin sperm webs (Gertsch 1949, p. 73) and some do

not (Savory 1928, p. 224).

Courtship is summarized as follows: on June 29, 1964, a male

was observed in the beginnings of courtship. The palpi, bent downward

at the patellae, were moved in a circular motion which, when viewed

from above, appeared clockwise in the right palpus, and anticlockwise

in the left. Simultaneously, the fir st pair of legs were lifted and extended

horizontally forward, then gradually relaxed while extended. When the

'tarsi of the first legs touched the ground, the palpi stopped churning.

The palpi started rechurning when the legs, brought back toward the cara-

pace and raised, started extending forward.

164 Spiders of Hazen Camp

9 12 16 20 24 28 2 6 10 14 18 22 26 30 3 7 II 15 19

June July August, 1964

Associated with these waving motions was a characteristic weaving

of the body. First, the waving motions were started in a position which

squarely faced the female, who was usually some 10 cm distant. The

male then turned 30° to the left and vigor ously made the waving motions,

turned toward the female and repeated the waving, then turned 30° to the

right and again repeated the waving motions, then centre, then left. . .

until within two cm of the female, at which point the weaving was cut to

two positions, each about 20° from centre. The right-left weave and

associated waving motions were continued vigorously until either the

female chased the male away, or until their legs touched, at which point

the female suddenly as sumed a defensive position with fangs spread open

Leech

165

and the first two pairs of legs raised up and forward. In this position,

the female char ged forward for about one cm, and the male fled for about

15 cm, then turned and again started the same advance procedures. Al-

most invariably, the males approached the females from the front.

On June 30, 1964, most of the males were dead. Mating had pre-

sumably taken place in the late afternoon of the previous day.

Previous authors have argued whether the male 's courtship reaction

was precipitated by sight, smell, contact, or a combination of all three.

It is my opinion that courtship by individuals of P. glaciaiis was initiated

by chemo and contact stimuli rather than by sight or touching of the

ground over which the females had passed. The opinion is based on the

following observations . Virgin males placed incages that had pr eviously

held females did not become "excited", but virginmales placed in cages

with females present became "excited" by leg contact with the females

and began courting within 25 minutes. The initial reaction of the males

after contact was to withdraw to a corner and start cleaning the whole

body, legs, and palpi. The cleaning usually lasted about 20 minutes.

The males then ventured out slowly, and at first fled at the approach of

either a male or a female. Courting began shortly thereafter. Osterloh

(1922) rightly concluded that the necessary stimulus for male spiders is

different in different species. The condition of the male and female

should be known when the observation was made. For instance, had ob-

servations at Hazen Camp been made only on males that had previously

touched females, the conclusions might have been that sight is the "trig-

gering" mechanism.

Mating was never observed in P ■ glaciaiis , but one male which had

repeatedly been shunned by all females, began courting a large male

Chironomid, which was lying on its side almost dead. The male even-

tually mounted the fly in the usual Pardosa manner (Kaston 1936, p. 167),

it then discovered the mistake and ate the fly.

Natural history - Egg laying - Within 48 hours after mating, the females

were ready to lay eggs. Four females were observed during the whole

egg-laying process. Because there were no observable differences during

the egg-laying, the general pattern is described as follows. The female

located a sheltered flat place which lay pocket-like between and under

several rocks. She then star ted making a thin flat sheet of webbing about

2.5 cms in diameter. In the centre of this the female made a small,

much-thicker patch of webbing about 0. 8 cm in diameter which was of a

different silk material than the main sheet web. The centre patch was

slightly green and opaque. These preparations took about 30 min.

The female then placed her genital region over the small centre

patch and started laying eggs at the rate of one egg each 50-60 seconds

until the usual 50-53 eggs were laid. Each female appeared to rest for

about tenminutes, thenmade the covering for the sac. The silk material

for the upper and lower halves of the egg sac was the same. The upper

half of the sac was attached to the lower half with such firmness that the

lower half became somewhat bowl- shaped when only half the upper cover-

ing was made.

The female made the cover for the upper half by attaching a thick

166

Spiders of Hazen Camp

silk line to one side of the centre patch, then swung the opisthosoma up

and over the eggs to the far side. As the silk lines were attached, the

female rotated about the eggs making a complete cover. This operation

lasted about 35 minutes. Again the female rested before cutting the egg

sac free with either the chelicerae or endites. The female then turned

and placed the spinnerets over the sac and attached them to it. The

female then waited for over an hour, emerged from the hole with the sac

attached, and the opisthosoma tilted up so that the sac did not drag on

the ground.

I was not able to keep any of the caged females alive until the young

emerged. Whenever a female died, I examined the egg sac to note the

development of the eggs, but in no case were any young seen. On August

19, 1964, a female was captured with young that had just hatched from

the egg sac. These were the only young seen of this species that hatched

in 1964.

Number of instars and longevity - On the basis of analysis of mensural

data obtained from 318 individuals of this species, it appears that there

are seven instars from the egg to the adult (fig. 6). The first instar is

spent inside the egg sac, and the second instar emerges from the egg

sac and crawls onto the opisthosoma of the mother. If it is assumed that

on the average each instar lasts one year, then the length of the lifecycle

of this species is about six years.

Solar escape orientation - Three groups of specimens were used for the

experiment, one group encountered in the field and left alone except for

the period of encounter, and two groups that were captured. One of the

captured groups was kept outside where the sun could be seen on clear

days. This group was maintained in order to assure having a control

group to contrast with the group kept in the laboratory. The second

captured group was maintained in the laboratory with a 100 watt bulb

shining from the geographic east about 25 cm from the cage. The pur-

pose of maintaining this group was to see if the solar orientation could

be interrupted or disturbed, and that this occurred is apparent from the

results summarized in figures 7, 8 and 9. The longer the specimens

were kept in the laboratory, the greater became their confusion when

released. Directions were often so unsure that a spider would turn almost

360° in a 5 cm circle before slowly going in a direction, and then it was

as often towards as away from the sun rather than at right angles to it,

as seen from the charts of the specimens kept outside or encountered in

the field (figs. 7, 8, 9). Each specimenused in the laboratory experi-

ment was kept inside for a minimum of nine days before beingused in the

experiment. In no case was the same spider tested more than once in

a three hour period, as it was found that individuals became tired and

gave different initial results at each escape.

Experiments were begun on July 2, 1964. Only adults or subadults

were used as the younger spiders showed preference for well-protected,

vegetated areas and their escape reactions were either to remain still

or else hide in the vegetation. Parasitized adults and subadults wer e also

omitted from the experiment when it was found that their escape r eactions

Leech

167

were considerably altered. The parasitized spiders show little or no

escape reaction. In fact some refused to move unless prodded with a

stick.

Fig. 6. Frequency distribution of

dimensions of Pardosa glacialis.

168

Fig. 7. Escape directions of Pardosa glacialis in the field observed under natural

conditions .

169

N

N

opacity 6 , no surety of direction

captured spec, kept outside

August 7, 1964 ; 1205-12' 15

opacity 6 , no surety of direction

captured spec, kept in laboratory

August 13, 1964 ; 11-40-11. 50

sunny , captured spec kept outside

Fig. 8. Escape directions of specimens of Pardosa glacialis kept in sunlight

and specimens exposed to a 100 watt lamp shining from the east .

170

August 13,1964-, 11:30-11: 35 AM.

no surety of direction

captured spec, kept in laboratory

Fig. 9. Escape directions of specimens of Pardosa glacialis kept in sunlight

and specimens exposed to a 100 watt lamp shining from the east.

Leech

171

The results of the Lake Hazen experiments are as follows:

Pardos a glacialis encountered in the field or kept under somewhat natural

conditions attempted to escape at approximately 90° right or left to the

sun's position. The group kept in the labor atory showed, in time, almost

complete disorientation of escape direction. Individuals encountered in

the field and placed in the shade before being startled, escaped directly

away from me, but upon entering the sunlight, turned and ran at 90° to

it. Aged or senile individuals tended to show less and less orientation

as the season progressed. Cool weather also inhibited escape reactions

as spiders apparently tried to get warm rather than escape.

I suggest that the escape direction of Pardos a glacialis in relation to

the sun - that is, to the right or left - is not intrinsic to each individual,

but a function of the direction in which the spider is facing, at rest,

immediately prior to the time of the escape. The spider 's resting position

is one that will allow it to present most of its body to the sun at one time.

In mid-afternoon, for example, this would mean on a line running north-

west to southeast, and from the graphs it can be seen that most of the

spiders tried to escape to the northwest (figs. 7.2, 4, 6; 8. 1) or south-

east (fig. 7.2, 4, 6) .

Papi and Syr jamaki ( 1963) have conducted similar experiments with

Arctosa cinerea (Fabr . ) (Lycosidae), but have obtained different results.

They have combined results of testing periods of rather long duration --

usually six hours. I feel that for these experiments long periods obscure

theresults. Therefore, I have recorded results for 15-25 minute periods

so that variations in the spiders' reactions can be more easily observed

in relation to variation in the sun's position. Tests were made in the

mid-morning and again in the late afternoon. Further, no theoretical

escape direction has been assumed or considered.

Food, parasites & predators - This species fed most readily on Chiron-

omidae, and when hungry, fed on the smaller cyclorrhaphous flies. Blow

flies and other flies of this size were left untouched. The species is also

highly cannibalistic, but was not observed feeding on other species of

spiders .

Remains of P. glacialis were found in the crops and gizzards of sev-

eral snow buntings and knots. No pr evious information about vertebrate

predators of this species was found.

Various degrees of parasitic castration in m a 1 e and female

P. glacialis by nematodes of the genus Hexamermis (Mermithidae) have been

observed at Hazen Camp. About one per cent of the specimens collected

were infected and most of these were females. Possibly more careful

examination of all the Hazen Camp specimens of P. glacialis might reveal

a considerably higher rate of parasitism, fora parasitic infection is very

hard to detect in the young spiders.

The ultimate effect of the parasite on the spider is death, as just

before Hexamermis emerges from the opisthosoma of the spider, the essen-

tial organs of the spider are eaten. Spiders examined just after a parasite

had emerged were found to be lacking in the main prosomatic muscles,

the entire digestive system, fat body, and the entire reproductive system.

An infected spider usually stopped feeding about one week before the

172

Spiders of Hazen Camp

parasite emerged, and during the last week such spider s were seen drin-

king quantities of water.

When the parasite was about to emerge, the spider crawled into a

dark hole or corner. It took about 20 minutes for Hexamermis to emerge

completely from the anterior end of the opisthosoma. The spider died

30-60 minutes before the Hexamermis first emerged.

Some of the obvious external morphological characteristics for

Hexamermis infectionare these: lopsided or greatly enlarged opisthosoma;

epigynum altered from the normal; legs shorter and thicker; sluggish

or inactive spider; and some secondary sexual characteristics of the

male not present or bar ely developed. A normal male appear s gaunt and

thin, but a parasitized male appears fat like a female full of eggs.

Parasitic castration in lycosids by Mermis has also been described

byAke Holm (1941) and some examples that might be caused by parasites

are cited and illustrated by Kaston (1961, 1963a, 1963b).

Material examined - Approximately 3383 adults of this species were

examined from Hazen Camp, one female from Payne River (59o30'N),

Quebec, four males from "Manitoba", one male from Umiat, Alaska,

four females and three males from Mesters Vig, E. Greenland, six

females from Holsteinborg (approx. 63°N), W. Greenland, and one fe-

male from Axel-Heiberg Island (Heinz Rutz, collector, 1963).

Distribution (fig. 10) - Greenland (East, 68-77°N; West, 61-75 °N;

Peary Land, 82-83°N). Ellesmere Island (76-82°N). Axel-Heiberg

Island (79°25 'N, 90°45'W). Baffinlsland. Southampton Island. Manitoba.

Umiat, Alaska. Payne River (59°30'N), Quebec.

Pardosa glacialis appears to be a nearctic species only.

Fig. 10. Distribution map of Pardosa glacialis .

Leech

173

Tarentula exasperans Pickard-Cambridge, 1877, p. 283 ; (Figs. 35-37 )

Arctosa exasperans: Bonnet 1955 p. 647; T. exasperans: Oliver 1963 p.

176, Braendegaard I960 p. 8.

Notes on taxonomy- There has been some confusion about the classifi-

cation of this species, mainly because of the scarcity of specimens in

museums. Gertsch (1934) and Braendegaard (I960) have correctly re-

placed this species in the genus Tarentula .

Description - Braendegaard (I960, p. 10) has described the female

of this species and has measured the sizes of a male and a female. I

have remeasured a Peary Land specimen and a number of the Hazen

Camp specimens, and find Braendegaard 1 s measurements a little more

than half of mine. Table 1 shows the measurements of individuals of

this species from Hazen Camp.

TABLE 1 - Mean dimensions of Tarentula exasperans in mm.

'Natural history - This species is a member of the arid arctic faunal

element, and is the most pronounced heliophile of all the species found

at Hazen Camp. T. exasperans was taken only on dry southwest and south-

facing slopes (rarely on southeast-facing) in and near clumps of Dryas

integrifolia . Where this species is abundant, P. glacialis was almost never

found except for occasional wandering males. T. exasperan's overwinters

by burrowing about 2. 5 cm into the ground at the bases of Dryas- integrifolia .

It was never found in the night shadow areas or areas of slopes of more

than 20°.

Figure 1 1 shows the main period of activity of the males and females

of this species. The females are never as active in wandering as the

males. Sexual activity of the males is between June 28 and July 6, though

they can be found before and after these dates. The adults are not known

to overwinter.

Newly emerged males and females were collected on the day they

emerged, and then kept in separate cages for one week. On June 29*

1964, a male was introduced to a cage containing four females. Upon

contact with a female, the male seemed to become excited. There was

a short sparring contact, then each fled in different directions. The

female went for about 5 cm and stopped, but the male began running about

in small circles and figure-eights as though injured, with the front two

pairs of legs drawn up against the carapace. Upon each contact with a

female, the scurryings were intensified.

174

Spiders of Hazen Camp

On June 30, 1964, two males and two females were placed in a

large cage outside. On July 5, the males began courting the females.

In all, five males were observed courting females, and there were no

obvious differences. In this pattern of courtship the male approached

the female, and at contact became more active and began circling and

scurrying. Thereafter, the male approached the female almost invariably

from behind, and tapped the female on the opisthosoma or fourth leg with

his first legs. The female merely lifted a leg, and the male scurried

off, but quickly returned and tried again. On the ninth or tenth try, the

male retired for about five minutes.

The above procedures were watched for over five hours continu-

ously, but no males were successful at mounting a female. The males

were found dead the following morning. There was no mating observed

for this species, nor even any partial attempts at mounting.

On July 6, 1964, a female and egg sac appeared in the outside cage.

The process for egg-laying is as follows. The female T. exasperans laid

eggs and made the egg sac in much the same way as Pardosa glacialis (see

p. 165). The difference was that T. exasperans dug a hole in the ground

about 2 cm deep and at the bottom of the hole made a round cavity about

1 cm in diameter. Once in the hole, the female closed over the entrance

with webbing. The hole was completely lined with silk. The whole pro-

cess, including the hole digging, took about four hours, but the female

often remained inside the hole for another three to six hours. When the

female emerged, the light brown egg sac was attached to the spinnerets.

The egg sacs were larger and rounder than those of Pardosa glacialis . There

are about 70 eggs per sac of T. exasperans . There is no great size differ-

ence between egg sacs.

The young were never observed clustered on the opisthosoma of

the female as were the young of P • glacialis . About 204 specimens of

various instars were examined to determine the number of instars and

length of life cycle. The results were poor, mainly because there were

a great many adults and penultimates, but very few of the preceding in-

stars. However, by inspection, it appears that there may be as many as

seven or eight instars, and a life cycle that may last six or seven years,

if it is assumed that each instar lasts about one year.

No escape orientation was observed in Tarentula exasperans males or

females, as the species relies on cryptic colouration rather than speedy

retreats to elude enemies and predators. The gray and black colour-

ation makes individuals almost invisible when they are in or near the

dead leaves of Dryas integrifolia . Specimens observed in the field did not

run at my approach but remained still. Even when the ground was shaken

under them, they moved only if their positions were somewhat precarious.

It was found by experiment that this species prefers the smaller

Diptera so abundant at Hazen Camp, though Collembola will be eaten if

caught. Small blowflies ( Phormia and Protocalliphora spp. ) were refused

even when the wings were cut off, perhaps because T. exasperans has very

small chelicerae for a spider of its size. No parasites or predators of

this species were observed, nor was cannibalism seen.

Material examined - About 397 adults of this species were examined

numbers of individuals

175

10 12 16 2 0 24 28 2 6 10 14 18 22 2 6 3 0 3 7 II 15 19

Fig. 1Z. Distribution map of Tarentula exasperans .

176

Spiders of Hazen Camp

from Hazen Camp, one male from Peary Land, Greenland, which was

loaned by the Zoological Institute in Copenhagen, three females and one

male from Umanak, Greenland, loaned by the American Museum of

Natural History, New York, one male and seven immatures from Tan-

quary Fjord, Ellesmere Island, 10 immatures from Axel-Heiberg Island

(Heinz Rutz, collector, 1963), and three males and 34 immatures from

Melville Island (J.E.H. Martin, collector, 1965).

Distribution (fig. 12) - Greenland (Peary Land; Umanak, 70° 40'N;

Saunders Island, 76°35’N, 69°45'W). Ellesmere Island (Discovery Har-

bour, 81°45'N; Hazen Camp; Tanquary Fjord, 81°28'N, 76°50'W). Axel-

Heiberg Island. Melville Island (74°58'N, 115°00'W). This species is

known only from the high Nearctic Region.

Linyphi idae

Collinsia spetsber gensis (Thorell) 1872 (Figs. 65, 69)

Erigone spetsbergensis Thorell 1872 p. 692. Typhochraestus spitsbergensis:

Bonnet 1959 p. 4747, C. spetsbergensis: Holm 1960b p. 512; C. spitsbergensis:

Braendegaard I960 p. 11.

o

Notes on taxonomy - According to Ake Holm (pers. comm. 1965),

Thorell's paper (1872) was written in Swedish, and since Spetsberg^n is

Swedish and Spitzbergen is German, the spelling spetsbergensis must stand

as valid, despite Bonnet's (1955, p. 74; 1959, p. 4747) comment to the

contrary.

Description - Female. Color: carapace brown, marked and shaded

with dark brown; chelicerae pale yellow-brown with brown specks on the

basal half; sternum brownish; labium brown, rimmed with gray; pedi-

palp coxae brownish, gnathobases pale gray, abdomen gray-brown; spin-

nerets gray-brown.

Structure: size, moderately small, about 2.25 mm long; carapace

distinctly longer than wide, about 0. 86 mm x 0. 69 mm, gradually rising

to the cephalic region, then sloping downward to the eyes; cephalic ldbe

lacking; clypeus height about 3. 5 to 4. 0 diameters of an anterior median

eye; posterior row of eyes slightly procurved; posterior medians slightly

smaller than the laterals and all about equally spaced at 2. 0 diameters

of one median; anterior row almost straight, but slightly recurved;

anterior medians slightly more than 2.0 diameters of one from the later-

als; median ocular quadrangle longer than wide and wider posteriorly;

promargin of cheliceral fang groove armed with five stout teeth; cheli-

cerae slightly reclined; legs moderately long.

Tibiae I — III each with two spines; tibia IV with one spine at 0.41;

trichobothrium (Tm) I about 0. 64; Tm II about 0. 61; Tm III about 0. 46;

Tm IV lacking.

Male. Color and structure: like those of the female, except for

the following: total length about 2. 0 mm; carapace longer than wide,

about 0. 80 mm x 0. 74 mm.

Tibia I— III with two spines; tibia IV with one spine; Tm I about

0.56; Tm II about 0.50; Tm III about 0.47 or 0.48; Tm IV lacking.

Leech

177

Natural history - This species is a member of the humid arctic faunal

element (Braendegaard 1946). Specimens are usually found in river

deltas in areas with fine, muddy sand covered with dense Equisetum and

some Salix . These areas are always very wet or damp, and are occasion-

ally flooded in the spring. If the ground begins to crack with dryness,

then C. spetsbergensis retreats into these cracks.

The adults are found throughout the season, though there is a slight

increase of captured adults at the end. The species overwinters at the

bases of the vegetation but not, so far as is known, as adults.

No parasites or predators were observed preying on this species,

but it can be assumed that the young are eaten by other species and by

their own adults.

Material examined - About 22 adults were examined from the Hazen

Camp material, two females from Marie Bay, Bathurst Island (Leonard

Hills, collector, summer, 1964), three females and two males from

Bailey Pt. , Melville Island (J. E. H. Martin, collector, 1965), 164 adults

from Weatherhall Bay, Melville Island (Larry Law, collector, summer,

1964), one male from Is achsen, Ellef Ringnes Island (J.F. McAlpine,

collector, I960), two females from Alert, Ellesmere Island (personal

collection, 1963) and four males and 17 females from Axel-Heiberg

Island (Heinz Rutz, collector, 1963).

Distribution (fig. 13) - Alaska (Arctic Coast). Marie Bay, Bathurst

Island. Weatherhall Bay and Bailey Point, Melville Island. Hazen Camp

and Alert, Ellesmere Island. Isachsen, Ellef Ringnes Island. Green-

land (Peary Land; E. Greenland, 62-65°N; W. Greenland, 70°N). Ice-

land. Spitsbergen. Sweden. Novaya Zemlya. Siberia. New Siberian

Islands. This species is high Holarctic in distribution.

Fig. 13. Distribution map of Collinsia spetsbergensis .

numbers of individuals

178

Spiders of Hazen Camp

Collinsia thulensis (Jackson) 1934^, p. 614 (Figs. 66-68 )

Coryphaeolanus thulensis Jackson 1934 pp. 614, 615, 618; C. thulensis :

Bonnet 1956 p. 1231. Collinsia thulensis: Holm 1958a p. 48; b p. 531 ,

1960a p. 112; Braendegaard I960 p. 12.

Natural history - This species is a member of the humid arctic faunal

element. Specimens are most commonly found in gravelly parts of river

deltas with scanty vegetation, mostly Dryas integrifolia and Salix arctica .

C. thulensis , in contrast to C. spitsbergensis , is active mostly during the early

part of the season. Overwintering forms were not found, but it appears

from the habitat that the species overwinter s on the surface of the ground,

perhaps under some of the stones or in the vegetation.

The active breeding period is indicated in figure 14. Note the low

number of adults caught at the end of the season. From this I assume

that the adults do not overwinter.

Captured specimens kept in cages fed readily on mites and Col-

lembola, but refused all flies offered, including very small Cerato-

pogonidae.

8 12 16 20 24 28 2 6 10 14 18 22 26 30 3 7 II 15 19

June July August, 1964

Leech

179

Material examined - About 100 adults of this species were examined

from Hazen Camp, six females and one male from Thule, Greenland

(personal collection 1964), and one female fro*n Axel-Heiberg Island

(Heinz Rutz, collector, 1963).

Distribution { fig. 15) - Kotzebue, Alaska. Hazen Camp, Ellesmere

Island. Axel-Heiberg Island. Greenland (Thule; Peary Land; and

between 70-75°N in E. Greenland). Spitsbergen.

This species appears to have an Holarctic distribution.

Fig. 15. Distribution map of Collinsia thulensis.

Cornicularia karpinskii (Piekard-Cambridge) 1873, p. 447 (Figs. 51-53)

Erigone karpinskii Pickard- Cambridge 1873 p. 447; C. karpinskii: Bonnet

1956 p. 1ZZ3; Holm 1960a p. 113, b p. 513.

Notes on taxonomy - As suggested by Holm (1958a), C. karpinskii seems

to be a complex of species whose components are not understood, or

else this is a polytypic species. None of the 30 males examined showed

any variation in the tibial apophysis, a feature that is variable in some

other populations of this species (Holm 1958a, pp. 53, 54). The only

observed variable feature in this species was the two lobes of the epigy-

num, whose proportions of length and width varied slightly.

* Examinations of the holotype of Cornicularia clavicornis Emerton (1882, Trans. Conn. Acad.

Arts Sci. 6 : 1-86) shows that the Lake Hazen material should be referred to this species. Details

will be published later.

180

Spiders of Hazen Camp

Description - Male. Color: carapace pale yellow-brown; eyes ringed

with dark brown; legs pale yellow; opisthosoma pale gray- green with four

red-yellow spots on the dorsum; sternum golden brown with brownmar-

gin; chelicerae golden brown; labium pale brown with gray margin.

Structure: size medium small, entire length about 2.31 mm;

carapace distinctly elongate, 0. 96 mm long x 0. 72 mm wide; carapace

gradually rising to the head part, dropping forward and down to the an-

terior median from the posterior median eyes, then the clypeus drops

vertically from the anterior medians; horn placed midway between the

anterior and posterior medians, projecting forward and upward, barely,

if at all, extending beyond the vertical face of the clypeus; horn with a

greater diameter distally than basally; height of clypeus about 3. 5 to

4. 0 diameters of an anterior median eye; chelicerae vertical or nearly

so, perhaps slightly reclined; stridulation organ on lateral sides of

chelicerae distinct; eyes equal or subequal in size; posterior row de-

cidedly procurved and all eyes equally spaced at about 1. 8 diameters of

one posterior median; anterior medians about 0.20 diameters of one

apart, and about 0.80 diameters of one from the laterals.

Sternum longer than wide and with a sparse cover of thin hairs;

legs not strikingly long or short; tarsal claws with a full complement of

teeth, and resembling a comb; the two tibial apophyses of the pedipalp

elongate and projecting forward and down atop the cymbium; median

apophysis curving down and forward under the lateral, then continuing

parallel but ventral to it; median apophysis bifid terminally, one broad,

flat, lateral projection with blunt spines pointing outward below and beyond

the lateral apophysis, and the other pointed and running parallel to the

terminal part of the lateral apophysis; the lateral apophysis curved

slightly to the median line, then turned outward at the terminal one-

third, ending in a blunt point; the embolus and other parts of the tarsus

within the cymbium as in figure 52. Anterior mar gin of the opisthosoma

protruding over the carapace; four small, pale red-yellow depressions

on the dor sum of the opisthosoma, the anterior pair closer together than

the posterior; average of five measurements of the opisthosoma is 1. 35

mm.

Tibia I - II with two spines; tibiae III-IV with one spine each at 0. 20

and 0. 19 respectively; Tm I about 0.53; Tm II about 0.50; Tm III about

0. 47; and Tm IV about 0. 31.

Female. Color and structure: Like the male except that the horn

is lacking; average length of five females is about 2. 55 mm, carapace

0. 93 mm, and the opisthosoma about 1. 60 mm; tibiae I — III with two

spines; tibiae IV with one spine at 0. 16 to 0. 17; Tm I about 0.46-0.47;

Tm II about 0.46 to 0.47; Tm III about 0.47; Tm IV about 0.50.

Natural history _ This species is a member of the humid arctic faunal

element. It lives in the cracks in the ground and ventures onto the ground

surface only when the relative humidity is above 90%. The soil is cal-

careous with sparse vegetation. The adults and young were found deep

in the cracks in the ground where therelative humidity approached 100%.

No overwintering sites were found, but I assume that individuals over-

winter fairly close to the ground surface.

numbers of individuals

181

June July August, 1964

Fig. 17. Distribution map of Comicularia harpinskii .

182

Spiders of Hazen Camp

Figure 16 shows the frequency distribution of this species during

the summer of 1964. There are no comparable d a t a from 1963.

C. karpinshii appears to be able to overwinter in the adult stage, as adults

were caught before the spring melt. No parasites or predators were

found for this species.

Material examined - About 78 specimens of this species were examined

from Hazen Camp, and two from the Aleutian Islands, Alaska.

Distribution (fig. 17) - Unalaska Island and Umnak Island, Alaska.

Banff, Alberta. Yellowstone Park, Wyoming. NewYork. Newfoundland.

Akpatok Island, Ungava Bay, N. W. T. Lake Hazen, Ellesmere Island.

East and West Greenland. Iceland. TheFaeroes. England and Scotland.

The Swiss Alps. Northern Scandinavia. Spitsbergen. Franz Joseph

Land. Novaya Zemlya. Waigatsch Island. Lake Baikal, Siberia. Kam-

chatka.

This species is circumpolar in distribution, though, as mentioned

in the notes on taxonomy, it is not certain that this distribution represents

only one species.

Erigone psychrophila Thorell, 1872ap ■ 689 (Figs. 42-44)

E. psychrophila: Bonnet 1956 p. 1772; Holm 1958a p. 52, b p. 532,

1960a p. 116, b p. 513; Braendegaard I960 p. 12; Oliver 1963 p. 176.

Natural history - This species is a member of the humid arctic

faunal element. Erigone psychrophila is restricted to vegetated, marshy

areas at the edges of ponds and quiet streams and to water-saturated,

vegetated slopes. These data do not quite agree with Holm (1958a, p.

53) who states that E. psychrophila belongs to both the dry and humid faunae.

The species apparently overwinters in the vegetation and can often be

found moving under water in the slush snow during the spring melt.

Figure 18 shows the main activity period and summer distribution

in numbers. The males are very active during the mating season, then

are scarce thereafter. The sharp drop in the number of females caught

in early July can be attributed mostly to the females secluding themselves

while egg laying. The drop-off in early August might be attributed to

death of the females and to inactivity because of cool weather. The adults

are apparently able to overwinter as they are found at the very beginning

of the season.

No parasites or predator s of this species were found, but I assume

that as in the case of all these small spiders, they are prey to the larger

spiders.

Material examin ed - About 1983 adult specimens of this species were

examined from Hazen Camp, nine females, thr ee males and six immatur es

from Cornwallis Island (Leonard Hills, collector, 1964), one male from

Thule, Greenland (personal collection, 1964), 355 adult specimens from

Melville Island (Larry Law, collector, 1964), and one male, one female

and one immature from Mould Bay, Prince Patrick Island ( J . E. H. Martin,

collector, 1965).

numbers of individuals

183

Fig. 19. Distribution map of Erigone psychrophila .

184

Spiders of Hazen Camp

Distribution (fig. 19) - Coastal Alaska (Arctic and Bering) . Weather-

hall Bay, 75°46'N, 106°56 'W) , Melville Island. Mould Bay, Prince Patrick

Island. Mari® Bay, Bathur st Island. Cornwallis Island. Alert and Hazen -

Camp, Ellesmere Island. Greenland (Peary Land; East Greenland,

67-77°N; West Greenland, 74-77°N). Iceland. Spitsbergen. Northern

Scandinavia. Novaya Zemlya. Waigatsch Island. Franz Joseph Land.

New Siberian Islands. Kamchatka. This species is circumpolar in dis-

tribution.

Hilaira vexatrix (Pickard-Cambridge) 1877, p. 280 (Figs. 45, 46)

Erigone vexatrix Pickard-Cambridge 1877 p. Z80. Hilaira vexatrix: Bonnet

1957 p. 2214; Holm 1958b p. 532; Braendegaard I960 p. 14.

Notes on taxonomy _ Schenkel (1950) lists a record of this species from

Banff, Alberta. Without seeing the Alberta specimens, I cannot agree

that this species has anything but a high arctic distribution. Holm (1956)

does not list Schenkel's reference, nor does he give any comment about

an Alberta record. All previous records for this species are above 70°N

latitude.

Natural history - This species is a member of the humid arctic faunal

element, as it is found only in damp, vegetated regions that are rarely,

if ever, inundated. Many specimens were collected in the damp, upper

edges of ponds and small streams that have dense vegetation and many

rocks under which they may crawl to overwinter. Specimens of

Hilaira vexatrix overwinter under rocks and in cracks in the ground about

one to two cm deep. They are active on the ground as soonas the ground

temperature is above freezing, even though the air temperature is well

below freezing.

Figure 20 shows the activity periods of this species during the

summer. The June 16 to 24 peak is the period of courting and mating,

and the peak at the end of the season is the increase of adults that will

overwinter. Therefore, the peaks belong to two distinct populations of

adults. The adults of this species overwinter.

Neither courtship nor mating were observed in this species; the

males died about five days after mating.

On June 15, 1964, two females laid eggs which were in small,

lenticular, white egg sacs. On June 18, a third female laid eggs. The

egg sacs were suspended in tangle webs about one cm above the ground.

The females remained with the eggs until after the young had emerged.

One of the females ate the male after mating.

On July 3, the small spiders were visible inside the egg sacs, and

on July 12, the young from the eggs laid on June 15 emerged. On July

20, the young from the third sac emerged. The three sacs contained 9,

11, and 8 eggs respectively. The egg sacs were kept at a constant 100%

relative humidity.

The females did not feed and were not fed for a period of six weeks,

and at the end of this period showed no signs of stress. Collembola wer e

introduced as food, but the females showed no inter est. All three females

were dead by August 10.

numbers of individuals

185

4 8 12 16 20 24 28 2 6 10 14 18 22 26 30 3 7 II 15 19

June July August, 1964

Fig. 21. Distribution map of Hilaira vexatrix .

186

Spiders of Hazen Camp

No parasites or predators of this species were found. A great

number of the immatures die by cannibalism.

Material examined. - About 2159 adults of this species were examined

from Hazen Camp, two males and eighteen females from Thule, Green-

land (personal collection, 1964) 63 males and 91 females from Axel-

Heiberg Island (Heinz Rutz, collector, 1963), and one male, 18 females

and 17 immatures from Melville Island (J. E. H. Martin, collector, 1965).

Distribution (fig. 21) - Greenland (Peary Land; Thule). Ellesmere

Island (Alert; Hazen Camp; Discovery Harbour). N. E. Coast of Baffin

Island. Axel-Heiberg Island. Melville Island. Arctic Coast of Alaska.

Banff, Alberta (?).

This species is Nearctic. The chances of it being found in the

Palearctic Region are slight, as Braendegaard (1958) has studied the

Iceland “material, Locket and Millidge (1951, 1953) have studied the

British material and Wiehle (1956, I960) has studied the German material.

*Meioneta nigripes (Simon) 1884, p. 439 (Figs. 62-64)

Microneta nigripes Simon 1884 p.439. Meioneta nigripes^B onnet 1957 p. 2756;

Braendegaard 1958 p. 80, I960 p. 15; Holm 1958a p. 56, I960 b p. 513.

Natural history - This species is a member of the humid arctic faunal

element. Individuals live deep in soil cracks or under medium to large-

sized stones on very dry south- to southwest-facing slopes. That is,

the macroclimatic conditions are dry, but the microclimatic conditions

are humid. Braendegaard (1946, I960) considers this species to be

euryoequous (euryecious), but examination of the microclimate leaves

no doubt that it is a humid arctic species.

Overwintered adults were collected on June 1, 1964, before the

spring thaw. Inactive females wer e collected from under rocks that were

frozen to the ground surface. These females became active within ten

seconds of the time they were collected and exposed to the sun. Figure

22 shows the peak of activity for the species at the beginning of the season.

Courtship was observed in several pairs of this species and no

variation was seen. Observations were started on June 1, 1964. Males

and females were placed in a small bottle with soil and rocks. Random

wandering was observed for several hours, after which time the males

selected areas that they would mildly defend. These were small areas

in which each male had built a small, horizontal, almost invisibly-thin

sheet web about 2x4 mm, and from which the males hung upside down.

Each male remained on this sheet for about 15 minutes. These were,

I believe, the sperm webs, though no sperm droplets were seen. Even-

tually, each male searched for and built a small tangle web near a female.

When the web was built, each male began a combination of activities as

follows: each palpus was jerked forward and back alternately, much like

two pistons. At the same time, the whole body was jerked back and for th.

Coordinated with these was a gradual approach towards the female. About

three mm from the female, the male began body- jerking and strumming

one front foot, then the other. Then the female started the same sort of

* Examinations of the holotype of Meioneta maritima (Emerton) New combination (1919, Rep. Can.

Artie Expedition 1913-18, 3 : 4H) shows that the Lake Hazen material should be referred to this

species.

Leech

187

motions. When their front legs touched, the female fled with the male

in jerky pursuit.

Fig. 22. Frequency distribution of

Meioneta nigripes , summer, 1964.

males

a— — females

n= 139

VA — A A A,

A

na— -a

-# — $

8 12 16 20 24 28 2

June

6 10 14 18 22 26 30 3 7 II 15 19

July August , 19 64

Fig. 23. Distribution map of Meioneta nigripes.

188

Spiders of Hazen Camp

During the next 45 minutes, the males and females engaged their

front legs for brief moments, then broke contact. At 90 minutes, one

of the females built a small tangle and sheet web, then hid in one corner

of the web. In a few minutes, the female was found by the male. The

male began improving the web and the two finished the preparations in

eight minutes. The web was made while both were upside down.

At this point the male and female stopped and remained still for

five minutes. Gradually, with increasing vigour , the male began jerking

in the web, and shortly after the female began strumming. Slowly, the

male stopped jerking and began strumming, and approached the female as

hedidso. Then the palpi began pushing back and forth like pistons . (Both

spiders were still hanging upside down from the sheet web.) When their

legs touched, the female did not scurry away, but relaxed the front two

pairs of legs so that the prosoma hung down from the web.

In mating, the male came forward so that his carapace touched the

sternum of the female. At the same time, the right palpus shot forward,

grappled with the epigynum, and lifted part of it away. The haematodocha

expanded and the embolus twisted spirally into the spermathecal duct.

The right palpus was engaged for one second, then the left palpus was

applied for the same length of time. During the next 18 min 25 sec the

male alternated the palpi 380 times. At no time was the male held captive

by the female.

The male left the female for 40 seconds and retired to a small

corner of the web where pos sibly the sperm were replenished in the palpi.

The male then rejoined the female and during the next 6 min 41 sec,

alternated the palpi 150 times before again returning to the corner to

replenish the sperm. Returning to the female, the male again alternated

the palpi on the epigynum, but now more slowly. In 6 min, only 40 al-

ternations; at 11 min, only 6 more times; and in the last 33 min, the

palp were alternated 37 more times. The last 6 couplings took about

two min each. The female effected the finish. Shortly thereafter, the

two fled in opposite directions.

More pairs were observed mating. The females seemed to be eager

to mate several times with any male and often tried, but no male could

be persuaded to engage a female - any female - more than once.

Four days after mating on June 5, 1964 two females laid eggs in

small, round, white egg sacs. The egg sacs were attached directly to

the side of the container. As with H. vexatrix , the eggs were kept in 100%

relative humidity. More eggs were laid by other females on July 12 and

August 24.

On July 3, 1964, eight young emerged from each egg sac laid on

June 5. During the development of the eggs, the females stayed within

two to three cm of the eggs.

On July 12, Collembola were introduced to the females and young

for food. The females fed immediately. I also observed that females

captured and bit several Collembola, leaving them inactive in the tangle

web about the egg sac, and later returned to feed on them. The young

spiders appeared to be too small to feed on Collembola. Most of the

young died by cannibalism. The females of this species refused to eat

anything but Collembola and Acarina, even though they were offered

Leech

189

small Diptera and spiders. No parasites or predators of this species

were found.

Material examined - About 157 adults of this species were examined

from Hazen Camp, and one male and 13 females from Bailey Point,

Melville Island (J. E. Martin, collector, 1965).

Distribution (fig. 23) - Ellesmere Island (Hazen Camp). Bailey

Point, Melville Island. Greenland (Peary Land; W. Greenland at 63°N;

E. Greenland from 63-70°N). Iceland. Spitsbergen. Jan Mayen Island.

The Faeroes. Scotland. North Sweden. Novaya Zemlya. The French,

Swiss, and Tyrolian Alps.

This species is mainly Palearctic in distribution, but the Hazen

Camp record makes it Holarctic. Judging frord the known distribution,

it is likely to be found across the high Nearctic.

Minyriolus pampia Chamberlin, 1948, p. 539 f Figs . 47-50 )

M. pampia: Oliver 1963 p. 176.

Notes on taxonomy _ Previous to the collections made at Hazen Camp,

this species was known only from one male from Clyde RiVer, Baffin

Island. In 1963 and 1964, about 520 males and females and an unknown

number of immatures of this species were collected. I have redescribed

the male and have provided illustrations. The female is here described

and drawn for the first time.

Description - Female. Color: carapace brown, splotched and

streaked with dark brown; chelicerae yellow-brown; sternum brown;

labium brown, but rimmed with gray; coxae of pedipalpi yellow brown,

but grayat the gnathobases; legs and pedipalpi yellowish brown, flecked

with gray-black spots near the joints; opisthosoma ovate, pubescent,

gray-black with fine green streaks and four small reddish spots on the

dorsum; spinnerets brown.

Structure: carapace rounded, cephalic region slightly elevated;

height of clypeus about 3. 5 diameters of an anterior median eye; eyes

small; posterior row slightly procurved; posterior medians about 2.0

diameter s of one apart, and about 1.5 diameters of one from the posterior

laterals; anterior row almost straight; anterior medians about half of

an anterior lateral in size.

Anterior medians about one diameter apart, and each about 1. 7

diameters from the anterior laterals; median ocular quadrangle longer

than wide and wider behind than in front; chelicerae reclined; sternum

only slightly longer than wide, and separating the hind coxae by almost

the length of one.

Total body length 1.85 ±0.15 mm; carapace length 0.62 ± 0.03

mm; car apace width 0. 59 ± 0. 02 mm; legs moderately short, metatarsi

slightly longer than tarsi; pedipalp tarsus lacking a spine or claw at the

tip; metatarsi each bearing one long trichobothrium at 0.74 or 0.75 ;

tibiae I — III with two spines, tibiae IV with one spine.

The color and structure of the male are like those of the female.

190

Spiders of Hazen Camp

except for the following points. Total length 1. 59 ± 0. 11 mm; carapace

length 0.63 ±0.01 mm; car apace width 0. 6 1 ±0.02 mm; tibiae I-IIIwith

two spines, tibiae IV with one spine; Tm IV at 0.78.

Natural history - This species is a member of the humid arctic faunal

element. It was found only on dens ely- vegetated slopes which are per-

manently water - saturated, and which are south- and southwest-facing.

It is further restricted to the night shadow area.

Figure 24 shows the main period of activity of the males of this

species. I am not able to explain the higher peak of the females which

coincides with the peak of the males. Adults apparently overwinter as

they were found in the slush ice at spring melt. No parasites or predators

of this species were observed.

Material examined _ About 581 adults of this species were examined

from Hazen Camp. The holotype was not seen.

Distribution (fig. 25)- This species is known only from Hazen Camp,

Ellesmere Island, and R iver Clyde, N. E. Baffin Island, N. W. T. , Canada

(7 0°N, 7 0°W) .

Savignya barbata (Koch), 1879, p. 60 (Figs. 58-61)

Erigone barbata Koch 1879 p. 60 . Savignia barbata.-'R oewer 1942 p. 623.

Typhochraestus barbatus: Bonnet 1959 p. 4745.

Notes on taxonomy _ The spelling of the generic name should be

" Savignya ", not "Savignia". The genus was named after Jules Cesar Savigny,

a French biologist of the early 19th Century, by Blackwall {1833).

Description - Female. Color: carapace brown with dark brown

markings; chelicerae pale yellow with a brownish tint; sternum dark

brown; legs pale yellow-brown with small brown splotches ; opisthosoma

gray-black with four small pale gray to reddish spots on the dorsum;

spinnerets brown; coxae brown, gnathobases gray; labium brown with

gray trim.

Structure: size small, about 1. 80 mm long; carapace broad and

rounded, slightly longer than wide, 0. 62 mm long x 0. 57 mm wide;

carapace raised behind the cephalic region, and sloping down to the eyes;

one anterior median eye about 0. 5 diameters of a lateral; anterior row

in a straight line; anterior medians about two diameters of one from the

laterals; posterior row slightly procurved; posterior medians slightly

more than two diameters of one apart, and about two diameters of one

from the laterals; posterior eyes equal or subequal in size; median

ocular quadrangle wider posteriorly than anteriorly; posterior medians

about as far apart as the quadrangle is long.

Chelicerae reclined; sternum wider than long, proportions are

2.1: 1; legs moderately short; tibia I - II with two spines, tibia III-IV

with one spine; Tm I about 0. 52; Tm II about 0. 46; Tm III about 0. 42;

Tm IV lacking.

The male is like the female in color and structure except for the

Leech

191

following features. Size small, about 1.68 mm long; carapace rounded,

0. 62 mm long x 0. 62 mm wide; carapace raised into a cephalic lobe;

cephalic pits opening out to horizontal grooves that run posteriorly the

full length of the cephalic lobe.

Fig. 24. Frequency distribution of

Minyriolus pampia , summer, 1964.

moles

8 12 16 20 24 28 2 6 10 14 18 22 26 30 3 7 II 15 19

June July August , 1964

Fig. 25. Distribution map of Minyriolus pampia.

192

Spiders of Hazen Camp

Eyes small; posterior row decidedly procurved; poster ior medians

at the top front edge of the cephalic lobe and almost five diameters of

one from the laterals; laterals about two diameters of a median in size;

anterior and posterior laterals on a small, common tubercle.

Clypeus height about ten diameters of an anterior median eye;

clypeus pubescent with short, stiff, straight, pale-colored hairs; tibia

I- II with two spines; tibia III-IV with one spine; Tm I about 0. 58; Tm

II about 0. 54; Tm III about 0. 50; Tm IV lacking.

Natural history - This species is a member of the humid arctic faunal

element. It was found only in the gravelly sections of river deltas with

scattered surface vegetation. The webs are built in the cracks in the

ground and rarely on the surface. It appears to overwinter on or near

the surface under rocks and in vegetation.

Figure 26 indicates the most active period of the summer season

for the males. It is not known if the adults overwinter.

Material examined - About 100 adults of this species were examined

from Hazen Camp, one female from Thule, Greenland (personal collec-

tion), two females from Bailey Point, Melville Island (J.E.H. Martin,

collector, 1965), five males and five females from Weatherhall Bay,

female from Axel-Heiberg Island (Heinz Rutz, collector, 1963).

Distribution (fig. 27) - Siberia (exact locality I could not find).

Novaya Zemlya. Spitsbergen. Greenland (Etah, and Thule). Ellesmere

Island (Hazen Camp) . Melville Island. Axel-Heiberg Island. This species

is Holarctic in distribution, but it is known only from the high arctic.

Typhochraestus latithorax (Strand), 1905 (Figs. 54-57)

Tarsiphantes latithorax Strand 1905 p. 2 3; T> latithorax: Bonnet 1959

p. 4262; Typhochraestus latithorax'. Holm 1960b p. 511.

Notes on taxonomy- In 1905, Strand erected the new genus Tarsiphantes ,

with the one species, latithorax . The species was described from one

damaged female and one subadult female. Holm (I960) synonymized

Tarsiphantes Strand, 1905, as a junior synonym of Typhochraestus Simon,

1884, based on a study of the holotype of latithorax. The holotype had in

the meantime become dried and even more damaged than when Strand

described it.

The genus Typhochraestus is determined and defined by the character s

of the palpus of the male, which has a large, spiral embolus with a small,

somewhat spiral basal apophysis ( see Holm 1943, and Wiehle I960). The

males of latithorax } here described and figured for the first time, have

these features .

Strand reports (1905, p. 23) that "Diese neue Gattung, deren Type

und einzigeArt die neue T. latithorax Strand ist, . . . . wurdeam Rice Strait,

30/6 1898 entdeckt ....". However, the ship "Fram" did not reach Rice

Strait until at least August 17, 1898, so either the year or the month is

in error. During June 1899, Dr. Johan Svenden, the "Fram's" doctor,

did some collecting at Fort Juliana (7 9°03 'N, 77°43'W). About August 12,

Leech

193

1899, the "Fram" left the Rice Strait for Payer Harbour, and later Jones

Sound (Bryce 1910, p. 245). Thus, it is more likely that Fort Juliana

is the actual collecting site for this species. Rice Strait (78°34'N, 74

45'W) is close to Fort Juliana.

8 12 16 20 24 28 2 6 10 14 18 22 26 30 3 7 II 15 19

June July August, 1964

Fig. 27. Distribution of Savignya barbata

194

Spiders of Hazen Camp

Description - Female. Color: carapace brown, marked and shaded

with dark brown; chelicerae brown; sternum brown; labium brown, but

marked with gray; coxae of pedipalpi brown, but gnathobases gray; legs

pale brown-yellow, distal part of each leg segment brownish; opisthosoma

brown-gray; spinnerets brown-gray.

Structure: size small, about 1. 90 mm total length; carapace longer

than wide, 0. 64 mm long x 0. 54 mm wide, carapace broad and rounded,

gradually rising to the ,low cephalic region; clypeus height about five

diameters of an anterior median eye; posterior row of eyes slightly

procurved; posterior medians about 1.5 diameters of one apart, and

closer to the posterior laterals than to each other; anterior row of eyes

slightly procurved; anterior medians les s than the diameter of one apart,

and about the diameter of one from the laterals; anterior laterals almost

twice the size of an anterior median; median ocular quadrangle about as

wide at posterior medians as long; chelicerae decidedly r eclined; ster-

num about as wide as long; legs moderately long; metatarsus IV about

1.45 times longer than tarsus IV; tibia I - III with two spines, tibia IV

with one spine at 0.29; Tm I about 0.63; Tm II about 0.54; Tm III about

0.48; Tm IV lacking.

Male. The male is colored like the female. Structure: size

small, about 1.45 mm; carapace longer than wide, about 0. 64 mm long

x 0. 54 mm wide; carapace structure like that of the female, except for

a small, post-ocular sulcus and lobe that is finely-bridged and connecting

behind each posterior median eye (see figures 55, 56); metatarsus IV

about 1.29 times longer than tarsus IV; tibia I-III with two spines, tibia

IV with one spine; Tm I about 0. 56; Tm II about 0. 53; Tm III about 0. 50;

Tm IV lacking. F or the characteristic details of the pedipalp of the male,

see figure 54.

Natural history - This species is a member of the humid arctic faunal

element. The species is widely distributed throughout soggy, vegetated

areas at and near pond edges, but is restricted to the night shadow areas .

Individuals have been collected under water in slush snow at the time of

the spring melt, and on wet, south-facing slopes and depr es sions aboun-

ding with sedges or mosses.

Figure 28 shows the abundance pattern of the species during the

summer of 1964. It is not known if the adults overwinter, but it can be

assumed that they do as the adults are found so early in the season.

Material examined _ About 100 adult individuals of this species were

examined from Hazen Camp and four males and two females from Axel-

Heiberg Island (Heinz Rutz, collector, 1963). The holotype was not seen.

Distribution (fig. 29) - This species is known from three localities

only, two on Ellesmere Island at Hazen Camp and either Rice Strait (78

34'N, 74°45 1 W) or Fort Juliana (79°03'N, 77°43'W), and Axel-Heiberg

Island.

Xysticus deichmanni Soerensen, 1898 p228 (Figs. 40, 41)

X. labradorensis: Bonnet 1959 p. 4883 (in part) ; deichmanni: Holm

numbers of individuals

Leech

195

1958b p. 533, Buckle and R edner 1964 p. 1139, Oliver 1963 p. 176,

Turnbull, Dondale, and R edner 1965 p. 1263.

12

10

8

6

4

2

0

— • — males

— a — females

n = 95

Fig. 28. Frequency distribution of

Typhochraestus latithorax , summer, 1961.

A

l\

*

8 12 16 20 24 28 2 6 10 14 18 22 26 30 3 7 II 15 19

June July August , 1964

Fig. 29. Distribution of Typhochraestus latithorax .

196

Spiders of Hazen Camp

Notes on taxonomy - Holm (1958b) and Buckle and R edner (1964) have

distinguished the species labradorensis from deichmanni . The distribution

records of the two species do not overlap.

Natural history - This species is a member of the arid arctic faunal

element. Individuals at Hazen Camp were found with Dictyna borealis under

and around Dryas integrifolia . The main differ ence observed about the habitat

of the two species is that X. deichmanni remains mostly on the ground under

and beside vegetation (occasionally in the Dryas flower s) , whereas/), borealis

is mostly on and in the vegetation. X. deichmanni was found mainly under and

near Dryas integrifolia clumps, but occasionally near Salix arctica and Kobresia

myosuroides .

Figure 30 shows the occur r ence of the adults of this species during

the summer of 1964. Data from 1963 are almost identical. The adults

are able to overwinter, as is indicated by the late season increase.

The Thomisidae have little or no courtship preceding mating (Kaston

1936), and Xysticus deichmanni is no exception. In all four cases observed,

the males mounted the females after a short contact or mounted directly

upon contact without any hesitation. The females offered no resistance

to any of the males. Once upon the females, the males tied down the

females with silk. Silk threads were attached from the carapace to the

patellae to the ground and back again many times. Once the female was

thoroughly tied down, the male began to mate.

Before the actual mating, the male appeared to clean and polish the

palpi in the chelicerae. Each palpus was very car efully rubbed and mani-

pulated. When this was done, the male crawled back along the female,

then around and under the posterior end of the opisthosoma of the female,

then mated by alternately placing the palpi upon the epigynum. When

mating, the pair are positioned venter to venter and facing the same

direction (see Kaston 1936). The duration of the matings were 5, 52,

55. 5 and 59 min each. In the case lasting five min, the male was suc-

cessful in placing each embolus within the epigynum once before leaving

the female. In the remaining cases, each palpus was placed on the epigy-

num for an average of about 14 min. The haematodocha was refilled and

embolus re-inserted once every 20 sec. When the haematodocha refilled,

the large dorsal and some smaller lateral spines on the legs of the male

became erect for about two sec, then gradually over the next three sec,

relaxed.

When the male was finished mating, he again crawled onto the dor-

sum of the female, polished the palpi in the chelicerae, paused, then

fled rapidly. The males died about four days later.

As of October 8, 1964, no eggs were laid. Thus it can be assumed

that the fertilized females overwinter and lay eggs in the following sum-

mer. Gertsch (1964, pers. comm.) has confirmed this. Sometime

between October 8 and 18, 1964, eggs were laid in a small sac on the

flat edge of a rock. There were 28 eggs. The sac was lenticular, about

six mm in diameter, and about two mm in thickness in the centre.

The main food for the species seems to be small Diptera, especially

Chironomidae and Culicidae. Oliver (1963, pers. comm.) found several

instances where this species was feeding on the first instar larvae of

numbers of individuals

Leech

197

Gynaephora rossi Curtis and G. groenlandica (Horn.) (Lymantr iidae, Lepidoptera)

as they emerged from the eggs on the cocoon of the female. The first

instar larvae are only about one mm long, and present no difficulty in

grasping for the spider. Oliver (pers. comm.) and I have also found

specimens hiding in Dryas flowers, presumably to catch visiting insects .

8 12 16 20 24 28 2 6 10 14 18 22 26 30 3 7 II 15 19 23

June July August, 1964

Fig. 31. Distribution map of Xysticus deichmanni .

198

Spiders of Hazen Camp

In 1963, one female was found with a parasite, Hexamermis species

(Nematoda, Mermithidae), inside the opisthosoma. The epigynum was

abnormal, indicating parasitic castration within.

The contents of the crop and gizzard of two snow buntings showed

remains of legs and carapaces of X. deichmanni . There were also many

observed cases of cannibalism.

Material examined - About 575 adults of this species were examined

from Hazen Camp, and one male and one female from Chesterfield Inlet,

N. W. T. , and two females from Tanquary Fjord, Ellesmere Island (Guy

Brassard, collector, 1964).

Distribution (fig. 3 1) - Gr eenland (N . E . Greenland between 7 0-78°N) .

Canada (Franklin District: near Ukpilik Lake, King's Bay, and Holman

Island, Victoria Island; Hazen Camp and Tanquary Fjord, Ellesmere

Island; Moose Bay, Bathur st I sland; Lake Harbour, Baffin Island; mouth

of the Aktinek River , Bylotlsland, 70°N, 78°W; Keewatin District; N. W.

side Aberdeen Lake; Chesterfield Inlet; Mackenzie District: Bathurst

Inlet; Coppermine; Bernard Harbour; Yukon Territory: Firth River,

16 miles from the Arctic coast; Swede Dome, 34 miles W. Dawson City).

Alaska (Mile 206, Richardson Highway; Nome; Point Barrow; Umiat;

Meade River; and Cooper Landing).

Notes: Buckle and R edner (1964, p. 1141) record the Richardson

Highway as being in the Yukon. This locality is really in Alaska, 206

miles north of Anchorage. The Holman Island locality referred to by

these authors is most likely the townsite of Holman Island on Victoria

Island, not the very small island off shore near the town. King's Bay,

Victoria Island, for all intents and purposes, is the same locality as

Holman Island*

ZOOGEOGRAPHICAL CONSIDERATIONS

In 1934, Gelting, after an analysis of the botanical and geological

evidence, proposed that the northeastern tip of Peary Land was ice free

during the maxima of the ice ages, and that some other areas on Green-

land were ice free during at least the Wisconsin Glaciation. The idea of

ice free areas or "refugia" in Greenland was not his though, as Kornerup

(1878) proposed this after a geological study, and Warming (1888) again,

after a botanical study. Heated controversy for and against the theory of

glacial refugia has occurred since then and the subject is still being

debated (Ball 1963, Lindroth 1963a, Benson 1958, Nordhagen 1963).

In 1937, Eric Hulten published a monumental work on phytogeo-

graphy in the arctic and boreal regions based on the premise of glacial

refugia. Since then, most biologists agr ee that ice-bound refugia existed

during glacial times (Savile 1961; Packer 1962, various author s in Lbve

and Love 1963, various authors in Lowther 1959, 1962, Lindroth 1957,

1961, 1963a, 1963 b, Larsson 1959, Harington 1964, Hammer 1955,

Bocher et al. 1957, Ball 1963, McPhail 1961).

The basic premise upon which biologists have based theories of

Leech

199

glacial refugia (other than on geological evidence) is a more or less

limited distribution of a species or of many species. The locality or

area of the distribution is also held as significant. The greater the num-

ber of species in an area that have not (yet) been found in other areas,

the greater the possibility that the area was a refugium. But the number

of known species from an area and the distribution of the species concerned

is most often directly proportional to the thor oughnes s of collecting that

has been done.

No area in Canada is as thoroughly collected as the Hazen Camp

area. This fact alone would ordinarily bias distribution patterns beyond

use for zoogeographic purposes unless some criterion other than tax-

onomy is used.

I am therefore introducing data from insects and spiders, based

on morphology, vagility, and biology, and some geological evidence, to

support a suggestion that the northern part of Ellesmere Island had ice

free areas during the Wisconsin Glacial division (and perhaps more divi-

sions within the Pleistocene epoch) that served as glacial refugia.

Studies at Hazen Camp have uncovered about 350 species of insects,

arachnids and Collembola, including several new species of Homoptera

(Richards 1963, 1964a, b), Diptera (Oliver 1963, and pers. comm.),

Coleoptera (W. J. Brown, unpubl. and J.A. Downes 1964), Hymenoptera

(W. R. M. Mason, pers. comm.), and Acarina (E. Lindquist, pers.

comm.). Except for two species of Ichneumonidae thatareover fivemm

long that are scarce, the bulk of the new material from Hazen Camp is

small (less than four mm long) and of generally cryptic, unstudied and

poorly collected groups.

There are about 75 species of flightless insects in the Hazen Camp

area. Most of them are brachypterous and some are apterous. Flight-

lessness in insects is not a rapidly developed feature, especially in peri-

pheral or marginal regions, where species cannot get the extra energy

necessary for morphological experimenting, and where emphasis is on

feeding and breeding. I therefore suggest that these flightless insects

have been on northern Ellesmere Island for part, if not all, of the Pleis-

tocene epoch. Gressitt (1964, p. 595), in contrary opinion, states that

selection favouring loss of wings in insects, particularly Diptera on

Campbell Is. ,(N. Z.), is probably proceeding at a rapid rate. At Hazen

Camp, however, no apterous Diptera were found, so either Gressitt's

theory is wrong or else the flightless condition develops in insects at

different rates in different areas (my inference).

To date, there are 14 species of Collembola and about 80 species

of Acarina known from the Hazen Camp area. However, I do not believe

that these two groups can be used for refugium analysis as they have the

ability to colonize readily in areas where no other arthropod can and they

do so very rapidly; the method of this rapid dispersal may be by wind

(Gressitt et al. 1963, and Gres sitt and Collaborator s 1964) and by indi-

viduals and/or eggs on clods of dirt on birds* feet. Thus there is no

way of calculating when these two groups came into an area.

Several species of spiders have probably remained on northern

Ellesmere Island during most or all of the Pleistocene epoch. Tarentula

exasperans was never observed to have a drag line, a feature that might

200

Spiders of Hazen Camp

be analogous to flightlessness in insects (H. W. Levi, pers. comm.), and

Pardosa glacialis and Xysticus deichmanni have drag lines that are so weak that

they would not support the weight of even a third instar, a feature that

might be analogous to brachyptery in insects (my inference).

In contrast, several immatures of Dictyna borealis were observed

ballooning in early July, 1964. Braendegaard (1937, 1938) has made

similar observations in Greenland and elsewhere. The remaining species

of spiders, all Linyphiidae, have strong drag lines, indicating possibly

recent immigration to the Hazen Camp area.

It appears that there are two basic zoogeographical groups of

spiders at Hazen Camp: one group of three species that has withstood

the Wisconsin Glaciation and probably most or all of the Pleistocene

epoch on northern Ellesmere Island, and a second group that may have

moved into the area recently.

The second group, that is, the recent immigrants, appears to have

had two sources, one from the arctic zone and the other from the boreal

or lowarctic zone. Two species of Linyphiidae, Typhochraestus latithorax and

Minyriolus pampia are confined to the night shadow area, and are thus

possibly not yet adapted to the arctic light conditions . These night shadow

areas are sunny during the day, but shaded during the period that would

be night in the temperate regions. The shadow regions are often cooler

"at night" than the sunny r egions, hence these two species appear to have

a diurnal rhythm. On this basis I suggest that these two species are

recent immigrants from the temperate or low arctic regions. The re-

maining species of Linyphiidae are not confined to the night shadow

regions. They display full adaptation to the arctic conditions of light.

Their general Holarctic distribution indicates this as well.

Taxonomic evidence that the nor them end of Ellesmere Islandmay

have been a refugium is based on an analysis of muskoxen skulls by

Harington (1964). Gjaerevoll (1963) states that there was a refugium

somewhere in the Queen Elizabeth Islands, though he gives neither

reasons nor references.

The geological evidence of a refugium on Ellesmere Island is

divided. Hatter sley-Smith (196l) suggests that any ice that might have

been on the plateau between Lake Hazen and Alert was protective rather

than erosive, as it is unlikely that the soft silts and lignite would have been

preserved in an area where they are in partcovered byapiedmont glacier

at the present time. On the other hand, widespread erratics have been

found in this area, though the date of deposition is not known.

I do not believe that during the Ice Ages of the Pleistocene epoch

conditions were ever as severe as most are led to believe. It is fully

possible that the conditions were so poor for several years in succession

that the ice and snow did not melt off the ground, but equally possible

that one season in four or five, or even ten, with favourable conditions

melted the ice and snow and permitted life and growth to continue. Thus,

even though there were about one and one-half million year s in the Pleis-

tocene epoch (Ericson etaL 1964), the effective time available for arthro-

pods to have been active may have been as little as three hundred thousand

years. If the Pleistocene epoch is shortened to three hundred thousand

years as some authors believe it should be, then these animals have had

Leech

201

even less time to evolve the flightless condition.

In summary, it appears that there is one fauna on northern Elles-

mere Island that has been there since before the Wisconsin Glaciation

and perhaps for the duration of the Pleistocene epoch, and another fauna

that may have immigrated to northern Ellesmere Island in post- Wisconsin

times.

ACKNOWLEDGEMENTS

My special thanks are to Dr. George E. Ball, chairman of my

committee, for help and guidance during this study. Appreciation is

extended to Dr. W. G. Evans, Entomology Department, University of

Alberta, for discussions during the progress of writing the thesis.

My special thanks are also extended to the following and their

respective institutions for the use of material and examination of speci-

mens while I visited their institutions and for material loaned to me by

them: Drs. C. D. Pondale and A. L. Turnbull (Agricultural Research

Institute, Belleville, Ontario); Dr. Willis J. Gertsch (American Museum

of Natural History, New York); Dr. Herbert W. Levi (Museum of Com-

parative Zoology, Harvard University); and Dr. Glenn B. Wiggins (Royal

Ontario Museum).

I thank Dr. Jens Braendegaard of Copenhagen, Denmark, for re-

prints, the loan of rare material and for helpful notes on synonymies;

Mr. Wilton Ivie for determinations of difficult and obscure spider species;

Dr. Ake Holm for reprints, identification of difficult species, and for

very lengthy correspondence during the course of this work; Drs. B. J.

Kaston, F. Papi, and Pierre Bonnet for reprints and correspondence;

and to fellow students Soenartono Adisoemarto and Donald Whitehead for

lengthy helpful discussions.

For identifications of the parasitic worms, I am indebted to Dr.

Harold Welch then of the Belleville, Ontario, Laboratories.

I express thanks to the National Research Council of Canada for

partial support in this project through a grant held by Dr. George E.

Ball.

Thanks are also extended to Drs. G. P. Holland and D.R. Oliver

of the Entomology Research Institute in Ottawa for allowing me to start

and complete this project at Lake Hazen; to Dr. G. Hatter sley-Smith

and the Defence Research Board of Canada for the use of Hazen Camp

and its supplies; to pilots and crewmen of the R.C. A.F. and Atlas Air

Lines for flying me and my equipment to and from Lake Hazen; to the

men of Alert and Eureka Weather Stations for accommodation and infor-

mation about the areas during the summers of 1963 and 1964.

I thank Larry Law (Dominion Observatory, Ottawa), Leonard Hills

(Geology Department, University of Alberta), and Guy Brassard (Queens

University) for making collections in areas that I could not reach.

And I thank "Little Mike", the radio operator at Eureka Weather

Base and Charles Harris, Edmonton Radio Ham Operator, for passing

messages to and from Hazen Camp.

202

Spiders of Hazen Camp

REFERENCES

Ball, G. E. 1963. The distribution of the species of the genus Cryobius

(Coleoptera, Carabidae, Pterostichus) : with species reference to

the Bering and ridge and Pleistocene refugia. In Gressitt, J. L.

(ed. ) Pacific basin biogeography, a symposium. Tenth Pacific

Science Congress. Honolulu, Hawaii. Bishop Museum Press,

pp. 133-151, 9 figs.

Banks, N. 1900. Arachnida of the expedition. In papers from the Har-

riman Alaska expedition. 11. Entomological results: 5, Arach-

nida. Proc. Wash. Acad. Sci. , 2 : 477-486, pi. 29.

Benson, R.B. 1958. Arctic sawflies and open habitat. Proc. int. Congr.

Ent. X. Montreal 1 : 693.

Bishop, S.C., and C.R. Crosby. 1935. Studies in American spiders:

Miscellaneous genera of Erigoneae. Part I. J1N.Y. ent. Soc.,

43:217-241; 255-281; 1 text fig. , pi. 5, 84 figs.

Bishop, S.C., and C.R. Crosby. 1938. Studies in American spiders:

miscellaneous genera of Erigoneae. Part II. J.N.Y. ent. Soc.,

46 : 55-107, pi. 6, 86 figs.

Blackwall, J. 1833. Character s of some undescribed genera and species

of Araneidae. Lond. Phil. Mag. Sci., 3(3) : 104-112; 187-197;

344-352; 436-443.

Bocher, T. W. , K. Holmen, and K. Jakobsen. 1957. Gronlands Flora.

P. Haase & Sons, Copenhagen.

Bonnet, P. 1945-61. Bibliogr aphia Araneorum. 3 tomes, 7 vols. Les

Freres Douladoure, Toulouse.

Braendegaard, J. 1936. R evisal of spider s from Ellesmer eland. Norsk

ent. Tidsskr. 4 : 128-130.

Braendegaard, J. 1937. Observations on spiders starting off on bal-

looning excursions . Vidensk. Meddl. Dansk naturh. Foren. 101:

115-117.

Braendegaard, J. 1938. Aeronautic spiders in the Arctic. Meddl. om

Gr^nland, 119, no. 5 : 1-9, 3 figs.

Braendegaard, J. 1940. 1. Spiders (Araneina) from northeast Green-

land between lats. 70°25> and 76°50'N. 1-23, 21 figs. 2. Onthe

possibility of a reliable determination of species of the females of

the genus Erigone . 24-31, 5 figs. Meddl. om Gr^nland, 125, no. 8.

Braendegaard, J. 1946. The spiders (Araneina) of east Greenland, a

faunistic and zoogeographical investigation. Meddl. om Gr^fnland,

121, no. 15 : 1-128, 49 figs.

Braendegaard, J. 1958. Araneida. In Fridriksson, A., and S. L.

Tuxen (managing editor s) . The zoology of Iceland. 3 (prt. 54):

1-113, 88 figs.

Braendegaard, J. I960. The spiders (Araneida) of Peary Land, north

Greenland. 159, no. 6 : 1-24, 5 figs.

Bristowe, W.S. 1958. The world of spiders. The New Naturalist,

Collins, St. James's Place, London. 304 p. , 36 pi.

Bryce, G, 1910. The siege and conquest of the North Pole. Gibbings

and Co. , Ltd. , London. 334 p.

Buckle, D. J. , and J.H. Redner. 1964. The Nearctic species of the

Leech

203

Xysticus labradorensis subgroup (Araneae : Thomisidae). Can. Ent. 96:

1138-1152, 13 figs.

Chamberlin, R. V. 1948. On some American spiders of the family

Erigonidae. Ann. ent. Soc. Amer. 41 : 483-562, 153 figs.

Chamberlin, R.V., and W. Ivie. 1947. The spiders of Alaska. Bull.

Univ. Utah, 37(10) : 1-103, 99 figs.

Chamberlin, R . V. , and W. J. Gertsch. 1958. The spider family Dicty-

nidae in America- north of Mexico. Bull. Am. Mus. Nat. Hist.,

116 (art. 1) : 1-152, 38 maps, 47 pi., 603 figs.

Christie, R.L. 1962. Nor theastern Ellesmere I sland, District of Fran-

klin. Geological Survey of Canada. Paper 62-10, 15 p. (map).

Crompton, J. 1950. Life of the spider. Illustrated Fontana Books.

252 p. , 24 figs.

Crosby, C.R., and S. C. Bishop. 1931. Studies in American spiders:

genera Cornicularia , Paracornicularia , Tigellinus, Walkenaera , Epiceraticelus ,

and Pelecopsis , with descriptions of new genera and species. J1 N.

Y. ent. Soc. 39 : 359-403, 6 pi. , 97 figs.

Crosby, C.R., and S. C. Bishop. 1933. American spiders: Erigoneae,

males with cephalic pits. Ann. ent. Soc. Amer. 24 : 105-182, 10

pi. , 236 figs.

Crosby, C.R., and S. C. Bishop. 1936. Studies in American spiders:

misc. genera of Erigonidae. In E. Strand (ed. ) Festschrift Zum

60 Geburtstag von Embrik Strand 2 : 52-64, pi. 4 &; 5.

Dahl, M. 1928. Spinnen (Araneae) von Nowaja Semlja. Norsk. Vid.

Akad. , Oslo. 37 p.

Dahl, M. 1933. Spinnen (Araneae) . In H.U. Sverdrup (ed. ) . TheNor-

wegian North Polar exped. with the "Maud", 1918-1925. Scien-

tific Results 5(162) : 3-4.

Day, J. H. 1964. Characteristics of soils of the Hazen Camp area,

northern Ellesmere Island, N. W. T. Defence Research Board of

Canada. D. Phys. R. (G) Hazen 24, 15 p. , 25 figs.

Dondale, C. D. 1961. Life histories of some common spiders from

trees and shrubs in Nova Scotia. Can. J. Zool. 39 : 777-787, 2

figs.

Dondale, C.D. 1964. Sexual behaviour and its application to a species

problem in the spider genus Philodromus (Araneae : Thomisidae).

Can. J. Zool. 42 : 817-827, 11 figs.

Downes, J.A. 1964. Arctic insects and their environment. Can. Ent.

96 : 279-301, 2 pi.

Ericson, D. B. , M. Ewing, and G. Wollin. 1964. The Pleistocene epoch

in deep sea sediments. Science 146 : 723-732, 4 figs.

Flint, R.E. 1957. Glacial and Pleistocene geology. Publishers John

Wiley and Sons, Inc. , New York, 553 p. , 127 figs.

Gelting, P. 1934. Studies on the vascular plants of east Greenland be-

tween Franz Joseph Fjord and Dove Bay (Lat. 73° 15,-75°20,N)

Meddl. om Grjefnland 101 : 1-340, 4 pi., 47 figs.

Gertsch, W. J. 1934. Notes on American Lycosidae. Am. Mus. Novit.

693 : 1-25.

Gertsch, W. J. 1939. A review of the typical crabspiders (Misumeninae)

of America north of Mexico. Bull. Am. Mus. Nat. Hist. 76(7): 277-

204

Spiders of Hazen Camp

422, 271 figs.

Gertsch, W. J. 1949. American spiders. D. Van No strand Co., Inc.,

New York. 285 p. , 32 pi. , 6 figs.

Gertsch, W. J. 1953. The spider genera Xysticus , Coriarachne , and Oxyoptila

(Thomisidae, Misumeninae) in North America. Bull. Am. Mus.

Nat. Hist. 102(4) : 417-482, 93 figs.

Gjaerevoll, O. 1963. Survival of plants on nunataks in Norway during

the Pleistocene glaciation. In A. & D. L'ove (ed. ) North Atlantic

Biota and Their History, pp. 261-283, 22 figs.

Gressitt, J. L. , R.E. Leech and K.A.J. Wise. 1963. Entomological

investigations in Antarctica. Pacific Insects 5(1) : 287-304, 9 figs.

Gressitt, J. L. , and collaborators. 1964. Insects of Campbell Island.

Pacific Insects, Monograph 7 : 1-663.

Hackman, W. 1957. Studies on the ecology of the wolf spider, Trochosa

rurica Dej. Soc. Scient. Fennica 16(6) : 1-34, 8 figs.

Hammer, M. 1955. Some aspects of the distribution of microfauna in

the Arctic. Arctic 8 : 115-126, 12 figs.

Harington, C.R. I960. A short report of snow and ice conditions on

Lake Hazen, winter 1957-58. Defence Res. Board of Canada. D.

Phys. R. (G) Hazen 7, 64 p.

Harington, C.R. 1964. Remarks on Devon Island muskoxen. Can. J.

Zool. 42 : 79-86, 5 figs.

Hatter sley-Smith, G. 1961. The ice cover of northern Ellesmere Island.

Ann. N.Y. Acad. Sci. 95 : 282-289.

Hatter sley-Smith, G. 1964. Bibliography of "Operation Hazen", 1957-

63. Defence Res. Board of Canada, D. Phys. R. (G), Hazen 20 :

1-5.

Holm, A. 1937a. Zur Kenntnis der Spinnenfauna Spitsbergen und der

baren Insel. Ark. Zool. 29a (18) : 1-13, 4 figs.

Holm, A. 1937 b. Notes on the spider fauna of East Greenland. Ark.

Zool. 29b(12) : 1-12.

Holm, A. 1939. Insecta and Araneae collected in the Kanger dlugsuak

region of East Greenland by the British expedition, 1935-36. Ann.

Mag. nat. Hist. ser. 11, 3 : 60-80, 4 figs.

Holm, A. 1941. Uber Gynandromorphism und Inter sexualitat bei den

Spinnen. Festskrift Tillagnad Professor Sven Ekman. Zool.

Bidrag, Uppsala 20 : 397-413, 3 figs.

Holm, A. 1943. Zur Kenntnis der Taxonomie, Okologie und Verbr eitung

der Schwedischen Arten der Spinnengattung Khaebothofax Sim.,

Typhochraestus Sim., und Latithorax n. gen. Ark. Zool. 34a (19): 1-32,

9 text figs, 3 pi. , 25 figs.

Holm, A. 1944. Revision einiger Norwegischen Spinnenar ten und Bemer-

Kungen liber deren Vorkommen in Schweden. Ent. Tidskr. 65:

122-134, 6 figs.

Holm, A. 1945. Zur Kenntnis der Spinnenfauna des Tornetraskgebietes .

Ark. Zool. 36a(15) : 1-80, 26 figs.

Holm, A. 1950. Studier eniiber die Spinnenfauna des Tornetraskgebietes .

Zool. Bidrag, Uppsala 29 : 103-213, 26 figs.

Holm, A. 1951. Araneae. In Per Brinck and K. G. Wingstrand (ed. )

The mountain fauna of the Virihaur e ar ea on Swedish Lapland. II.

Leech

Z05

O

Special account. Lunds Universitets Arsskrift 46(2) : 138-149, 8

figs.

Holm, A. 1956. Notes on arctic spider s of the genera Erigone Aud. , and

Hilaira Sim. Ark. Zool. 9(20) : 453-469, 20 figs.

O

Holm, A. 1958a. The spiders of the Isfjord region of Spitsbergen.

Zool. Bidrag, Uppsala 33 : 29-69, 20 figs.

Holm, A. 1958b. Spiders (Araneae) from Greenland. Ark. Zool. serie

2, 11(31) : 525-534, 10 figs.

Holm, A. 1958c. Spindlar fran Harjedalens fjaltrakter. Ent. Tidskr.

79(3-4) : 216-230.

o

Holm, A. 1960a. On a collection of spider s from Alaska. Zool. Bidrag,

Uppsala 33 : 109-138, 42 figs.

Holm, A. 1960b. Notes on Arctic spider s . Ark. Zool. serie 2, 12(2) :

511-524, 3 figs.

Hulten, E. 1937. Outline of Arctic and bor eal biota during the Quaternary

Period. Bokjorlags Aktiekolaget Thule, Stockholm. 168 pp. , 14

figs. , 43 pi.

Jackson, A. R. 1930. Results of the Oxford University expedition to

Greenland, 1928. - Araneae and Opiliones collected by Major R . W.

G. Hingston; with some notes on Icelandic spiders. Ann. Mag.

nat. Hist., ser. 10, 6 : 639-656, 1 pi., 14 figs.

Jackson, A.R. 1934. Notes on Arctic spiders obtained in 1933. Ann.

Mag. nat. Hist. , ser. 10, 14 : 611-620.

Jackson, A.R. 1938. Notes on Arctic spiders obtained in 1933-1936.

Proc. zool. Soc. Lond. 1937, ser. B, 107 : 543-551.

Jackson, C.I. 1959, The meteorology of Lake Hazen, N. W. T. Part

I, Analysis of the observations. Defence Res. Board of Canada,

D. Phys. R. (G), Hazen 8 : 1-194.

Jackson, C.I. I960. The meteorology of Lake Hazen, N. W. T. based

on observations made during the International Geophysical Year

1957-58. Parts II, III, IV. Synoptic influences, local forecasting,

bibliography. Arctic Meteorology R esearch Group, McGill Univer-

sity, Montreal. Meteorology No. 16. Defence Research Board of

Canada, Ottawa: Report D. Phys. R. (G), Hazen 9, 195-295.

Kaston, B. J. 1936. The senses involved in the courtship of some vaga-

bond spiders. Ent. Amer. 16 : 97-167, 23 figs.

Kaston, B. J. 1948. The spiders of Connecticut. Conn. State Geol.

nat. Hist. Survey. Bull. 70 : 1-874, 2144 figs.

Kaston, B. J. 1961. Spider gynandromorphs and intersexes. J1 N. Y.

ent. Soc. 69 : 177-190, 18 figs.

Kaston, B. J. 1963a. Deformities of external gerptalia in spiders. J1

N.Y. ent. Soc. 71 : 30-39, 23 figs.

Kaston, B. J. 1963b. Abnormal duplication of the epigynum and other

structural anomalies in spiders. Trans. Amer. Micr. Soc. 82 :

220-223, 10 figs.

Koch, L. 1879. Arachniden aus Siberienund Nowaja Semlja, eingesam-

melt von der Schwedischen Expedition im Jahr 1875. Kongl. Svenska

Vet. - Akad. , Handl. , 16(5) : 3-136, 8 pi.

Kornerup, 1878. Geologiske Iagttegelser fra Vestkysten af Grjdnland.

Medd. Grpnland, 1, Kobenhavn.

206

Spiders of Hazen Camp

Lar s son, S. G. 1959. Coleoptera 2, general remarks. In Fridriksson,

A., and S. L. Tuxen (managing editors) The Zoology of Iceland,

3, part 46b : 1-85, 13 figs.

Lenz, H. 1897. Groenlandischen Spinnen. Biblio. Zool. , 20(3) : 73-76.

Lindroth, C. H. 1957. The faunal connections between Europe and North

America. John Wiley & Sons Inc., Stockholm and New York. 344

p. , 61 figs.

Lindroth, C. H. 1961. Verbindungen und Barrier en in der Zirkumpolar

der Insekten. Proc. int. Congr. Ent. XI Wien, 1 : 438-445.

Lindroth, C. H. 1963a. The fauna history of Newfoundland as illustrated

by Carabid beetles. Opusc. ent. Suppl. 23 : 1-112, 96 figs.

Lindroth, C. H. 1963b. The problems of late land connections in the

North Atlantic area, 73-85 p. , 6 figs. In A. & D. Lbve (ed. )North

Atlantic Biota and Their History.

Locket, G. H. , and A. I. Millidge. 1951. British spiders. Vol. 1. Ray

Society, London. 310 p. , 142 figs.

Locket, G. H. , and A. I. Millidge. 1953. British spiders. Vol. 2. Ray

Society, London. 449 p. , 254 figs.

Love, A., and Lbve, D. (ed. ) 1963. North Atlantic biota and their

history. Pergamon Press Co. , New York. 430 p.

Lowther, G. R . , (ed. ) 1959. Problems of the Pleistocene and Arctic.

Vol. 2, 67 p. McGill Univ. Public. , Montreal.

Lowther, G. R . (ed. ) 1962. Problems of the Pleistocene and Arctic.

Vol. 2, 67 p. McGill Univ. Public. , Montreal.

McAlpine, J. F. 1965. Insects and related terrestrial invertebrates of

Ellef Ringnes Island. Arctic 18(2) : 73-103, 7 figs.

McPhail, J.D. 1961. A systematic study of the Salvelinus alpinus complex

in North America. J. Fish. Res. Bd. Canada, 18(5) : 793-816, 6

figs.

Nordhagen, R. 1963. Recent discoveries in the south Norwegian flora

and their significance for the understanding of the history of the

Scandinavian mountain flora during and after the last glaciation.

In A. & D. Love (ed. ) Atlantic biota and their history. 241-260

12 figs.

Oliver, D. R. 1963. Entomological studies in the Lake Hazen area,

Ellesmere Island, including lists of species of Arachnida, Collem-

bola, and insects. Arctic 16 : 175-180.

Oliver, D.R. and P. S. Corbet. 1966 (in press). Aquatic habitats in a

high Arctic locality : Hazen Camp study area, Ellesmere Island,

N. W. T. Defence Research Board, Ottawa, Canada.

Osterloh, A. 1922. Beitrage zur Kenntnis des Kopulationsapparates

einiger Spinnen. Z. wiss. Zool. 119 : 326-421, 42 figs.

Packer, J. G. 1962. The taxonomy of some North American species of

Chrysosptenium L. , section alternaria Franchet. Can. J. Bot. 40:

85-103, 11 figs.

Papi, F. 1955. Astronomische Orientierung bei der Wolfspinne Arctosa

perita (Latr.). Z. vergl. Physiol. 41 : 481-489, 4 figs.

Papi, F. , and J. Syrjamaki. 1963. The sun- orientation rhythm of wolf

spider s at differ ent latitudes . Arch. ital. Biol. 101:59-77, 5 figs.

Papi, F. , and P. Tongiorgi. 1962-63. Or ientamento astronomico verso

Leech

207

nord: una capacita innata dei ragni del gen. Arctosa . Monit. zool.

italiano 70-71 : 485-490, 1 fig.

Papi, F. , and P. Tongiorgi. 1963. Innate and learned components in the

astronomical orientation of wolf spiders. Ergeb. Biol. 26 : 259-

280, 15 figs.

Petrunkevitch, A. I. 1911 (ed. J.A. Allen). A synonymic index catalogue

of spiders of North, Central and South America, with all adjacent

islands, Greenland, Bermuda, West Indies, Tierra del Fuegq

Galapagos, etc. Bull. Am. Mus. nat. Hist. 29 : 1-793.

Pickard-Cambridge, O. 1873. On some new species of Araneida, chiefly

from oriental Siberia. Proc. zool. Soc. Lond. 435-452, pi. nos.

40, 41, 14 figs.

Pickard-Cambridge, O. 1877. On some new and little known spiders

from the arctic regions. Ann. Mag. nat. Hist, ser . 4(20): 273-285, pi. 8.

Powell, J.M. 1961. The vegetation and microclimate of the Lake Hazen

area, northern Ellesmere Island, N. W. T. Defence Res. Board of

Canada. D. Phys. R. (G), Hazen 14 : 112 p.

Richards, W. R . 1963. The Aphididae of the Canadian Arctic (Homop-

tera). Can. Ent. 95 : 449-464, 3 figs.

Richards, W.R. 1964a. A new arctic aphid (Homoptera: Aphididae).

Can. Ent. 96 : 1027-1029, 2 figs.

Richards, W.R. 1964b. The scale insects of the Canadian Arctic (Homop-

tera: Coccoidea). Can. Ent. 96 : 1457-1462, 16 figs.

Roewer, C.F. 1942-1954. Katalog der Araneae Tome I, Tome II, parts

A & B. Tome I, 1040 p. Bremen Zeitun^N.S. - Gauverlag

Weser - Ems GmbH. , Bremen, 1942. Tome II, part A, 1-923,

part B, 927-1751. Patrimoine de I'Institut royal des Sciences

naturelle de Belgique, Bruxelles, 1954.

Savile, D. B. O. 1961. The botany of the northwestern Queen Elizabeth

Islands. Can. J. Bot. 39 : 909-942, 2 figs.

Savile, D.B.O. 1962. Personal and unpublished notes.

Savile, D.B.O. 1964. General ecology and vascular plants of the Hazen

Camp area. Arctic 17 : 237-258, 2 figs.

Savory, T. H. 1928. The biology of spiders. Sidgwick and Jackson,

Ltd., London. 376 p. , 121 figs.

Schenkel, E. 1950. Spinnentiere aus dem Westlichen Nordamerika,

gesammelt von Dr. Hans Schenkel - Rudin. Verh. naturf. Ges.

Basel 61 : 28-92, 34 figs.

Simon, E. 1884. Les arachnides de France. Tome 5, parts 2 and 3 :

180-808, pi. 26 and 27. Paris.

Soerensen, W. 1898. Arachnida Groenlandica (Acaris exceptis).

Vidensk. Meddl. Dansk. naturh For en. , Kjobenh. 176-235.

Strand, E. 1905. Araneae. In Report of the second Norwegian Arctic

expedtion in the "FRAM". Kristiania, 3 : 22-30.

Strand, E. 1906. Die arctischen Araneae, Opiliones und Chernetes.

Fauna Arctica, 431-478, 3 figs.

Thorell, T. 1872 a. Om Arachnider fr§.n Spetsber gen och Beer en-Eiland.

Ofvers. Kongl. Vet. Akad. Forh., 28(6) : 683-702.

Thorell, T. 1872 b.Om Nagra Arachnider fr cin Grbnland. Ofvers. Kongl.

Vet. Akad. Forh., 29 : 147-166.

208

Spiders of Hazen Camp

Thorell, T. 1875. Notice of some spiders from Labrador. Proc.

Boston Soc. nat. Hist. 17 : 490-504.

Tongiorgi, P. 1963. Ricerche ecologiche sulgi artropodi di una spiaggi

sabbrosa del litorale Tirrenico. Redia 48 : 165-177, 7 figs-.

Turnbull, A. L. , C.D. Dondale and J. H. Redner. 1965. The spider

genus Xysticus C. L. Koch (Araneae : Thomisidae) in Canada. Can.

Ent. 97 : 1233-1280, 179 figs.

Warming, E. 1888. Om Gr^dnlands vegetation. Meddl. om Grpnland

12, Kobenhavn.

Wiehle, H. 1956. Spinnentier e oder Arachnoidea (Araneae). 28. Fam-

ilie Linyphiidae - Baldachinspinnen. In Die Tierwelt Deutschlands

44 : 337 p. , 551figs.

Wiehle, H. I960. Spinnentiere oder Arachnoidea (Araneae). 11.

Micryphantidae - Zwer gspinnen. ' In Die Tierwelt Deutschlands 47 :

620 p. , 1147 figs.

Yong, R. 1961. A preliminary study of Lake Hazen soil, Ellesmere

Island, N. W. T. Defence Res. Board of Canada. D. Phys. R. (G),

Hazen 16 : 10 p.

Yong, R., S. J. Windisch, and L. Limperis. 1962. A study of Lake

Hazen soils. Part I. (Ellesmere Island). Dept. Civil Eng. and

Appl. Mechanics, McGill University. Soil Mechanics, ser. no. 5 :

76 p. , 6 pictures, 24 figs.

Some of the papers cited above are not referred to in the text but

are included because of their general importance in the literature. On

synonymies and taxonomy see Bishop and Crosby 1935, 1938; Crosby

and Bishop 1931, 1933, 1936; Dondale 1964; Gertsch 1939, 1953; Locket

and Millidge 1951, 1953, and Petrunkevitch 1911. On distributions of

spiders see Banks 1900; Braendegaard 1940, 1958; Chamberlin and

I vie 1947; Dahl 1928, 1933; Holm 1937a, 1937b, 1944, 1945, 1950,

1951, 1958c; Jackson 1930, 1938; Lenz 1897; Strand 1906, and Thorell

1875. On solar orientation see Papi 1955; Papi and Tongiorgi 1962-63,

1963; Tongiorgi 1963. On general biology of spiders see Bristowe 1958;

Crompton 1950; Dondale 1961, and Hackman 1957 . And on the Pleistocene

epoch see Ericson et al. 1964 and Flint 1957.

Leech

209

INDEX

DICTYNIDAE

Dictyna borealis Pickard-Cambridge, 1877 160

LYCOSIDAE

Pardosa glacialis (Thorell, 1872) 163

Tarentula exasperans Pickard-Cambridge, 1877 173

LINYPHIIDAE

Collinsia spetsbergensis ( Thor ell, 1872) 176

Collinsia thulensis (Jackson, 1934) 178

Cornicularis harpinshii (Pickard-Cambridge, 1873) 179

Erigone psychrophila Thorell, 1872 182

Hilaira vexatrix (Pickard-Cambridge, 1877) 184

Meioneta nigripes (Simon, 1884) r 186

Minyriolus pampia Chamberlin, 1948) 189

Savignya barbata (Koch, 1879) 190

Typhochraestus latithorax (Strand, 1905) 192

THOMISIDAE

Xysticus deichmanni Soerensen, 1898 194

PLATES

Figs. 32-34. Pardosa glacialis ventral view, palpus d; ventral view, epig-

ynum; dorsal view, carapace and opisthosoma.

Figs. 35-37. Tarentula exasperans ventral view, epigynum; ventral view,

right palpus dorsal view, carapace and opisthosoma.

Figs. 38-39. Dictyna borealis ventral view, left palpus d* ; ventral view,

epigynum .

Figs. 40-41. Xysticus deichmanni ventral view, epigynum; ventral view,

right palpus cf.

Fig. 42. Erigone psychrophila lateral view, right palpus d*.

Figs. 43-44. Erigone psychrophila ventral view, epigynum; posterior view,

epigynum.

Figs. 45-46. Hilaira vexatrix posterior view, epigynum; lateral view, right

palpus cf.

Figs. 47-50. Minyriolus pampia ventral view, epigynum; dorsal view,

carapace; dorsal view, opisthosoma; lateral view, right palpus <f.

Figs. 51-53. Cornicularia harpinshii ventral view, epigynum; lateral view

right palpus d\ lateral view, carapace d*.

Figs. 54-57. Typhochraestus latithorax lateral view, right palpus d; dorsal

view, carapace d; lateral view, carapace d; ventral view, epigynum.

Figs. 58-61. Savignya barbata lateral view, rightpalpus; front view, ceph-

alic region, carapace d*; lateral view, carapace d; ventral view, epi-

gynum.

Figs. 62-64. Meioneta nigripes lateral view, right palpus ventral view,

epigynum; posterior view, epigynum.

Figs. 65-66. Collinsia spetsbergensis ventral view, epigynum; lateral view,

right palpus d*.

Figs. 67-69. Collinsia thulensis lateral view, right palpus posterior view,

epigynum; ventral view, epigynum.

210

4.0mm

211

0.4 mm

212

0.2 mm

213

Book Review

ANNUAL REVIEW OF ENTOMOLOGY. Volume 11, 1966. Annual Re-

views Incorporated, Palo Alto, California, in cooperation with the En-

tomological Society of America . viii + 596 pp. 3052 refs, 27 figs. $8.50

U.S.A., $9.00 elsewhere. With some comments on the fir st ten volumes .

This eleventh volume in the series is no less essential than any of

its ten predecessors; in fact, perhaps the> extra hundred or so pages

over some of the earlier volumes make it more so. That it is essential,

however, makes it particularly important that attention should be drawn

to some of its shortcomings.

The title of a review article should, in a sense, be a review of its

contents. Several titles here are not; the worst is the last reviewin the

book, entitled "Pest Control", which covers little besides the control of

industrial and domestic pests, an area last covered in Volume 1 of the

Review. Jacobson's "Chemical insect attractants and repellents" ends

abruptly after the heading Synthetic Repellents, an area of wide current

interest, yet includes a treatment of phagostimulants - but only those in

plants. The authors explain these things in part, but it would be better

if they didn't have to.

The indexes to author s and subjects are invaluable . That to chapter

titles could be much better organized; for years it has opened with the

solecism ACARACIDES, while this is cross referenced to Insecticides,

the only reference thereunder is spelt acaricides. More importantly,

the main headings are not mutually exclusive: 'Application of Insecticides '

and 'Resistance to Chemicals' should surely come under 'Insecticides

and Toxicology'; 'Apiculture and Pollination' and 'Vectors of Plant Patho-

gens ' should come under 'Agricultural Entomology ' , 'Population Ecology '

under 'Ecology', and 'Nutrition' under 'Physiology'. While many chapter

titles need duplicate entry under this system, only a few have been ac-

corded this .

If the chapters of the first eleven volumes are tabulated under the

following tentative system, they fall as indicated by the number s in brac-

kets, and very few titles present difficulties: -

Historical and General Entomology ( 1) ; Morphology (7) ; Taxonomy

(20: general 12, apterygotes 0, exopterygotes 3, endopterygotes 5);

Physiology (45); Ecology (47); Applied Entomology - General (45); Ap-

plied Entomology - Agricultural and Forest (25); Applied Entomology -

Medical and Veter inary (22); Applied Entomology - Industrial and Dom-

estic (2); Applied Entomology - Benefactory (9). This process reveals

some imbalance in coverage, beyond that necessitated by the imbalance

of the current research efforts of entomologists. Specifically, we could

do with many more general articles (c.f. Remington and Remington,

Volume 6); substantially more on morphology - including that revealed

by electron microscopy; more taxonomy, especially of the apterygotes

and exopterygotes; and more on industrial and domestic entomology -

perhaps especially in the context of population ecology and new approaches

to control. This same system suggested for indexing might well be used

to impose some sequence on the chapters themselves; the random ar-

rangement used hitherto, without even chapter numbers, has nothing to

214

recommend it to the user.

Several author s in Volume 11 find itnecessary to apologise for this

or that omission from their reviews, attributing this to space limitations .

In so doing they squander space whichmight have beenused to repair the

omission. Most users of Annual Reviews know by now that space is

limited.

Although "it is often (always?) easier to review our ignorance than

to repair it" as Weaver says (page 79), we all owe an immense debt of

gratitude to those who undertake the time-consuming task of writing these

reviews. I particularly enjoyed Evans' delightful "Behaviour patterns

of solitary wasps " . Hoogstraal's "Ticks and human viral diseases" with

355 references is as complete and cosmopolitan a treatment as anybody

could wish. The fine study of the Triatominae by U singer and his allies ,

an original work as well as a review, is at the other end of the scale with

28 references. The 3052 references (less duplications) in the book as a

whole comprise a substantial part of a year's accretion to the entomolo-

gical literature. Other particularly timely and important contributions

areKroeger and Lezzi's "Gene action ininsect development", Madelin's

botanical approach in "Fungal parasites of insects", and perhaps the

most hopeful sign for the future of applied entomology, Geier's "Manage-

ment of insect pests".

The establishment of a special price for students is to be applauded.

Brian Hocking

CORRIGENDA

P . 8, para. 5, 1. 6 delete "line", substitute: "reservoir and the genital

opening, replacing the term ejaculatory duct".

P. 63, para 3, 1. 6, delete "Odontotar sini", substitute "Odontoscelini" .

Publication of Questiones Entomologicae was started in 1965 as part

of a memorial project for Professor E. H. Strickland, the founder of the

Department of Entomology at the University of Alberta in Edmonton

in 1922.

It is intended to provide prompt low-cost publication for accounts of

entomological research of greater than average length, with priority given

to work in Professor Strickland’s special fields of interest including ento-

mology in Alberta, systematic work, and other papers based on work

done at the University of Alberta.

Copy should conform to the Style Manual for Biological Journals

published by the American Institute of Biological Sciences, Second

Edition, 1964, except as regards the abbreviations of titles of periodicals

which should be those given in the World List of Scientific Periodicals,

1964 Edition. The appropriate abbreviation for this journal is Quaest. ent.

An abstract of not more than 500 words is required. All manuscripts will

be reviewed by referees.

Illustrations and tables must be suitable for reproduction on a page

size of 9%X6% inches, text and tables not more than 7 inches,

plates and figures not more than 8^X5 inches. Reprints must be ordered

when proofs are returned, and will be supplied at cost. Subscription rates

are the same for institutions, libraries, and individuals, $4.00 per

volume of 4 issues, normally appearing at quarterly intervals; single

issues $1.00. An abstract edition is available, printed on one side of

paper or on one or both sides (according to length) of 3X5 inch index

cards (at $1.00 per volume) or on 5X8 inch standard single row punched

cards ($1.50 per volume).

Communications regarding subscriptions and exchanges should be

addressed to the Subscription Manager and regarding manuscripts to:

The Editor, Quaestiones Entomologicae,

Department of Entomology,

University of Alberta, Edmonton, Canada.

J

Quaestiones

entomologicae

A periodical record of entomological investigations,

published at the Department of Entomology, Uni-

versity of Alberta, Edmonton, Canada.

VOLUME 11

NUMBER 3

JULY 1966

QUAESTIONES ENTOMOLOGICAE

215

A periodical record of entomological investigations, published at

the Department of Entomology, University of Alberta, Edmonton, Alberta.

Volume 2 Number 3 4 July 1966

CONTENTS

Editorial 215

Corrigenda 216

Nimmo - The arrival pattern of Trichoptera at artificial

light near Montreal, Quebec 217

Sharplin - An annotated list of the Formicidae (Hymenoptera)

of central and southern Alberta 243

Shamsuddin - A Bathymermis species (Mermithidae : Nematoda)

parasitic on larval tabanids 2 53

Book review 256

Editorial - Bridge builders?

It is a common weakness of peoples and generations brought up

without history to attach undue importance to the discoveries of their own

lifetimes. Amid the excitement over DNA and the endoplasmic reticulum

and other revelations of the electron microscope and the ultracentrifuge,

those of us who are aware we are aging may take comfort in the thought

that an insect is still an insect and a plant still a plant and in the assur-

ance that the solid body of knowledge handed down from previous gener-

ations, despite the thrilling rate at which it is being added to, remains

essentially unassailable. Morphology remains morphology, though its

horizon dips to the molecular level, into the limbo designated by that

most unfortunate misnomer "ultrastructure”.

Physiology remains physiology; it hasn't become biochemistry

despite the Cinderella status of biophysics and especially biomechanics;

the newper spectives in this field do not alter its dependence on morph-

ology. Indeed the revelations of molecular structure serve to emphasize

more strongly than ever the necessity for a thorough understanding of

structure before functional speculation or even experimentation is in-

dulged in.

Those of us who are not yet aware that we are aging should be dis-

comforted by the reflection that history and aging go hand in hand but that

the one repeats itself and the other doesn't. It is, indeed, the repetitions

of history which give it the perspective generating quality that makes it

an essential part of the study of any subject.

Even a molecular biologist needs a name for a species he works

with, and needs in consequence to be aware of the vagaries of names and

their application and indeed of the principles of systematics. It makes

no more sense at the molecular level than at any other to study the func-

tions of an organ one cannot find, in a species one cannot name. And

names, of course, have histories, even if we have decreed that those

proceeding further back than 1758 should have academic status only, and

iSmvmw

216

these histories have a future. It might, in fact, be said that the purpose

of most biological work, including that at the molecular level, is to con-

tribute to the future of names. For evolution is no more than the history

of life, and both as an end in itself and as a contribution to human welfar e,

nothing could be more central to biology than a detailed and accurate

understanding of the past of evolution. This is the true aim of system-

atics; its attainment should permit predictions as to faunal future to

replace the chaotic mayhem man occasions today. The provision of

handles for taxa is but a serendipitous appendage.

Given this grand aim it is doubly unfortunate that biologists should

attach to themselves such belittling prefixes as 'micro- 1 and 'molecular '.

Surely better names can be found; perhaps it is even the bearing of these

names that contributes to the very evident rift which has developed in

recent years between micr o-thinking groups in both the traditional and

the molecular areas of biology, to the detriment of both. We cannot

afford such little luxuries.

There is no better bridge between the macro and the micro than the

entomologist; his subjects of study force him into both camps, they are

both small enough and hardy enough to be superior subjects for micro

and molecular study, and ubiquitous enough and diver s e enough in struc-

ture, function, and relationships to compel attention from the tradition-

alist. People come to resemble the subjects of their studies; while we

may describe the brains of insects as small, both relatively and absolute-

ly, let us as entomologists stretch ours to build this bridge.

Brian Hocking

C OR R I GENDA

P. 209 (Vol. II No. 2). For: Figs. 65-66, read Figs. 65 & 69.

(C. spetsbergensis )

For: Figs. 67-69, read: Figs. 66-68.

(C. thulensis)

217

THE ARRIVAL PATTERN OF TRICHOPTERA AT ARTIFICIAL LIGHT

NEAR MONTREAL, QUEBEC *

ANDREW P. NIMMO

Department of Entomology Quaestiones entomologicae

University of Alberta, Edmonton, Alberta 2 : 217 — 242 1966

The arrival pattern of Trichoptera at artificial light at lie Ste. Helene, in the St. Lawrence River

opposite Montreal, Quebec, is examined. A Robinson trap with a mercury vapour light bulb was combined

with a Lafrance trap which changed containers hourly. Ten minute catch periods were used to examine the

evening peak in detail, containers being changed manually. Numbers and sex ratios for each of 78 species

taken are given. One species not then described, and two other doubtful forms, females, were noted. Thir-

ty one genera and thirteen families are represented. The pattern in each of 7 species examined in detail

is noctumally bimodal, with only a small morning peak. The roles of light, temperature, wind, relative

humidity, and saturation deficit in determining total catches per night and fluctuations of numbers within

any one night, are examined. Temperature and wind are the primary factors, with light fixing the time of

the evening and morning peaks. Neither relative humidity nor saturation deficit seemed to be of any sig-

nificance, at the values experienced. A differential effect of wind on flight, depending on species size, is

shown. Sex ratios throughout the night are briefly examined, and it is concluded that no one sex of any of

the seven major species is alone responsible for any peak. It is considered that the pattern of arrival at

light reflects a natural pattern of flight activity.

It has previously beenfound that in East Africa (Corbet Tj^nneland

1955) the numbers of Trichoptera at lights vary throughout the night ac-

cording to a pattern. This study is an investigation of the arrival pattern

of Trichoptera at artificial light at night at lie Ste. Helene in the St.

Lawrence River opposite Montreal, Quebec. The effects of meteoro-

logical factors, including natural light, on the pattern were examined.

While the study deals with the pattern of arrival at artificial light, and

the results can only be dir ectly interpreted within the observed conditions,

some attempt is made in the discussion to relate the pattern found to the

natural pattern of flight activity, independent of artificial stimuli.

METHODS AND EQUIPMENT

To examine patterns of animal activity relative to time, the time

involved must be subdivided to a number of equal periods, here called

'catch periods' or just 'periods'. The population at light was sampled

during successive catch periods. To compare patterns between nights

and obtain meaningful average patterns for the summer, it is necessary

to start any one chosen catch period at the same solar time each evening.

* Publication No. 6 resulting from the World Exhibition Shadfly Project:

Canada Department of Agriculture, R e search Branch; Provincial Depart-

ment of Agriculture, Quebec; and Canadian Corporation for the 1967

World Exhibition.

218

Trichoptera at Artificial Light

Two solar times were used: sunset (solar elevation minus 0. 83°) and

civil twilight (solar elevation minus 6°). These times, translatedto local

clock time for each night, were obtained from tables prepared by the

Dominion Observatory, Ottawa, and the Meteorology Branch of the Canada

Department of Agriculture, Ottawa. The times were prepared for the

latitude and longitude of lie Ste. Helene (45°31'N, 73°32'W).

It was decided to examine the pattern of the entire night using 1 hr

catch periods, the first of which was to start one half hour prior to sun-

set, and to examine the evening peak in more detail, using 10 minute

periods, starting one hour prior to civil twilight. Civil twilight was used

for the 10 min catches as it was noted after running several nights at 1

hr periods that the massive upsurge to the evening peak generally oc-

curred in the second period, in which civil twilight also occur red. It was

felt that civil twilight might be of significance to the evening peak of flight

activity and the 10 min sessions were designed to determine the timing

and form of the evening peak, and perhaps the factors controlling it.

The 1 hr catching period nights always ran for a total of 12 hrs

as this was the capacity of the automatic trapused andcovered the entire

night, ending well past sunrise. As many as 19 ten minute periods were

run on some nights, but 13 was decided on as the minimum which sufficed

to cover the period of peak flight activity. On one night trapping was

stopped after 9 periods due to cold; the data areonlyused in the results

when considering catch nights individually.

Trapping Methods and Equipment

Trapping was done at the old Fort (figs. 1 & 2), The 1 hr catches

were taken with a mechanical trapping device designed and built by Mr.

J. Lafrance (1965) of the Canada Department of Agriculture Laboratory,

St. Jean, Quebec, and loaned for this study. It is capable of hourly (±2

min) changing of catch canisters, but can be adjusted for other periods.

Capacity is 12 canisters, and the killing agent used was 7 0% ethanol.

The insects reached the cans by way of a large funnel on top of the

trap body, with the spout passing through the roof to the cans below. On

top of the funnel was placed themetal cone of a Robinson trap (Robinson

& Robinson 1950), which bears a socket for an 'Osram1 125 watt, high

pressure mercury vapour light bulb (230-240 volts; model MB/V). The

light from this bulb is particularly rich in UV light, and is highly attrac-

tive to Trichoptera in consequence (Williams 1951). The spectral com-

position of light from a similar source is given in table 1. A cylindrical

collar, 7" high by 13" diameter, made of file-folder card was placed

around the upper edge of the Robinson cone to reduce the intensity after

anabortive first use of the trap in which somany Trichoptera arrived as

to swamp the cans and neces sitate rejection of the catches for that night.

It was used for both 1 hr and 10 min catches. Even so, some of the

catches taken on some nights were beyond the capacity of the machine.

When this was so the entire night's catches were discarded. Intrinsic

to the R obinson trap cone are 4 vanes set at 90° to each other on the inner

surface, which serve to stun the insects as they spiral inwards and down-

wards towards the light; they then fall into the ethanol below. The vanes

also served to hold the collar in place and hold the light bulb socket.

This part of the Robinson trap was retained and temporarily coupled to

the Lafrance trap.

Nimmo

219

220

Nimmo

TABLE 1- Intensity of the radiation from a high pres sure mercury vapour

bulb corrected for transmission through lime - soda glass

(from Rossler 1939).

* Lower limit of visible radiation in this spectrum.

The same equipment was used to take the 10 minute catches except

that as the Lafrance trap could not be set for ten minute intervals, the

jar was changed manually every ten minutes. Other conditions were as

identical as possible to those of the 1 hr catch periods. The nights on

which trapping took place, and other relevant data, are set out in table

2 (1 hr catches) and table 3 (10 min catches).

TABLE 2 - Dates and times (eastern daylight saving) of the first of 12

consecutive 1 hr catches of Trichoptera at lie Ste. Helene

Montreal, summer 1964.

Catch 12 not obtained.

Trichoptera at Artificial Light

221

TABLE 3 - Dates and times (eastern daylight saving) of the fir st of series

of 10 minute catches of Trichoptera, at lie Ste. Helene,

Montreal, summer 1964.

* One hour prior to civil twilight.

Other Observations

All the 1 hr catches were taken in the period for which continuous

records of temperature, relative humidity, wind, and rainfall were made

on the island, except that wind recording started after the first trapping

night, June 13. The records used here are from the West Station (see

fig. 1), about 20 ft horizontally from the edge of the river, but about 50

ft above the water and 1/4 mile from the trapping site. The station was

setup and maintained by the Canada Department of Transport, in conjunc-

tion with the Shadfly Project.

Temperature and relative humidity were read from the charts ob-

tained at the mid-point of each 1 hr catch period; wind speed, in miles

per hour, is averaged for each catch period; saturation deficit was ob-

tained from temperature and relative humidity by tables. Zenith light

intensity readings were taken at the mid-point of each 10 min period.

It was later discovered that the light readings could not be converted to

foot-candles as the precise spectral sensitivity of the sensor unit was

unknown and could not easily be determined. It has been possible, how-

ever, to obtain a curve of light intensity in arbitrary units (p. 238). As

time permitted, notes on cloud, rainfall, and wind were taken.

Treatment of Catches and Data

Generally, all Trichoptera taken were determined to species and

sex. Without exception all were counted. But occasionally a species or

group of species, Hydroptilidae especially but also Hydr opsychidae and

Protoptila muculata (Hagen), were taken in such numbers that the proportions

of each species and sex had to be estimated from a sub-sample . Selection

of sample size was arbitrary but in general the larger the total numbers,

the smaller the per cent sampled.

222

Nimmo

To reduce the effect of fluctuations due to environmental factors,

the arithmetic values (n) may be transformed to the logarithmic value

(log n) . To bypass the difficulty of zeros, for which there is no logar-

ithmic value, Williams (1937) suggested adding one (1) to all values in a

time series, and transforming the resulting values (n + 1) to logarithms.

If the log (n + 1) values for all periods of any one time series, or equi-

valent periods in terms of solar time of several time series are averaged,

[2 log (n + 1)] / N and the antilog taken, an approximation to the geometric

mean of the series is obtained. This approximation is knownas Williams*

mean (Haddow I960), and is symbolized as Mw. The value 2 log (n + 1) /

N, when obtained for equivalent periods of several time series and plotted

against time gives an average pattern for these time series.

RESULTS

Numbers of Specimens Examined

Table 4 lists the species taken, in descending order of numbers

taken per species in all catches. The total number of specimens of

Trichoptera was 297, 967 for all species. A total of 78 species,

in 31 genera and 13 families was taken, plus 2 doubtful forms. One of

these, Cheumatopsyche montrealensis Nimmo ( 1966) was not then described. The

following species were selected for detailed examination of data, andare

given in order of abundance: Hydroptila spatulata Morton, Cheumatopsyche speciosa

(Banks), Protop tila mac ulata (Hagen), Hydropsyche recurvata Banks, Psychomyia flavida

Hagen, Athripsodes cancellatus (Betten), and Athripsodes tarsipunctatus (Vorhies).

Agraylea multipunctata Curtis was not selected although more numerous than

Athripsodes tarsipunctatus because the catch was spread over 27 nights whereas

that of A. tarsipunctatus was concentrated into 18. Also in table 4, the total

numbers per species are broken down to sexes, sex ratios (per cent

males) as in Henderson, Henderson & Kenneth ( I960) , and finally the range

of dates on which each species was taken is given, concerning which it

must be remembered that trapping started on June 2 and ended August 31.

The ratios may be artifacts of the trapping method, due to differ-

ential attraction of the sexes. One remarkable fact emerges from table

4, however. P. flavida shows a sex ratio of virtually zero. Marshall

(1939) obtained similar results, but cautioned about the possible differ-

ential attraction. Betten (1934) states that he had only 2 specimens,

females, but that Sibley (1926) took 893, in an unspecified manner, all

of them female. If this is the natural ratio, then P. flavida must be usually

parthenogenetic . Crichton (I960) considers in detail only the ratios of

species with over 100 individuals taken, which is also done here. Only

26 species qualify of which 17 give ratios above 50, 3 of them close to

this: Leptocella Candida { 52.00), ' Agraylea multipunctata (53.05) and Athripsodes ancylus

(53. 78).

Pattern of Arrival at Artificial Light

The data will be examined first in the form of total numbers of

Trichoptera per catch, after which the separate data on the previously

selected species will be examined, for both 1 hr and 10 min catches.

Trichoptera at Artificial Light

223

TABLE 4 - Species of Trichoptera taken at lie Ste. Helene, Montreal,

summer 1964 in descending order of numbers taken in both

1 hr and 10 min catches.

224

Nimmo

TABLE 4 (cont. )

Grand total 297, 967

Trichoptera at Artificial Light

225

Species patterns follow closely the total number s patterns . The average

pattern for each species for the summer, will, however, be presented.

Only 1 hr catches will be considered relative to weather, as it is im-

possible to read values accurately from the meteorological charts for

intervals as small as 10 minutes.

1 Hour Catches - Total Numbers per Catch

In fig. 3 a peak occurs generally in the second period. In 2 of 15

nights the peak occurred in period three. This may be due to extrinsic

factors obscuring or delaying the peak. A second peak, slight or other-

wise, is found in the first or second period immediately prior to sun-

rise. This is the morning peak.

Between the two peaks, evening and morning, it is seen that adults

are taken, occasionally in very nicely deer easing s eries, as in fig. 3(35-

26 August), but often in widely varying numbers.

The average pattern for the summer, as determined by the nights

on which trapping was carried out, is shown by fig. 4 (total numbers) .

The distinct evening peak is seen, but the morning peak is not manifest.

This is due to sunrise shift during the season in relation to sunset and

hence also in relation to the catch periods.

It remains now to demonstrate the dependence of the peaks on natural

light intensity, and to explain the intervening period of gradual decline,

or fluctuation as the case may be.

Dependence of the Peaks on Natural Light Intensity

The least fluctuating nightly graphs are selected for visual exam-

ination of the concomitant environmental factors. These graphs of fig.

3 (17-18 June, 13-14 July, and 25-26 August) show the peaks well. Table

5 presents the meteor ological data for these three nights and examination

shows that temperature is either declining throughout the night, usually

slowly, or holding steady, but never increasing. Next, wind holds steady

for at least the first two periods, in which the evening peak occurs.

Finally, relative humidity and saturation deficit seem to fluctuate er-

ratically. On the night of 13-14 July, however, they held steady for 5

hour s preceding and including the morning peak. This seems to rule out

these two factors as influencing the peak, at least at the values encoun-

tered here.

Thus meteorological factors are either declining fairly evenly,

holding steady, or fluctuating and showing no correlation with log (n + 1),

yet the peaks occur outstandingly. The first catch is fixed on time of

sunset, and the evening peak always occurs in the same period, the second.

The morning peak may occur in period 9, 10, or 11, depending on the

season, but always in the pre- sunrise period.

Table 5 omits only light intensity. In the evening, with all factors

generally declining, there is a peak in numbers of Trichoptera caught.

In the morning, the same conditions prevailing, there is another peak in

numbers caught. The only factor which has equal values in both evening

and morning is light intensity. Obviously from the graphs, the light values

involved occur after sunset and before sunrise. Thus the causal (or

triggering) factor of both peaks appears to be a certain light intensity.

226

Nimmo

1-2 July

4

3.

2

1.

0

4

31

2

1

III

18-19 July

16-17 June

,IL ,

4-5 July

1 5 10

27-28 June

bk

13-14 July

3-4 August

25-26 August

hi

1 5 10

Figure 3 - Total hourly numbers of Trichoptera at UV light, lie Ste.

Helene, Montreal, summer 1964. Abscissae 1 hr periods, ordinates

log (n + 1), plotted at the period mid-points: Suns et coincident with first

period mid-point. Arrows indicate approximate sunrise.

Trichoptera at Artificial Light

227

Figure 4 - Total numbers and numbers of 7 species of Trichoptera separ-

ately, taken at He Ste. Helene, Montreal, in a UV light trap on 16 nights

(or fewer), 1964; average values per 1 hr period for the summer. Ab-

scissae 1 hr periods, ordinates Williams ' mean (Mw) transformed to %.

Sunset coincident with the first period mid-point.

228

Nimmo

TABLE 5 - Meteor ological data and log (n+1) of total number s of Trich-

optera taken per 1 hr period at lie Ste. Helene, Montreal,

on three selected evenings, summer 1964.

June

17-18

July

13-14

August

25-26

* S.D. = saturation deficit, in inches of mercury.

** Peaks.

Trichoptera at Artificial Light

229

Effects of Other Environmental Factors

The night as a whole _ Examined here are those periods of each night

between and including sunset and sunrise.

Means of each factor, temperature, wind speed, relative humidity,

and saturation deficit were obtained for each night. Each factor was then

plotted against the mean log (n + 1) values for each night. As seen in

fig. 5, temperature and log(n+l) are clear ly cor related; no other factor

showed any significant correlation. That is, on an evening of high tem-

peratures (7 0-80 F) one can expect a large catch, the opposite also holding

true. Cor relation coefficients are not calculated here as this is prelim-

inary to an analysis of the data for the periods without sunlight. It should

be noted that catches may be low despite high temperatures, due to re-

lative seasonal scarcity of Trichoptera. This explains some lack of

correlation in fig. 5. Other unrecorded or unrecognized factors may

also be involved. Wind speed shows little apparent correlation with log

(n + 1), although it can have a distinct effect on flight activity. The best

available example is the night of 18-19 July (table 6):

TABLE 6 - Total number s of Trichoptera taken, andmeteorological data

for 1 hr catch periods 1-9, 18-19 July, lie Ste. Helene,

Montreal.

Temperature was high and decreasing slowly. While a morning peak is

clear, the evening peak is represented only by the small stubs in periods

2 and 3. As the wind dropped, rather abruptly, there was an upsurge in

numbers of Trichoptera taken. This catch is one of the most useful of

all catches taken. It will be noted that in fig. 5 the plotted position of

18-19 July is one of the poorly correlated catches. If the evening had

been 'normal', wind being low or absent, the point would probably have

been located well to the right.

Wind on the night of 16-17 June varied from 17 to 22 mph and the

pattern is clear (fig. 3). It varied from 13 to 18 mph on 23-24 July and

the pattern appeared. On 1-2 August the wind varied from 2 to 8 mph,

but the temperature fell 14°F; the pattern is obscured by fluctuations

between the peaks. Thus, if the effect of wind be removed, it is seen

that temperature plays a major role in determining flight activity. The

fact that most Trichoptera species fly en masse at night and not during the

day when temperatures are higher, suggests an inhibiting effect of either

or both of temperature or light. Wind can only make it difficult, or im-

possible, for the insects to fly, and thus to come to the trap, no matter

how much theymay be encouraged to fly by high temperature. The pat-

tern will remain clear but numbers will be reduced. Thus wind and

230

Nimmo

-80°F

-70

-60

#16 Jn

1

#18 Jy

1 Jy

* s*

k8 Au

27 Jn*

•3 Au

#17 Jn

V

13 Jn

%

*4 Jy

1 Au

►19 Au

#12 Au

2i 3i

log (n + 1 )

Figure 5 - Mean night (plus sunset & sunris e periods) temper atur e plotted

against mean log (n + 1) of the hourly catch of Trichoptera.

-80°F

-70

-60

#16 Jn

#18 Jy

25 Au '

8 Au •

• 3 Au

#17 Jn

nV #13 Jn

#6 Jy

• @4 Jy

23 Jy

#19 Au

#1 Jy

• 1 Au

#12 Au

2, 3i

log ( n + T )

Figure 6 - Mean night (inter-peak) temper atur e plotted against the mean

log (n + 1) of the hourly catch of Trichoptera.

m.p. h.

Trichoptera at Artificial Light

231

temperature seem to be major factors in determining the overall catch

for any one night. But the peaks, bar ring such exceptional circumstances

as occurred on the night of 18-19 July, will remain detectable.

The periods between the peaks - To determine more precisely the effects

of environmental factor s the peak period data must be omitted from con-

sideration, and the inter-peak periods examined more closely. The

periods involved here are numbers 3 to 7, 3 to 8, or 3 to 9, depending

on the time of sunrise.

Saturation deficit is not considered as it fluctuates from one hour

to the next, with no apparent cor relation with Trichoptera numbers. Figs.

6 and 7 illustrate mean log (n + 1) plotted against mean temperature and

mean wind speed, for the appropriate sets of periods on collection nights .

Fig. 6 shows a strong cor relation of temperature with mean log (n + 1).

Wind speed (fig. 7) shows some negative cor relation as expected, but this

is obscured since wind is secondary to temperature. Figs. 6, 7, and 8

collectively demonstrate the role of wind in disrupting the effect of tem-

perature on the flying population. Fig. 8 gives mean log (n + 1) plotted

against mean temperature times mean wind speed. It will be seen that

the distribution of nights is similar to that of fig. 7; as temperature is

also' involved, the vertical spread in fig. 8 is slightly gr eater than in fig.

7. In fig. 6, showing mean log (n + 1) against mean temperature, the

distribution is entirely different, and the correlation is much improved,

and positive. This is further evidence of the overriding effect of tem-

perature and the subsidiary disrupting effect of wind on flight of Trich-

optera in the time between the peaks.

r2Q ® 16 Jn

-15

-10

-5

*12 Au *23 Jy

#25 Au

*8 Au

• 18 Jy

Jl

• 17 Jn

• 4 Jy

*19 Au

• 27 Jn

1 Au

*3 Au *6 Jy # •! Jy

• 13 Jy

2, 3,

log ( n + 1 )

Figure 7 - Mean wind speed against mean log (n + 1) of inter-peak catch

of Trichoptera.

( s00l ) H'duJ * do

232

Nimmo

-10

• 16 Jn

-8

• 23 Jy

#18 Jy

#25 Au

#12 Au

#8 Au

-6

-4

#17 Jn

#4 Jy

27 Jn

• *19 Au

-2

• 3 Au

#6 Jy

#13 Jy

# i Jy

•l Au

2, 3,

log ( n +l)

Figure 8- Mean temperature X mean wind speed against mean log (n + 1)

of inter-peak catch of Trichoptera.

Fig. 9 presents log (n + 1) plotted against temperature for each

separate period, as limited in this section, for all catch nights. Again

a definite correlation is seen, in more detail. The cros ses represent the

5 catches from the night of 18-19 July and it will be obs erved that periods

3, 4, and 5, before the wind dropped, show low catches relative to the

rest of the scatter.

In a statistical analysis of the data from inter-peak periods, the

method of multiple correlation as set out in detail by Croxton & Cowden

(1955) was used. Values of log (n + 1) are designated in the following as

X^; of temperature, X^; of wind speed, X3. Details of the calculations

are omitted, suffice it to summarize the results, X^ being the dependent

variable:

Using one independent variable: X£ or X3

Total variation of X^, £ x ^ = 69.7634

Variation explained by use of only = 32.4282

Standard error of estimate = 0. 6789

Coefficient of correlation, r12 = +0.6817

Thus variation in temperature serves to explain 68% of the variation in

log (n + 1), or 68% of the changes in number s are associated with changes

in temperature.

Trichoptera at Artificial Light

233

[90 °F

Periods 3 +

+ 4

5 +

80

18-19 July

70

60

50

lL

2i

I og (n + 1 )

Figure 9 - Temperature against numbers of Trichoptera (as log (n + 1))

taken at an ultra-violet light trap at lie Ste. Hfelene, Montreal, summer

1964. Each 1 hour inter-peak period for all catch nights plotted separ-

ately.

Variation explained by use of X3 only = 14. 3372

Standard error of estimate = 0. 8272

Coefficient of correlation = -0.4533

Thus variation in wind, serves to explain 45% of the variation of log (n + 1) ;

i.e. 45% of the changes in log (n + 1) are associated with changes in wind

speed.

Using two independent variables: X£ and X3

Total variation, xr^ = 69.7634

Explained variation, xc. 23 = 39. 6811

Coefficient of correlation, ^1.23 = +0.7541

Thus temperature and wind speed together account for 7 5% of the variation

of log (n + 1); i. e. 75% of the changes in log (n + 1) are associated with

changes in temperature, wind speed, or both.

As the combined effect of these two factors on numbers of insects

taken is to the extent of 75%, the interaction of temperatur e and wind may

be assumed to be 38% (i.e. 45- (75-68) = 38). The remaining 25% of

variation may be attributed to saturation deficit and other unmeasured

and unrecognized factors. An application of the F test for the reliability

of ^1.23 shows this to be clearly significant, that is, the correlation

between X^, and X£ ^ 3 appears to be very good.

234

Nimmo

The pattern at the species level- For each species the graphed pattern of

only one night is used as the species patterns follow the total numbers

pattern closely. The night chosen for each species was that which showed

the pattern most clearly. Varying seasonal occurrence prevented the

same night being used for all species, but only two nights were needed:

13-14 June and 25-26 August. Species and night are given in fig. 10.

It will be seen in these graphs that all seven species tend to follow the

pattern, with differences, of course, but these are minor.

Included in fig. 10 are 2 additional graphs, for H. recurvata and

C. speciosa . They are both for the night of 18-19 June on which the

speed suddenly decreased. In these two figures an evening peak is dis-

cernible, especially in H. recurvata . The overall depression of the first

half of the night shows, but the species numbers rose to a peak, then fell

away; the wind dropped and the number s recover ed to the assumed 'nor-

mal' for that night. On this night Athripsodes cancellatus also showed a peak,

but not so well. The inter esting point here is, that it was the three large

species (body length > 4 mm) which produced discernible evening peaks

of flight activity despite the high wind. Three other species, Psychomyia

flavida , Protoptila maculata , and Hydroptila spatulata , showed no evidence of a peak at

all: they are all micr o - Trichoptera. Thus sizeis seen to be of importance

to a species in maintaining the pattern of night time activity, if winds are

high and fluctuating. This diver sity due to size may well be another factor

in the 25% variation in log (n + 1) remaining to be explained.

In fig. 4 are graphs of the patterns using mean values of log (n + 1) .

Allowance should be made for sunrise shift.

Sex Ratios

Sex ratios were examined to determine if the sexes were active at

different times. In all seven species considered in detail, at least fifty

per cent of the total numbers of each night arrived by period three. In

most cases the ratio is about 50 throughout the night.

It seems safe to conclude that the pattern of total number s of Trich-

opter.a taken at light is due neither to any one sex of any one or more

species, nor to any species as a whole.

Ten Minute Catches — Total Numbers

Fig. 11 shows patterns for individual nights using log (n+1) values,

and fig. 12 shows the average pattern for the summer , using mean values

of log (n + 1).

The points to observe are as follows. For the first six periods

there were no catches or, at most, small numbers: occasionally catches

in periods 4 to 6 were substantial. The peak of flight activity generally

occurred in catch period 7 but sometimes in period 6, 8, or even 9.

F rom figs . 11 and 12 it will be seen that the peak follows immediately

after civil twilight. Fig. 13 shows that a sharp change in rate of decline

of light intensity occurs at civil twilight.

The curve of activity generally starts to rise prior to civil twilight,

indicating a response either to low, or a lowering of, light intensity.

However, the sudden upsurge to the peak seems to be associated with the

sudden change in rate of decline of light at civil twilight. Harker (1961)

235

A. tarsipunctatus ‘JJ recurvata C. speciosa

Figure 10 - Pattern of arrival on selected nights of seven species of

Trichoptera at a UV light trap at lie Ste. Helene, Montreal, summer

1964. Two additional graphs are included for reasons given in the text.

Abscissae in 1 hr periods, ordinates numbers as log (n + 1). Sunset

coincident with the first period mid-point. Sunrise indicated byarrows.

points out that ". . . , it is rare for activity to occur as an immediate

reaction to change in light intensity". It could be, therefore, that the

peak is a delayed reaction to the light values of earlier periods, which

are themselves vastly lower than normal daytime values. Nielsen's

(1963) summary of the situation in poikilotherms : "The releasing factor

may be a certain low level of illumination, or it might be a certain rate

of change of intensity or a combination of both" s eems appropriate to the

uncertainty concerning the role of natural light in producing the peaks in

Trichoptera. One further possibility is that the change in the rate of

decline of light intensity induces’ an immediate increase in flight activity.

236

Nimmo

Figure 11 - Total number of Trichoptera taken at a UV light trap at lie

Ste. Helene, Montreal, summer 1964, for each night on which 10 min

catch periods were used. Ordinates in log (n + 1) , abscissae in 10 minute

periods, log values plotted at the period mid-points. Civil twilight indi-

cated by the arrows.

DISCUSSION

Previous Studies of Nocturnal Activity Patterns in Trichoptera

Light trap studies of the nocturnal flight activity rhythms of insects

of immediate interest, are those by Williams (1935 and others), Stage

Chamberlin (1945), Southwood (I960), Corbet and Tj^nneland (1955),

and Brindle (1957 a, b, and 1958). Most papers mention Trichoptera in

passing, if at all. Trichoptera have been studied seasonally (Crichton

I960, Marshall 1939), rather than hourly, as here; such studies are

consequently of little interest in the present context. It is unfortunate

that there appear to have been no studies of Trichoptera using non-at-

tractive traps, other than that of Lewis & Taylor (1965).

Trichoptera at Artificial Light

237

T otal nos

JLU

2

1.5J

1

0.5]

0

J//.5p

atulata

-ill

C. speciosa

j|

2

1.51

1

0.51

0

H. recurvata

J

P. maculata

illJ

P. flavida

A. cancellatus

2

1.5

1.

0.5

0

A. tarsipunctatus

I

Figure 12 - Means of numbers of Trichoptera taken per 10 min period

at an ultra-violet light trap at He Ste. Helene, Montreal, for those equi-

valent periods of each night on which trapping was carried out, summer

1964. Abscissae in 10 min periods, ordinates in log (n + 1) with values

plotted at period mid-points. The arrows mark civil twilight.

238

Nimmo

The Pattern at Montreal

The pattern found in this study has the following characteristics:

bimodal, the evening peak relatively much more pronounced than the

morning peak; the pattern of numbers in periods exclusive of the peaks

forms a gradually decreasing slope from the evening peak till the slight

rise to the morning peak; the interpeak slope may be punctuated by fluc-

tuations of varying degrees, dependent on meteorological factors; the

morning peak is terminated by an abrupt drop-off in numbers to zero,

or almost zero. R eferring to Corbet & Tj^nneland's (1955) classification

of relative development of the two peaks in East African Trichoptera,

the present 7 species seem to fit their class 2 well; "Both peaks dis-