The Pan-Pacific Entomologist

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THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly by the Pacific

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PAN-PACIFIC ENTOMOLOGIST

68(1): 1-7, (1992)

EVOLUTION OF OVIPOSITION HABITS IN APHODIUS

DUNG BEETLES (COLEOPTERA: SCARABAEIDAE)

Nobuyo Yoshida and Haruo Katakura

Zoological Institute, Faculty of Science, Hokkaido University,

Sapporo, 060, Japan

Abstract .—Oviposition habits of nine species of Aphodius dung beetles common in Sapporo,

Hokkaido, northern Japan, were studied under laboratory conditions using glass cages supplied

with soil and fresh cattle dung. Four types of oviposition habits were recognized. Type I: Eggs

were laid singly in the dung on the ground. Type II: Eggs were laid singly in the soil beneath the

dung. Type III: Each egg was laid in a small dung mass stuffed in a shallow burrow excavated

beneath the dung. Type IV: Each egg was laid in the soil near the terminal end of a sausage¬

shaped dung mass buried beneath the dung. Although Types III and IV were similar in that

females provided food for larvae, behavioral sequences of oviposition and provisioning were

distinctly different between the two types. In Type III, an egg was laid after a dung mass was

provided; whereas, in Type IV, an egg was laid before a dung mass was buried. Provisioning

habits of Type III and Type IV seemed to have evolved independently from more primitive

Types I or II, and from Type II, respectively. Oviposition habits of Aphodius were compared

with those of two major groups of scarabaeid dung beetles, Geotrupinae and Scarabaeinae. Our

Type III oviposition habit is analogous to those of certain species of Geotrupinae and Scara¬

baeinae, and Type IV to some Geotrupinae, indicating parallel evolution of dung burying habits

in several lines of scarabaeid beetles.

Key Words.—Insects, Scarabaeidae, Aphodius, dung beetles, oviposition, behavioral sequence,

evolution

Reproductive biology of dung beetles belonging to the subfamily Aphodiinae

has had little attention until recently, in spite of their dominance in the number

of species and individuals in the north temperate zone (Balthasar 1964). Scattered

records show that oviposition habits of Aphodiinae are diverse. Many species lay

eggs directly in dung on the soil surface, or in the soil beneath the dung (Hafez

1939, White 1960, Landin 1961, Hanski 1980). A few species bury dung masses

for larval food in the soil under droppings (Paik 1968; Hosogi et al. 1979, 1980).

Obligatory or facultative kleptoparasitic species are also known (Hammond 1976

[and the references therein], Klemperer 1980, Kiuchi 1987).

Due to this diversity, the Aphodiinae may offer invaluable information for

studying evolutionary origin of more elaborated oviposition habits found in two

major groups of dung beetles: Geotrupinae and Scarabaeinae (for reviews, Halffter

& Matthews 1966, Halffter & Edmonds 1982, Doube 1990). Unfortunately, how¬

ever, oviposition habits of Aphodiinae species have not been studied in detail,

except for a few species such as Aphodius rufipes (L.) (Madle 1934, Holter 1979,

Klemperer 1980). For many species, only the scattered descriptions of egg site

and/or larval feeding sites were available. Furthermore, there have been no quan¬

titative studies specifically dealing with oviposition behavior or evolutionary trends

in Aphodinae species.

In this paper, we will examine four types of oviposition habits distinguished

for nine Japanese species (all belonging to Aphodius ) on the basis of the results

obtained by rearing under laboratory conditions.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

Table 1. Types of oviposition habits and egg distributions of nine Aphodius species together with

their reproductive periods and female body size. S: desiccated dung surface, U: moist upper half layer

of dung, L: moist lower half layer of dung, M: margin between dung and soil, G: soil under dung.

Type

Species

% of eggs laid in

U L M

Total 0

Repro¬

duction 11

Body

size 0

(mm)

A. brachysomus

23 d

1.2

76.1

22.2

0.4

0.0

1417

spring

7.8 ± 0.58

A. haemor-

rhoidalis

35 e

4.4

26.5

38.9

23.0

7.1

113

spring

4.7 ± 0.23

A. breviusculus

18 e

16.7

40.3

37.5

5.6

0.0

spring

4.6 ± 0.34

A. pratensis

22 e

11.4

48.6

37.1

2.9

0.0

autumn

4.6 ± 0.31

A. sordidus

28 e

0.0

100.0

autumn

6.5 ± 0.43

A. rectus

59 e

0.0

4.9

7.6

87.6

185

spring

5.8 ± 0.42

A. pusillus

38 e

0.0

2.8

8.3

11.1

77.8

spring

3.7 ± 0.28

III

A. elegans

45 d

0.0

100.0

1576

autumn

11.6 ± 0.92

A. haroldianus

0.0

100.0

329

spring

9.4 ± 0.81

a Total number of eggs laid in all glass cages.

b Reproductive period (all species are univoltine in Japan).

c Female body length (mean ± SD, n > 30).

d Number of females separately reared.

e Number of adults reared (sex unknown).

Materials and Methods

The nine Aphodius species listed in Table 1 were reared. Each species was

collected, at the peak of reproductive activity (Yoshida & Katakura 1985), from

cattle dung at pastures in Hokkaido Agricultural Experiment Station (42°59' N,

141°24' E) in Sapporo, northern Japan, approximately 10 km from the laboratory

of Hokkaido University, where the rearing was performed.

Of these species, A. haroldianus Balthasar and A. elegans Allibert have been

known to bury dung and lay eggs near (A. haroldianus) or in (A. elegans) the

buried dung masses (Paik 1968; Hosogi et al. 1979, 1980). Other species are

considered to lay eggs directly in droppings on the ground, or in the soil under

the droppings (Yoshida & Katakura 1985; M. Kiuchi, personal communication).

Beetles were reared in glass cages consisting of two vertical glass plates and a

narrow wood frame, which formed the bottom and the two sides of the cage. One

of the glass plates was fixed to the wood frame, but the other plate was removable

permitting food changes and periodical inspections. The lower half of the cage

was filled with humid sand, and then fresh cattle dung was placed on sand to a

depth equal to one-quarter volume of the cage. The top of each cage was sealed

with cotton cloth and 3 mm mesh nylon net to prevent escape of beetles. Several

rearing cages were placed together in a wood box and were shaded by black sheets

so as to keep them dark except for the top. Two sizes of glass cages were used

according to the body size of beetle species (length x width x height: large cage,

19.0 x 1.4 x 25.0 cm; small cage, 12.5 x 0.6 x 10.0 cm).

The females collected at the peak of reproductive activity were assumed mated

prior to collection (Yoshida & Katakura 1985). Then, a female of each of the

three larger species (A. elegans, A. haroldianus and A. brachysomus Solsky, which

were easily sexed) was released separately and individually into large glass cages.

The other six species were difficult to sex externally, and so one to four adults

(unknown sex) per small cage, or five to ten adults (unknown sex) per large cage,

were released.

1992 YOSHIDA & KATAKURA: APHODIUS OVIPOSITION EVOLUTION

Figure 1. A scheme of oviposition sites of four types of oviposition habits distinguished for nine

Aphodius species. I-IV: type of oviposition habits defined in the text; la, A. brachysomus-, lb, A.

haemorrhoidalis, A. breviusculus, A. pratensis.

The dung and soil in each cage were changed every two or three days, after they

were thoroughly examined to detect the oviposition sites and the number of laid

eggs. In addition, position of laid eggs was traced, when necessary, on the trans¬

parent Saran Wrap®, which was placed on the glass plate. For the two species that

bury dung for larvae, the dung burying process was also observed through the

glass plate.

Species that reproduce in the spring (Table

1) were reared under long day

C, and species that reproduce in

1° C or

conditions (LD 16:8) at 15 ± 1° C or 18 ± 1°

the autumn were reared under short day conditions (LD 12:12) at 18

23 ± 1° C.

Results

Oviposition habits of the nine Aphodius species were classified into four types

according to egg sites and presence or absence of provisioning for larvae (Table

1, Fig. 1).

Type I. —Four species were classified as Type I: A. brachysomus, A. haemor¬

rhoidalis (L.), A. breviusculus (Motschulsky), A. pratensis Nomura & Nakane. Eggs

were laid singly in round or oval spaces in the dung on the soil surface (Fig. 1,

la, lb).

The following is a summary of oviposition habits of A. brachysomus, the most

intensively studied species of this type: Most eggs were laid in the middle part of

cattle dung placed in the rearing cages (Table 1). Almost all eggs were laid in oval

spaces (98.3%, n = 1264); the remainder were laid directly on the surface of

tunnels left behind by the adults’ passage. These spaces (Fig. 1, la) are probably

egg chambers prepared by the mother beetles for oviposition. The spaces were

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

relatively large (6.7 ±1.16 mm long, 4.4 ± 0.66 mm wide; mean ± SD, n = 47)

and their inner wall was smooth, except for one side that was rough and protruded

somewhat inwards into the space. No such space had more than one egg. Each

egg was stood on end at the side opposite to the rough side of the space. It seems

that the mother beetle makes an egg chamber, lays an egg, and closes it with rough

dung fragments.

Eggs of the remaining three species of this type were also found singly in round

or oval spaces, 1.3-2.4 mm diameter, inside of the dung (Fig. 1, lb; Table 1).

Whether these spaces were specially prepared egg chambers or mere spaces left

behind by the adults’ movement was not determined for these smaller species.

Type II. — Three species were classified as Type II: A. sordidus (Fabr.), A. rectus

(Motschulsky), A. pusillus (Herbst). Eggs were laid singly in spaces in the soil

beneath dung (Table 1; Fig. 1, II).

The spaces were simple and round, 1.5-3.0 mm diameter and not coated with

dung. Because these spaces were apart from tunnels that were left behind by the

adults’ passage, they were probably specially prepared egg chambers. Most of the

eggs were deposited at a depth shallower than 30 mm as follows: A. sordidus, 13.5

± 11.28 mm; A. rectus, 11.1 ± 10.14 mm; A. pusillus, 10.3 ± 9.73 mm. Some

eggs of A. rectus and A. pusillus were laid at the boundary between dung and soil,

and a few others were in the lowest part of dung.

Type III. — One species was classified as Type III: A. elegans. Each egg was laid

in a small dung mass stuffed into a shallow burrow in the soil beneath the dung

(Fig. 1, HI).

Dung masses were found shallower than 30 mm deep, and its top usually

connected with the bottom of the above dung. The periphery of buried dung

masses was mixed with grains of soil. Dung masses were 17.3 ± 3.6 mm long

and 9.6 ± 1.6 mm wide (n = 83). There was a space in each mass, in which a

single egg was usually laid (97.0% of masses with one egg, 0.4% with two eggs,

and 2.6% with no egg, n = 533). The space (egg chamber) was 10.5 ± 2.79 mm

long and 5.9 ± 1.37 mm wide (n = 45). The inner surface of the egg chamber

was smooth at the bottom and sides, but rough at the top, sometimes protruding

inwards. The dung wall was thin at the bottom (1.7 ± 1.41 mm) and sides (2.6

±0.92 mm) but thick at the top (4.4 ± 1.85 mm). Most eggs were laid on the

lower one-half of the wall of egg chambers (58.5% of laid eggs; n = 94) or on the

bottom (36.6%). Sometimes a few dung masses were fused together at their sides.

In the provisioning and oviposition processes, a female excavated a shallow

burrow in the ground beneath dung, carried dung fragments from above, and

plastered the wall of the burrow with dung so as to form a chamber. Although

we could not trace subsequent processes, the female must smooth the inner surface,

lay an egg and close it from above with dung fragments, judging from the condition

of buried egg masses.

Type IV. — One species was classified as Type IV: A. haroldianus. Each egg was

laid in the soil beneath the dung; after oviposition, the parental females buried a

dung mass near the egg (Fig. 1, IV).

Buried dung masses were found in the soil up to 8 cm deep. They were similar

to a sausage and often somewhat curved; 34.8 ± 8.4 mm long and 14.3 ± 3.2

mm wide (n = 106). Dung sausages were made of stratified compact dung and

had no spaces within them. An egg was laid in a space in the soil 5-8 mm apart

from the terminal end of each dung sausage. The space was on the average 6.6

1992 YOSHIDA & KATAKURA: APHODIUS OVIPOSITION EVOLUTION

(IV)

BOP

BPO

(in)

Figure 2. Suggested evolutionary relationships among four types of oviposition habits in Aphodius.

I-IV: type of oviposition habits, O: oviposition, B: burrowing soil, P: provisioning dung into burrow.

mm long, 4.0 mm wide, and without any coating. Two or three masses sometimes

fused together at the sides or at the terminal ends, at least in the narrow rearing

cages.

In the provisioning and oviposition processes, a female excavated a vertical

shaft beneath the dung and constructed an egg chamber near the end of the shaft.

(Then, she probably lays one egg in the chamber and closes the chamber with

soil, but these processes could not be confirmed in this study.) After oviposition,

she filled the shaft with dung, mixed her own excrements with dung, and then

she closed the shaft with soil. Sometimes, however, the dung sausage was not

entirely buried but remained in contact with above unburied dung.

Discussion

Evolutionary Trends among Oviposition Habits. — Although it is yet uncertain

which behavior type is most primitive among the four, Types I and II are evidently

simpler and more primitive than Types III and IV. In Types III and IV, dung is

supplied for the young, but the behavioral sequence of provisioning and ovipo¬

sition is distinctly different between the two species. In Type III, a dung mass is

first prepared in the soil, and then an egg is laid in the dung mass. On the other

hand, in Type IV egg is laid in the soil before dung is buried to form a dung

sausage. Because the sequence of oviposition and provisioning is thus inverted

between the Types III and IV behaviors, it is likely that these two types evolved

independently (Fig. 2).

Type IV behavior seems to have evolved from Type II; in both behaviors eggs

are laid in the soil near (but not in) the larval food resource, and the behavioral

sequence of oviposition habits in Type IV can be easily evolved from the simpler

Type II, by adding provisioning behavior to the behavioral sequence of the latter

(+P).

On the other hand, two alternative interpretations are possible for the evolution

of Type III behavior: (1) this type may have evolved from Type II, by inserting

provisioning behavior (P) between burrowing (B) and oviposition (O); or (2) Type

III behavior may be introduced from Type I, by inserting burrowing and provi¬

sioning to the behavioral sequence of the latter (+BP). The first interpretation

seems more parsimonious, but the second interpretation may be supported by

the fact that in both Type III and Type I behaviors eggs are laid in the larval food

resource.

Parallel Evolution of Dung Burying Habits. — Klemperer (1983) coined the term

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

“rummagers” for the Aphodius species that lay their eggs directly in droppings

(synonymous with endocoprids sensu Hanski 1986, not sensu Bomemissza 1969),

and contrasted them with “buriers” (paracoprids) that excavate burrows and fill

them with dung masses in each of which an egg is laid, and with “rollers” (telocop-

rids) which roll away a ball of dung some distance before being buried in a chamber

where the ball is either eaten or converted to a brood ball. According to this

classification, species with Type I behavior are typical rummagers. Type II be¬

havior species can also be called rummagers because their larvae freely feed on

the dung on the soil surface, although they lay eggs outside of the dung.

On the other hand, species of Types III and IV behavior are buriers, in that

both place dung in the soil for larvae. However, the prepared dung masses are

small in behaviors of both Types III and IV, as described above. Larvae of species

with Type III behavior cannot complete the growth with buried dung masses

(second and third instars eat freely in droppings on the soil; Hosogi et al. 1979;

NY, unpublished data). Larvae of species with Type IV behavior can complete

the growth in the buried dung masses, but often depart from it and eat freely in

unburied dung (third instars; NY, unpublished data). Accordingly, it seems more

appropriate to treat behaviors of Types III and IV as intermediate states between

rummagers and typical buriers completing growth only with buried dung masses.

Anyhow, behaviors of Types III and IV could be regarded as representing two

basic types of oviposition habits found in buriers. Type III is analogous to the

oviposition habits of many species of Scarabaeinae (e.g., the Oniticellini and

Onthophagini; in particular, Oniticellus egregius Klug) and Geotrupinae (e.g.,

Geotrupes spiniger Marsham, G. cavicollis Bates) laying eggs in buried dung masses

(Halffter et al. 1985, Klemperer 1979, Davis 1989). Type IV is essentially the

same as the oviposition habits of some Geotrupinae, typically represented by

Typhoeus typhoeus (L.) (Palmer 1978, Brussaard 1983) and Ceratophyus hoff-

mannseggi Fairmaire (Klemperer 1984), which lay each egg near the terminal end

of a dung sausage buried in the soil.

Diversification of brood caring habits is prominent in two major groups of

scarabaeid beetles, Scarabaeinae and Geotrupinae (Halffter & Matthews 1966,

Halffter & Edmonds 1982). Almost all species of these two groups are presocial,

and even in the most primitive type, adults provide foods for larvae. Ironically,

this makes it difficult to seek the evolutionary origin of their presociality among

these two groups of beetles. For the evolutionary origin of brood caring in dung

beetles to be clarified, it is more preferable to concentrate our effort to the groups

which include both species showing brood caring and those not. The present study

showed that Aphodius beetles are particularly suitable for such purpose. We expect

that closer ethological and ecological studies of Aphodius beetles will thus facilitate

our unbiased understanding of the diverse brood caring habits evolved in coproph-

agous beetles of the family Scarabaeidae.

Acknowledgment

Makoto Kiuchi kindly informed us of his unpublished observations on ovi¬

position habits of some Aphodius species. Terumitsu Miyashita and other mem¬

bers of the staff of the Hokkaido Agricultural Experiment Station made available

the facilities of the Experiment Station. The rearing of Aphodius beetles was carried

out at Center for Experimental Plants and Animals, Hokkaido University.

1992 YOSHIDA & KATAKURA: APHODIUS OVIPOSITION EVOLUTION

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Received 31 January 1991; accepted 20 May 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 8-11, (1992)

SURVEY OF MYZUS PERSICAE (SULZER)

(HOMOPTERA: APHIDIDAE) INFESTATIONS ON

BEDDING PLANTS FOR SALE IN EASTERN IDAHO

Susan E. Halbert and Thomas M. Mowry

Southwest Idaho Research and Extension Center, Department of Plant, Soil and

Entomological Sciences, University of Idaho, Parma, Idaho 83660-6699

Abstract. — A survey of bedding plants commercially available in all potato seed production areas

of eastern Idaho revealed that they remain a potential source of infestation of Myzus ( Necta -

rosiphon ) persicae (Sulzer), the green peach aphid. Cole crops, eggplant, forget-me-not, peppers,

and petunia showed 18, 53, 36, 42, and 41% infestation, respectively. A survey involving 476

green peach aphids showed that none transmitted PLRV to test plants.

Key Words.— Insecta, Aphididae, Myzus persicae, bedding plants, PLRV

Potato leafroll virus (PLRV) can be a major problem in Idaho potato production,

especially for certified potato seed growers. PLRV is spread by vegetative prop¬

agation of infected potato seed or transmission by aphid vectors, the most im¬

portant being Myzus {Nectarosiphon) persicae (Sulzer), the green peach aphid.

Myzus persicae has a broad summer host range, but in Idaho it overwinters

holocyclicly only on peach and apricot trees (Bishop & Guthrie 1964). These

overwintering hosts will not withstand the severe winters in most Idaho potato

seed production areas. Work by Tamaki et al. (1979) suggested that anholocyclic

overwintering may occur in Washington during mild winters, but this is very

unlikely under severe Idaho winter conditions in seed production areas. Prior to

the 1960s, it was assumed that aphid infestations on seed potatoes were initiated

each year by alatae that immigrated from warmer areas; however, subsequent

work indicated that this might not be the case. Bishop (1965) showed tht seed

potato fields closest to towns had the most M. persicae and highest PLRV inci¬

dence. Home gardens in the towns were found to be sources of both virus and

vector. Bishop & Guthrie (1964) strongly implicated bedding plants imported

from surrounding states as the major source of M. persicae and showed that virus

inoculum built up in home grown potatoes as gardeners used their own crop for

seed year after year.

A successful integrated pest management program for potato seed production

involving elimination of winter hosts, distribution of free certified seed potatoes

to home gardeners, and insecticide treatment of bedding plants was implemented

in the Grace area of Idaho in the 1960s (Bishop 1967). More recently, prophylactic

insecticide treatments have supplanted this integrated program. Because of new

restrictions on insecticide use, assessment of the feasibility of integrated pest

management has become attractive. The source of M. persicae in seed production

areas that are too cold for survival of primary hosts is thus a matter of scientific

and applied interest. Though bedding plants have been suspected as a major source

of aphids, there is no published survey objectively documenting frequency of

infested bedding plants in a major potato seed production area. There is also no

published information as to whether or not aphids on bedding plants are virulifer-

1992

HALBERT & MO WRY: MYZUS PE RSI CAE INFESTATIONS

Table 1. Myzus persicae infestation of commercially available bedding plants in eastern Idaho seed

potato production areas. Survey conducted 14-18 May 1990.

City/area a

Plant examined

No. salable

units

examined

No. infested

with GPA

PLRV

transmission 11

American Falls/Aberdeen (2)

eggplant

0/30

pepper

0/30

Arco/Mackey (4)

cole crops

0/30

pepper

0/50

Ashton/St. Anthony (4)

cole crops

0/5

eggplant

—

forget-me-not

—

pepper

0/10

petunia

—

Burley (2)

cole crops

—

eggplant

0/9

pepper

0/17

Idaho Falls (7)

cole crops

0/10

eggplant

0/47

pepper

109

0/37

Pocatello (5)

cole crops

—

eggplant

0/10

pepper

0/32

Resburg (6)

cole crops

0/25

eggplant

—

forget-me-not

—

pepper

0/40

Soda Springs/Grace (3)

cole crops

0/4

eggplant

0/10

pepper

0/20

petunia

—

Teton basin (3)

cole crops

—

forget-me-not

0/30

pepper

0/30

petunia

—

a Number in parentheses is the number of commercial outlets surveyed. The information is pooled

by community/area. Not every business had infested plants.

b Number of aphids transmitting/number tested. Readings on concurrent controls were: aphids with

no access to PLRY, 0/20; no aphids, 0/20; aphids fed on known PLRV source, 12/20.

ous. This survey tried to determine if commercially available bedding plants are

presently a source of M. persicae and if they are a potential source of PLRV

inoculum in Idaho’s major seed production areas.

Methods and Materials

The survey was carried out from 14-18 May 1990, in all major eastern Idaho

potato seed production areas and nearby larger cities (Pocatello, Idaho Falls and

Rexburg). Bedding plants originating from nurseries in Idaho and surrounding

states were inspected in 36 commercial outlets in 14 communities.

Only those plants known to be good hosts of M. persicae were examined: cole

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

Table 2. Myzus persicae infestation of commercially available bedding plants in eastern Idaho seed

production areas summarized by plants. Survey conducted 14-18 May 1990.

Plant

Salable units

No. examined

No. infested

% infested

Cole crops

245

18.4

Eggplant

150

52.7

Forget-me-not

36.4

Pepper

490

204

41.6

Petunia

40.6

Totals

939

349

37.2

crops, eggplants, peppers (green and chili) and certain ornamentals. Results were

scored in terms of salable units rather than individual plants (i.e., if plants were

sold in trays of six plants, the number recorded would be number of trays infested

out of number examined).

Aphids infesting bedding plants might originate on hosts other than those ex¬

amined at commercial outlets allowing for the possibility, however slight, of

previous PLRV acquisition. To examine this possibility, at each location where

infested plants were found, three infested plants were purchased, individually

caged and returned to the laboratory. The aphids were then transferred to Physalis

floridana L. seedlings for a 72 h inoculation access period. After removing the

aphids with an insecticide spray, the plants were held for three weeks in the

greenhouse and observed for PLRV symptoms.

Results and Discussion

Plants infested with M. persicae were found in every community/area surveyed

(Table 1), but not in every store. In all, infested plants were found in 72% of the

outlets surveyed. Percentages varied from 25% in Ashton/St. Anthony to 100%

in American Falls/Aberdeen, Burley and Pocatello. Overall, approximately 42%

of the peppers and 53% of the eggplants were infested with green peach aphids

(Table 2). Forget-me-nots and petunias (mostly very young petunia plants) were

similarly infested, and cole crops (Brussels sprouts, broccoli, cabbage and cauli¬

flower) were less frequently infested. This survey indicates that commercially

available bedding plants remain a major source of M. persicae in the eastern Idaho

seed production areas.

None of the plant species examined in this survey are known hosts of PLRV.

Although these plants provided a significant source of M. persicae, they were not

a source of viruliferous aphids that may have originated on PLRV hosts. Testing

of 476 aphids for PLRV transmission revealed none transmitted the virus to P.

floridana seedlings.

As restrictions on insecticide use increase, there is a need to return to an

integrated approach to aphid management. Success of Bishop’s (1967) pilot pro¬

gram depended, in part, upon control of aphids on bedding plants. It is likely that

interdiction of infested bedding plants, coupled with elimination of volunteer

peach and apricot seedlings and distribution of free certified potato seed to home

gardeners would significantly reduce aphid infestation levels and PLRV incidence

in Idaho seed production areas.

1992 HALBERT & MO WRY: MYZUS PERSICAE INFESTATIONS 11

Blackman (1987) and Blackman & Paterson (1986) have shown that M. persicae,

in its historical sense, is actually a complex of morphologically similar species.

It would be problematic if sibling species in the M. persicae complex have

differential vectoring abilities for PLRV. Future work should be done to assess

the ability of aphids originating on bedding plants to colonize potato and their

efficiency in transmitting PLRV.

Acknowledgment

We thank Joyce Sorrell for technical assistance and the Idaho Potato Com¬

mission for financial support. This is Idaho Agricultural Experiment Station sci¬

entific paper 90762.

Literature Cited

Bishop, G. W. 1965. Green peach aphid distribution and potato leafroll virus occurrence in the seed

potato producing areas of Idaho. J. Econ. Entomol., 58: 150-153.

Bishop, G. W. 1967. A leafroll control program in Idaho’s seed potato areas. Am. Potato J., 44:

305-308.

Bishop, G. W. & J. W. Guthrie. 1964. Home gardens as a source of the green peach aphid and virus

diseases in Idaho. Am. Potato J., 41: 28-34.

Blackman, R. L. 1987. Morphological discrimination of a tobacco-feeding form from Myzus persicae

(Sulzer) (Hemiptera: Aphididae), and a key to New World Myzus {Nectarosiphon) species. Bull.

Ent. Res., 77: 713-730.

Blackman, R. L. & A. J. C. Paterson. 1986. Separation of Myzus (. Nectarosiphon ) antirrhinii (Mac-

chiati) from Myzus (N.) persicae (Sulzer) and related species in Europe (Homoptera: Aphididae).

Syst. Ent., 11: 267-276.

Tamaki, G., L. Fox & B. A. Biott. 1979. Ecology of the green peach aphid as a factor of beet western

yellows virus of sugarbeets. USD A Technical Bulletin, 1599.

Received 4 March 1991; accepted 15 May 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 12-14, (1992)

RATES OF PREDATION BY CHRYSOMYA RUFIFACIES

(MACQUART) ON COCHLIOMYIA MACELLARIA (FABR.)

(DIPTERA: CALLIPHORIDAE) IN THE LABORATORY:

EFFECT OF PREDATOR AND PREY DEVELOPMENT

Jeffrey D. Wells and Bernard Greenberg

Department of Biological Sciences, University of Illinois at Chicago,

Chicago, Illinois 60680

Abstract. — Chrysomya rufifacies (Macquart) is a blow fly that was recently introduced to North

America. Because the larvae of this species are facultative predators on other maggots, native

North American carrion flies probably will be negatively affected by the invasion. Cochliomyia

macellaria (Fabr.), the native calliphorid with the greatest bionomic similarity to the invader,

was selected as the prey species for a laboratory study of predatory behavior. We investigated

the influence of both predator and prey development on predation rates when single predators

and prey were paired in the laboratory. Third instar C. rufifacies consumed third and, at a lesser

rate, second instar C. macellaria. Earlier instars were not predaceous. Both relatively small and

relatively large third instar C. rufifacies consumed the same number of mid-size prey.

Key Words. —Insecta, Cafliphoridae, Chrysomya invasion, predatory behavior, Cochliomyia prey,

effect of development

The Old World blow fly Chrysomya rufifacies (Macquart) was apparently in¬

troduced to Costa Rica around 1978 (Jiron 1979). Since that time it has been

collected in Baja California and Caifomia (Greenberg 1988), Texas (Richard &

Ahrens 1983), and Arizona (Baumgartner 1986). Chrysomya rufifacies is an im¬

portant parasite of newborn calves in extremely wet areas of Hawaii (Shishido &

Hardy 1969) and concern has been expressed about the economic impact of this

species in its new range (Schmidt & Kunz 1985). Chrysomya rufifacies larvae are

facultative predators on other maggots including parasitic species (Fuller 1934).

Because of this habit, and because C. rufifacies is typically a secondary invader

of both carrion and live mammals (Fuller 1934, Norris 1959), the net economic

effect of this fly is often unclear.

Carrion arthropod species display a continuous succession in a carcass (Schoenly

& Reed 1987). Two species occupying the same carcass may avoid particular

interactions simply because the necessary developmental stages do not meet. It

is of interest, then, to know what interactions are possible between various life

stages. Both second and third instar Chrysomya rufifacies have been described as

predaceous (Goodbrod & Goff 1990), but the relative behavior of the different

instars and the size of the prey that can be subdued have not been reported. As

part of a study of the ecology of this fly and its impact on native Diptera, we

investigated the effect of both predator and prey development on C. rufifacies

predation rates in a laboratory setting. The prey species was Cochliomyia ma¬

cellaria (Fabr.), the North American fly with the greatest bionomic similarity to

the invader (Nicholson 1934, James 1947, Hall 1948, Bohart & Gressitt 1951,

Denno & Cothran 1975, unpublished data), and presumably its closest ecological

homolog.

1992

WELLS & GREENBERG: PREDATION BY CHRYSOMYA

Methods and Materials

Experiment 7.—Single C. rufifacies and C. macellaria larvae were confined

together in 55 x 13 mm plastic petri dishes lined with moistened filter paper.

Dishes were placed on a laboratory bench at 23° C with lights on. Larvae from

laboratory colonies had been reared on an excess of ground beef. Treatments were

the nine possible combinations of the three larval instars of each species. Ap¬

proximate body lengths of the larvae used were 2.3, 7.4 and 10.5 mm for first,

second and third instar C. rufifacies, and 2.3, 7.0 and 16.8 mm for first, second,

and third instar C. macellaria. Twenty pairs were created for each treatment. The

dishes were simultaneously arranged in a random pattern within 20 rows and nine

columns. The larvae were constantly scanned for 5 h and instances of successful

predation (C. macellaria consumed) by C. rufifacies were recorded for every hour

(C. rufifacies curls around and pierces its prey which struggles violently in re¬

sponse). Following this, the larvae were left in place with the lights off for 17 h

and again examined for evidence of predation.

Experiment 2.—Larvae were confined as in experiment 1, but in this case we

examined the effect of predator size when third instar C. rufifacies attack third

instar C. macellaria. Predator size was either relatively small (approx. 10.5 mm)

or relatively large (approx. 16.2 mm) matched with one prey size (approx. 12.5

mm). Care was taken that post-feeding larvae were not used for the larger pred¬

ators. Again, 20 dishes for each treatment were set up and arranged at random

within a pattern of 10 rows and four columns. Because the great majority of

predaceous acts in experiment 1 occurred within the first hour (see below), the

larvae were constantly scanned for 1 h and instances of predation were recorded

for each 0.5 h.

Results and Discussion

The numbers of dishes with predation in experiment 1 were 17 of the paired

third instars, seven of third instar C. rufifacies with second instar C. macellaria,

and zero for all other treatments. All acts of predation between third instars

occurred within the first hour, but some from the second treatment occurred in

hours two (two dishes) and three (one dish). After the dishes were left overnight

in darkness a second instar C. rufifacies was observed feeding on a dead second

instar C. macellaria. This may have been either predation or scavenging. In

experiment 2 there was no difference in the number of prey taken by small versus

large C. rufifacies (15 each).

The conditions in this investigation were, of course, highly artificial and might

not represent true predation rates within a carcass. Still, the relative differences

in behavior seen here may exist in the field. Although second instar C. rufifacies

may be predaceous as reported, they were much less so than third instars.

Chrysomya rufifacies typically behaves as a secondary fly in that oviposition

occurs on carcasses already occupied by other larvae (Fuller 1934, Bohart &

Gressitt 1951, Early & Goff 1986). Our results suggest that an ecological refuge

exists for native Diptera that reach the post-feeding stage before third instar C.

rufifacies are present. We have found that all C. rufifacies instars are present in

goat carcasses when the food is exhausted (unpublished data), indicating that no

similar refuge could exist for species following C. rufifacies in succession.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(1)

Literature Cited

Baumgartner, D. L. 1986. The hairy maggot blow fly Chrysomya rufifacies (Macquart) confirmed in

Arizona. J. Entomol. Sci., 21: 130-132.

Bohart, G. E. & J. L. Gressitt. 1951. Filth-inhabiting flies of Guam. Bernice P. Bishop Mus. Bull.,

204.

Denno, R. F. & W. R. Cothran. 1975. Niche relationships of a guild of necrophagous flies. Ann.

Entomol. Soc. Am., 68: 741-754.

Early, M. & M. L. Goff. 1986. Arthropod succession patterns in exposed carrion on the island of

O’ahu, Hawaiian Islands, USA. J. Med. Entomol., 5: 520-531.

Fuller, M. E. 1934. The insect inhabitants of carrion: a study in animal ecology. Counc. Sci. Ind.

Res. Bull. (Aust.), 82: 4-63.

Goodbrod, J. R. & M. L. Goff. 1990. Effects of larval density on the rates of development and

interactions between two species of Chrysomya (Diptera: Calliphoridae) in laboratory culture.

J. Med. Entomol., 27: 338-343.

Greenberg, B. 1988. Chrysomya megacephala (F.) (Diptera: Calliphoridae) collected in North Amer¬

ica and notes on Chrysomya species present in the New World. J. Med. Entomol., 25: 199—

200 .

Hall, D. G. 1948. The blowflies of North America. Thomas Say Foundation.

James, M. T. 1947. The flies that cause myiasis in man. USDA Misc. Pub., 631.

Jiron, L. F. 1979. Sobre moscas califordidas de Costa Rica (Diptera: Cyclorrapha). Brenesia, 16:

221-223.

Nicholson, A. J. 1934. The influence of temperature on the activity of sheep-blowflies. Bull. Entomol.

Res., 25: 85-91.

Norris, K. R. 1959. The ecology of sheep blowflies in Australia. Monogr. Biol., 8: 514-544.

Richard, R. D. & E. H. Ahrens. 1983. New distribution record for the recently introduced blow fly

Chrysomya rufifacies (Macquart) in North America. Southwest Entomol., 8: 216-218.

Schmdit, C. D. & S. E. Kunz. 1985. Reproduction of Chrysomya rufifacies (Macquart) in the lab¬

oratory. Southwest. Entomol., 10: 163-165.

Schoenly, K. & W. Reed. 1987. Dynamics of heterotrophic succession in carrion arthropod assem¬

blages: discrete seres or a continuum of change? Oecologia, 73: 192-202.

Shishido, W. H. & D. E. Hardy. 1969. Myiasis of new-born calves in Hawaii. Proc. Haw. Entomol.

Soc., 20: 435-438.

Received 16 January 1991; accepted 18 May 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 15-26, (1992)

NEST BIOLOGY OF OSMIA (DI CERA TOSMIA)

SURF ASCI AT A CRESSON IN CENTRAL TEXAS

(HYMENOPTERA: MEGACHILIDAE)

John L. Neff 1 and Beryl B. Simpson 12

Central Texas Melittological Institute, 7307 Running Rope,

Austin, Texas 78731

department of Botany, The University of Texas, Austin, Texas 78713

Abstract.— Nests and provisioning behavior of Osmia (Diceratosmia) subfasciata Cresson, a

widely distributed, polylectic, cavity nesting bee were studied in central Texas. Most study nests

were in borings in pine blocks, but field observations suggest snail shells also are used. Nest

plugs and partitions are formed of a mixture of masticated plant material and coarse sand with

both materials collected and mixed on the same foraging trip. Provisioning series consist of a

mix of long and short foraging trips, with six to ten long trips required to provision a cell. Details

of the structure of the four layered cocoon are discussed and figured. Biologically novel features

included the initiation of the outermost cocoon layer while the larva is still feeding with the

margins of this layer being extended along the cell wall as feeding continues. Larvae spend part

of the summer in an extended prepupal diapause before pupating and eclosing to overwinter

within their natal cocoons. Floral pollen and nectar sources are listed and observations on the

biology of Chrysura pacifica (Say) (Chrysididae), a nest parasite, are presented.

Key Words. — Insecta, bee, nest biology, provisioning rate, cocoon, diapause

The subgenus Diceratosmia has been considered the most generalized group of

North American Osmia (Sinha 1958). Although American workers long consid¬

ered Diceratosmia to be a distinct genus, it was reduced to a subgenus of Osmia

by Sinha (1958) in a treatment followed by most modem workers (Mitchell 1962,

Hurd 1979, Michener 1979). Osmia (Diceratosmia) subfasciata Cresson is a wide¬

spread but little studied member of this assemblage with most observations on

its biology consisting only of brief reports of its use of beetle burrows in wood as

nest sites (Linsley 1946, Mitchell 1962), and collection records suggesting its use

of mud wasp nests and plant stems as nest sites (Cockerell 1911). The only report

yielding any details on its nesting biology is a brief note by Krombein (1967: 311-

312) that described a single nest from Arizona, but gave little indication there is

anything distinctive about the biology of this bee. Our observations agree in

general outline with those of Krombein but differ significantly from his obser¬

vations in several aspects of nest biology such as materials used in nest construc¬

tion and duration of prepupal diapause. These differences, coupled with new data

on provisioning behavior and timing of cocoon formation, suggest that O. subfas¬

ciata is biologically more interesting than previously indicated.

Materials and Methods

Observations on foraging behavior and nest construction were conducted from

1979 to 1990 at several sites in central Texas with most of the studies done

between 1986 and 1990 at the Brackenridge Field Laboratory (BFL) of the Uni¬

versity of Texas, Austin, Texas. Artificial pine block trap-nests with diameters of

2.8, 3.2, 4.8 and 6.4 mm were set out at BFL as well as at Sayersville, Bastrop

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

Co., and Charco, Goliad Co. Additional nests with bores of 5.8, 7.9 and 9.5 mm

were also set out at BFL. All 2.8 mm and some 3.2 mm diameter nests had bore

depths of 45 mm. Most 3.2 mm nests and nests of all other diameters were bored

to a depth of 120 mm. Nests were set out in both large, shaded domiciles with

40 to 60 nests per domicile or in smaller exposed clusters of 12 to 16 nests at

BFL. Roughly equal numbers of nests of each diameter were used in each cluster

or domicile. Nests in the large domiciles, but not in the small clusters, were

replaced when empty nests were filled. The domiciles were placed 1 m from the

ground on poles equiped with various sticky traps and moats to exclude fire ants.

Nest clusters were taped or wired to tree limbs at 1 or 2 m above the ground as

well as mounted on stakes at a height of 15 cm above the ground. Four sets of

nest clusters were set out at Charco and Sayersville with two nest clusters on

stakes and two on tree limbs at each site. All nests were aligned with their long

axes horizontal to the ground.

Individual nests, or provisioning females, from BFL were coded by the year

followed by the nest number. Data on phenology and floral hosts were based on

the general collections and observations of JLN in central Texas from 1979 through

1989. Data are presented as the mean ± one standard deviation. Plant nomen¬

clature follows Correll & Johnston (1970) and Johnston (1988). Insect vouchers

are deposited at the Brackenridge Field Laboratory, Austin, Texas and the Snow

Museum, University of Kansas, Lawrence Kansas.

Observations on larval development were based on trap-nests opened in the

laboratory and maintained at room temperature (24° C to 8° C summer and 15°

C to 22° C winter). Torchio (1989) found that in several Osmia spp., the first

larval molt occurs before hatching so that the first evident instar is actually the

second yielding a total of five larval instars. Our methods were inadequate for

detecting if such an early molt occurs in O. subfasciata so our discussion refers

to the number of instars assuming the eclosing larva is the first instar although

we recognize this may be incorrect.

Pollen collection records were based on field observations and analyses of sam¬

ples of scopal loads and nest provisions. Pollen from scopal loads was mounted

in glycerine for microscopic examination and sorted to plant morphospecies with

the aid of a reference pollen collection. Crude pollen volume per morphospecies

was estimated from pollen volume per grain times number of grains per sample

(minimum of200 grains for total sample). As a conservative measure, only pollens

of species constituting 10% or more of the volume of a sample were considered

to have been actively collected because some pollen will be picked up incidental

to nectaring and many flowers may show high levels of “contamination” of foreign

pollen which may be unintentionally collected by a bee.

Results

Phenology and Floral Hosts. — Osmia subfasciata is a strictly vernal species in

central Texas. Collection records suggest the species is weakly protandrous with

females having much greater longevity than males (Fig. 1). We have collected

males from 3 Mar to 14 May in central Texas with most records from the last

half of March, although females were collected from 14 Mar to 19 Jun with most

activity from the last half of March through April. Lifespans of individual bees

are unknown.

1992

NEFF & SIMPSON: NEST BIOLOGY OF OSMIA

1-Mar 2-Mar 1-Apr 2-Apr 1-May 2-May 1-Jun 2-Jun

DATE

Figure 1. Central Texas collections records for O. subfasciata indicating pattern of seasonal abun¬

dance. Males = hatched bars. Females = solid bars. Horizontal axis in 2 week periods.

Females of Osmia subfasciata are clearly polylectic, collecting pollen from a

wide array of plants. We recorded O. subfasciata visiting flowers of 32 genera in

14 families in central Texas (Table 1). Hurd & Michener (1955) listed an additional

three families and nine genera of plants visited by O. subfasciata in Texas. NonTexas

records (Hurd & Michener 1955, Mitchell 1962) bring the total to 18 families

and 48 genera. Osmia subfasciata collected pollen of at least 13 genera in nine

families in central Texas (indicated by the letter P in Table 1). Analysis of pollen

loads and nest contents from trap-nests indicated 54% (14 of 26) of the scopal

loads contained mixtures of pollens (two or more pollens each representing at

least 10% of pollen load volume).

Nest Structure. — Considerable variation in body size is seen in central Texas

O. subfasciata with males usually smaller than females. Males were 7.1 ± 0.8

mm (range 5.8-8.7; n = 25) in length with thoracic widths of 2.2 ± 0.2 mm

(range, 1.8-2.6; n = 25). Females were 8.5 ± 0.8 mm (7.0-9.7; n = 25) in length

with thoracic widths averaging 2.7 ± 0.5 mm (2.2-3.1; n = 25). Females initiated

nests in wooden traps with bore diameters of 3.2 ( n = 4), 4.8 (n = 11) and 6.4

mm ( n = 2). Only the long 120 bores were used by O. subfasciata. Our sample

of nests suggests diameters of 3.2 to 4.8 mm, a size best fitting observed female

cross-sectional area, were the preferred nest diameters for O. subfasciata. Unlike

many taxa collected during trap-nesting programs, O. subfasciata is apparently

not gregarious as we never found more than one female active at a given trap-

nest station.

As is common in Osmia, but not in some trap-nesting Hoplitis, nest walls were

unlined, even in nests with diameters significantly larger than that of the cross-

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

Table 1. Central Texas floral records for Osmia subfasciata.

Plant family

Plant taxon

Visited by

Asteraceae

Chaetopappa bellioides (A. Gray) Shinners

Crepis sp.

Engelmannia pinnatifida Nuttall

Coreopsis nuecensis Heller

Coreopsis basalis var. wrightii (A. Gray) Blake

Erigeron sp.

Gaillardia pulchella Fougeroux

Hymenoxys scaposa (DC) Parker

9P, 3

Krigia occidentalis Nuttall

Lindheimera texana Gray & Engelmann

Berberidaceae

Berberis swaseyi Buckley

Brassicaceae

Lesquerella grandiflora (Hooker) Watson

Lesquerella argyraea (A. Gray) Watson

Cactaceae

Opuntia macrorhiza Engelmann

Commelinaceae

Tinantia anomala (Torrey) Clarke

Fabaceae

Cercis canadensis L.

9,3

Desmanthus velutinus Scheele

Lupinus texensis Hooker

Prosopis glandulosa Torrey

9P, 3

Vicia villosa Roth

Hydrophyllaceae

Nama hispidum A. Gray

Nemophila phacelioides Nuttall

9P, 3

Phacelia congesta Hooker

Phacelia patuliflora (Engelmann & A. Gray) A. Gray

Lamiaceae

Brazoria sp.

Monarda citriodora Cervantes

Teucrium cubense Jacquin

9,3

Malvaceae

Callirhoe leiocarpa Martin

Callirhoe involucrata (Torrey) A. Gray

9P, 3

Sida abutifolia Miller

Sphaeralcea lindheimeri A. Gray

Onagraceae

Oenothera speciosa Nuttall

Rubiaceae

Hedyotis nigricans (Lamark) Fosbery

Sapindaceae

Ungnadia speciosa Endlicher

Solanaceae

Chamaesaracha sordida (Dunal) A. Gray

Verbenaceae

Verbena bipinnatifida Nuttall

Verbena officinalis L. ssp. halei (Small) Barber

9,3

1 P indicates known pollen host.

sectional diameter of the bee. Partitions, basal plugs and entrance plugs were

constructed of a mixture of coarse sand and masticated plant material (Figs. 2,

3). The entrance plug was single layered, 3-6 mm thick, and had a smooth, concave

outer surface. The margins of the plug were usually flush with, or more rarely

recessed 3-6 mm from, the nest entrance. The entrance plug was followed by one

or two vestibular cells of 6-47 mm in length. The partition between the vestibular

cells, or the outermost cell partition, if only one vestibular cell was present, was

consistently thicker than the inner brood cell partitions and sometimes thicker

than the entrance plug. Partitions between inner cells were quite thin medially

(0.2-0.3 mm) but thicker along the cell wall (0.5-1.0 mm). In one case, a female

was found to have used a nest previously occupied by a eumenid wasp and had

1992

NEFF & SIMPSON: NEST BIOLOGY OF OSMIA

Figure 2. Female O. subfasciata adding soil to masticated leaf material (7 x).

used portions of some of the remaining mud partitions as bases for her own nest

partitions.

Nests consisted of linear series of 2-12 brood cells. These cells varied from

5.5-12.0 mm in length or 60.0 to 322.0 mm 3 in volume depending on nest

diameter. Lengths of brood cells averaged 8.4 ± 0.6 mm (7.5-9.7; n = 21) in

nests with diameters of 3.2 mm and 7.9 ± 1.3 mm (5.5-12.0, n = 52) and 7.9 ±

1.4 mm (6.5-12.0, n— 11) for 4.8 and 6.4 mm diameter nests respectively. No

consistent pattern was evident between brood cell length and its position within

a nest. Too many bees escaped from nests without being sexed or were detroyed

by parasites to allow accurate estimates of sex ratio or position of sexes within

nests. Limited available data suggest that, as is common in sexually dimorphic,

trap-nesting species with large females, females are produced primarily in the

innermost cells. Only males were produced from the two 3.2 mm diameter nests.

The provision mass was firm and uniformly moist (except for dry loose pollen

along the walls), filling most of the cell. The outer face of the provision mass

slanted towards the cell entrance, the upper part of the mass being the furthest

from the entrance. Females inserted the posterior end of the egg in a moistened

area on the upper third of the slanted face with the anterior portion of the egg

remaining free but curving down over the provisions.

In addition to trap-nests, we observed females of O. subfasciata investigating

THE PAN-PACIFIC ENTOMOLOGIST Vol. 68(1)

Figures 3-4. O. subfasciata cell and cocoon structure. Figure 3. Cross-section of anterior portion

of cocoon with attached nest partition (12.2x). (a) partition of coarse sand and masticated plant

material, (b) outer cocoon layer adhering to partition, (c) outer sheets of inner cocoon, (d) fibrous layer

of nipple, (e) middle layer of inner cocoon, (f) innermost layer of inner cocoon. Figure 4. Cross-section

of cocoon nipple (49 x). (a) elaboration of middle layer of inner cocoon, (b) fibrous inner layer of

nipple, (c) innermost layer of cocoon.

various small, empty snail shells at BFL where a single-celled bee nest was found

in one of these shells by A. Hook (personal communication). Although the nest

was destroyed when opening the shell, the plug was of the characteristic sand/

plant mastic mix. This, along with its small size, suggests it was a Disceratosmia

nest. Osmia subfasciata is the only Diceratosmia known to occur at BFL, and

other small megachilines occurring there use different materials in their nest

partitions. It thus seems likely that O. subfasciata, like Osmia conjuncta Cresson

(Rau 1937), occasionally constructs nests in snail shells.

Nest Construction. — Nests were initiated by placing a thin layer of the sand/

masticated plant material mix at the end of the burrow. Sand and plant material

were collected on the same trip. At Sayersville, we repeatedly observed female O.

subfasciata chewing leaf margins of Helianthemum georgianum Chapman (Cis-

taceae), forming a small ball which was held beneath the mandibles. The bee then

moved to an area of loose sand and dropped the ball of chewed leaf material on

the sand surface where she proceeded to chew and knead the ball as she rolled it

over the soil surface incorporating soil/sand particles into the mass (Fig. 2). In¬

dividuals were observed to return to the same small one or two m 2 areas for five

or more consecutive leaf-soil loads. Source of the plant material at other sites is

unknown but presumably a variety of plant taxa is utilized. Upon returning to

the nest, the female laid down a low rim of the soil/leaf masticate mix which

served as the base of the partition separating the first two cells of the nest. Con¬

struction of such a rim, known as Fabre’s threshold (Malyshev 1936) is widespread

among megachiline bees (Frohlich 1983; JLN, personal observation). After pro¬

visioning and ovipositing in the first cell, the female closed the cell delimited by

the initial rim and constructed another rim. This pattern continued until the final

cell of the nest was completed.

Bee 87-7 required 11 trips and 97.7 minutes to finish a partition and construct

a new rim. During this period she averaged 6.3 ± 1.73 min (2.82-9.63; n= 11)

in the nest and 2.58 ± 2.34 min (0.76-8.78; n = 11) away from the nest. Female

O. subfasciata apparently are able to construct and provision two cells per day.

1992

NEFF & SIMPSON: NEST BIOLOGY OF OSMIA

The female observed in 1989 at BFL took six days to complete a 12 cell nest (89-

7) and then six days for a nine cell nest (89-18).

Provisioning Behavior. —Because females in the small 3.2 and 4.8 mm diameter

nests had to exit the nest to turn around to deposit pollen, time in the nest after

pollen trips could be separated into nectar and pollen deposition components.

Female 89-7 averaged 0.76 ±0.19 min (« = 11) in the nest for nectar deposition

and 1.00 ± 0.30 {n = 11) for pollen deposition. Female 87-7 was somewhat faster,

averaging 0.31 ±0.14 {n = 9) for nectar deposition and 0.71 ± 0.21 for pollen

deposition. The preceding times for pollen deposition exclude the long final period

of a provisioning sequence which presumably represents final pollen mass prep¬

aration and oviposition. This final period was 5.35 min for bee 87-7, and 4.45

min for bee 89-7.

Interpretation of foraging patterns was complicated by the lack of consistency

in the duration of pollen trips. A complete provisioning series for a male cell in

nest 89-7, provisioned on 19 Apr 1989 required 192 minutes. This entailed 10

or possibly 13 pollen trips. The variable estimate stems from the fact that most

pollen trips were relatively long, averaging 16.05 ± 2.78 min (12.04-22.29; n =

9) but there were also three short trips averaging only 2.74 ± 1.22 min (2.02-

4.15; n = 3) when the bee returned with pollen. It is likely the short trips were

not true pollen trips but we cannot be sure as we were not able to tell if the bee’s

scopa still contained pollen when she exited the nest. The female behaved as if

she were depositing pollen prior to leaving for the apparent short pollen trips.

Another cell provisioned on 1 Apr 1987 showed the same pattern. This cell

required only 88 minutes and six long trips with a mean of 15.07 ± 4.66 min

(6.12-19.90) and four short ones with a mean of 1.46 ± 0.85 min (0.89-2.71).

A similar mix of long and short trips was observed in other partial provisioning

series noted in 1987, 1989, and 1990. The short trips may have been nectar trips

or perhaps represent a form of defensive behavior against nest parasites which

oviposit in open cells.

Development and Cocoon Structure. —Larvae fed without moving from the

initial position of egg insertion during the first three instars. The conspicuously

setose fourth instar occurs six to seven days after hatching and then begins moving

over the provision mass. Defecation is initiated one to two days after molting to

the fourth instar. The pale, flattened fecal pellets are initially placed on the distal

portions of the cell, particularly the distal walls and outer margins of the cap.

Individual pellets free from the wall are curved, have tapered ends, and are 0.6-

0.7 mm long. The curved, ventral surface of an individual pellet had a weakly

defined, shallow, longitudinal groove. Pellets produced later become strongly flat¬

tened, even ribbon-like, as they are closely appressed to the cell wall or outer

layer of the cocoon.

The cocoon consists of four layers in two distinct structures, an inner and outer

cocoon (Fig. 3). The outer cocoon is single-layered and is initiated two to three

days after molting to the final larval instar, well before the completion of feeding

as two-thirds to three-fourths of the provision mass remains at that time. The

initial portion of the outer cocoon layer consists of a tough translucent membrane

placed over, and closely adhering to, the cell cap and anterior portions of the cell

wall up to the edges of the remaining provisions. This portion of the cocoon is

laid over the initial layer of fecal pellets. The outer cocoon is extended along the

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

cell walls, sometimes after the deposition of more fecal pellets, as the walls are

exposed by continued feeding. Additional fecal pellets are deposited on the inner

surface of this cocoon layer. The provisions are completely consumed roughly

four to five days after the initiation of the outer cocoon. The outer cocoon is

completed by covering the base of the cell after the completion of feeding.

The inner cocoon, initiated after the completion of the outer, required two or

three days to construct and consisted of three layers. The outermost is formed by

a series of thin, translucent sheets with a few embedded threads. These sheets are

apparently laid down as overlapping series which are attached both to one another

and to the outer cocoon. The innermost of these translucent sheets forms the

foundation for the relatively thick (0.02 mm), tough, brown, opaque middle layer.

Closely appressed to the inner surface of the middle layer is a thin, translucent

inner layer.

The anterior end of the inner cocoon is elaborated into a nipple (Figs. 3, 4), a

complex structure which apparently serves as an air exchange mechanism for the

otherwise impervious cocoon. Seen from within, the innermost portion of the

nipple appears as an opaque, smooth brown disk surrounded by a paler ring. In

cross section, it can be seen that the disk is formed by a tightly packed region of

the tough brown fibers which fill the mesal, hemispherical region of the nipple.

The thick layer of tough threads is covered by, and grades into, the thickened

continuation of the opaque middle layer of the cocoon. This thickened opaque

layer is not continuous, having a central opening of approximately 0.3 mm. This

central opening is filled with a continuation of the inner layer of coarse threads

which in turn intergrades with dense translucent sheets covering the outer surface

of the nipple.

Osmia subfasciata is univoltine in central Texas. After the completion of cocoon

spinning, there is a relatively long larval diapause of approximately 90 to 110

days before pupation. Larvae completing cocoon construction by 6 May pupated

by 5 Aug when kept in the lab at room temperature (23.3°-27.7° C). However

larvae from a nest constructed at the same time but left in the field did not pupate

until 30 Aug. Individuals overwinter as adults within their natal cocoons.

Mating Behavior.— Mating was rarely observed although males are commonly

observed patrolling female pollen and nectar sources such as flowers of Cercis,

Nemophila or Berberis during the initial portions of the flight season. Mating,

including a period of post-copulatory mate guarding, was prolonged and was

observed only at flowers. Males were never observed at nest sites or areas of

female emergence when occupied trap nests were placed in the field. It is likely

that female O. subfasciata mate only once, or at most a few times, during a brief

receptive period.

Predators and Parasites. — Nest parasitism was not extensive with only two

species of nest parasites reared. Bombyliid larvae were reared from two outer cells

of BFL trap-nests. Pupal morphology indicated they were Anthracinae, presum¬

ably Anthrax, but neither individual was able to pierce the nest closure to exit

and both failed to eclose. The other nest parasite was the small chrysid wasp,

Chrysura paciftca (Say) which was reared from nests from Sayersville set out in

1986 and 1989. Chrysurapacifica has not been collected at BFL. Our observations

of C. pacifica larval development agreed with earlier observations of C. pacifica

attacking Osmia pumila Cresson (Krombein 1967: 446). The four-celled Sayers-

1992

NEFF & SIMPSON: NEST BIOLOGY OF OSMIA

ville 89 nest included six C. pacifica: two cells with two C. pacifica, and two with

one. The eggs have a thick, tough chorion and are deposited near the basal portion

of the provisions. In a cell where larval development was followed completely,

the C. pacifica egg hatched one day after the O. subfasciata egg (three to four days

after O. subfasciata oviposition). The C. pacifica larva remained quiescent for

three days before beginning to move forward over the provision mass, aided by

an unusual bifurcate caudal segment. After two days of movement it reached and

attached itself with its mandibles to the still sessile, third instar O. subfasciata

larva. A second C. pacifica larva, previously hidden on the opposite side of the

cell, attached itself to the O. subfasciata larva on the following day. The O.

subfasciata larva molted to the fourth instar the following day (seven days after

hatching) and both larvae were detached, with only one reattaching to the now

setose, free moving O. subfasciata larva. The remaining first instar C. pacifica

larva remained inactive but attached to the O. subfasciata larva for an additional

12 days, during which the O. subfasciata larva completed constructing its cocoon.

It then began feeding on the O. subfasciata larva and its own growth became

obvious. Feeding continued for five days, after which the O. subfasciata larva was

badly shriveled. Unfortunately, the C. pacifica larva was killed by a laboratory

infestation of Chaetodactylus mites which almost certainly came from infested

nests of Osmia ribifloris Cockerell stored nearby. Mites developed from egg to

adult on provisions and feces of O. subfasciata but no infestations were observed

in field collected nests nor were mites present on adults.

Miltogrammine flies were common at the BFL nest site but apparently were

associated with various sphecid and eumenid wasps using the domicile as none

were observed at bee nest entrances. The surface of the pollen mass from one cell

in nest 89-18 in which the egg failed to hatch was encrusted with the black fruiting

bodies of Ascosphaera. The fungus was not noted in other cells. The only obser¬

vation of predation was a female captured by the common red assassin bug,

Apiomerus spissipes (Say), at flowers of Verbena officinalis Linnaeus ssp. halei

(Small) Barber, at Pedemales Falls State Park, Blanco Co., Texas.

Discussion

Our observations on nest structure and cocoon structure of O. subfasciata agree

with those of the only previous study by Krombein (1967) in many features of

general nest architecture but differ in a number of important aspects. Krombein

reported that partitions in his nest from Scottsdale, Arizona, were constructed

only of masticated leaf material rather than the sand/plant mix we found in Texas.

Use of a combination of sand or soil and masticated leaf material for nest con¬

struction is common in the Hoplitis-Anthocopa-Proteriades complex, particularly

in the subgenera Penteriades, Hoplitina, Xerosmia, Acrosmia, Atoposmia, Ere-

mosmia and Hexosmia (Parker 1975, 1978a, b). However, it apparently is un¬

common in Osmia (sensu Sinha 1958). Published accounts indicate the vast

majority of Osmia species use either soil or plant masticate in nest construction

but not both (Iwata 1976, Rust 1974). Exceptions among American species include

Osmia ( Cephalosmia ) californica Cresson, which constructs nest plugs and par¬

tition of a mix of mud and masticated plant material (Levin 1966, Rust 1974,

Torchio 1989), and Osmia (Trichinosmia) latisulcata Michener, which uses a

sand/plant masticate mix for cell partitions and plant masticate and pebbles for

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68 ( 1 )

nest plugs (Parker 1984). Both mud and plant masticate are used in constructing

the um-like cells of some members of subgenus Acanthosmioides (Rust et al.

1974). The use of a sand/plant masticate mix in nest construction appears to be

unusual even within Diceratosmia. Nest construction materials in various species

of Diceratosmia have been described as leaf paste ( O. conjuncta [Rau 1937]), leaf

pulp ( Osmia gallarum Spinola [Iwata 1976]), and green putty ( Osmia versicolor

Latreille [Fabre 1915]). If Diceratosmia is actually the sister group of all other

Osmia (sensu Sinha 1958), it is tempting to suggest that the sand/plant pulp

mixture is the primitive nest construction material for this lineage, but the oc¬

currence of mud, plant masticate, or various mixes as nest construction materials

by taxa scattered throughout the genus renders this highly speculative.

Initiation of cocon spinning before the completion of feeding also appears to

be unusual among megachilines. Although production of a few threads, which aid

in holding the feces in place prior to cocoon completion, is common among

megachilines (Stephen et al. 1969, Torchio 1989), production of a distinct sheet

as produced by O. subfasciata is not. Unfortunately, there are relatively few studies

of development which would allow one to ascertain accurately when cocoon¬

spinning is initiated. The only other megachiline we are aware of that regularly

initiates cocoon construction well before completing feeding is Hoplitis ( Robert-

sonella) simplex (Cresson) (JLN, personal observation).

There is considerable variation in cocoon structure within Osmia. The cocoon

of O. subfasciata is a distinctive combination of elements found elsewhere in the

genus. The tough, nippled, multilayered cocoons of Osmia s. str. (Rust 1974; JLN,

personal observation) appear to be virtually identical to the inner cocoons of O.

subfasciata. The presence of a well formed anterior layer of the outer cocoon is

apparently more unusual in Osmia although a very similar structure is present in

cocoons of O. latisulcata (Parker 1984). A well defined anterior portion of the

outer cocoon is present in at least some species of Hoplitis and Chelostoma (Parker

1988; JLN, personal observation).

The extended prepupal diapause we observed in O. subfasciata also appears to

be unusual among megachilines which overwinter as adults. A review of published

reports suggests most Osmia larvae pupate within thirty days of cocoon comple¬

tion. Significant exceptions are the two year individuals of parsivoltine species,

which typically spend their first winter as prepupae (Torchio & Tepedino 1982)

and larvae of Osmia nigrifrons Cresson that have prepupal diapause lasting 130—

150 days (Rust et al. 1974). In addition, we have found that central Texas pop¬

ulations we studied of Osmia ribifloris have an extended prepupal diapause of

100-150 days (unpublished data) even though populations from northern Nevada

have a brief prepupal diapause (Rust 1986). Although we are unaware of conclusive

proof, it is generally believed that the prepupal stage of bees has lower metabolic

requirements and is more resistant to environmental stress than the adult. Pre¬

sumably this explains why the prepupal stage is the one most commonly used for

overwintering or extended diapause.

Nonetheless, overwintering as an adult is common among cool-temperate bees

which emerge in the early spring (Stephen et al. 1969). Overwintering as an adult

presumably facilitates early emergence and avoidance of pupation under cold or

otherwise unsuitable conditions. However, overwintering as an adult by vernal

bees may regularly expose diapausing adults to potentially lethal temperatures

1992

NEFF & SIMPSON: NEST BIOLOGY OF OS MIA

and/or moisture stress in areas of high summer temperatures. A recent study has

shown that diapausing adult Osmia cornifrons Radoszkowski kept at 22° C had

a survivorship rate (76%) nearly twice that of individuals maintained at 30° C

(39.9%) (Maeta 1978). Excluding Osmia, most osmiine bees overwinter as pre¬

pupae so it is possible that the extended prepupal diapause we observed in O.

subfasciata simply reflects retention of the plesiomorphic state. However, we

believe that in Osmia, extended prepupal diapause is a derived condition facili¬

tating tolerance of extended hot conditions, particularly those with high night

temperatures such as are commonly encountered during central Texas summers.

Average minimum monthly temperature exceeds 18° C from May through Sep¬

tember in central Texas (Conway & Liston 1974). We expect extended prepupal

diapause will be found in other Osmia spp. of central Texas as well as in other

regions with high night temperatures. Osmia subfasciata larvae from Scottsdale,

Arizona reared in a Washington, D.C. laboratory had a prepupal diapause of

roughly 45 days (Krombein 1967). The pattern of overwintering as an adult during

the winter before spring emergence, appears to be universal in Osmia and may

be an important factor limiting the southern distribution of the genus. Two year

old individuals in parsivoltine species of Osmia spend their first winter as pre¬

pupae, but their second as adults (Torchio & Tepedino, 1982). These cases rep¬

resent an extreme form of extended prepupal diapause for some individuals.

Extended prepupal diapause appears to be a mechanism which minimizes ex¬

posure of diapausing adults to the temperature and moisture stress in warm climes

yet retains the flexibility for early spring emergence.

Acknowledgment

We thank Gregg Dieringer for assistance with SEM work, and the Texas De¬

partment of Parks and Wildlife for permission to collect at Pedemales State Park

(permits 17-89 and 10-90). Gregg Dieringer also read the manuscript and made

many helpful suggestions.

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U.S. Nat. Mus., 40: 241-264.

Conway, H. M. & L. L. Liston (eds.). 1974. The weather handbook (rev. ed). Conway Research Inc.,

Atlanta, Georgia.

Correll, D. S. & M. C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research

Foundation, Renner, Texas.

Fabre, J. H. 1915. Bramble-bees and others. Dodd, Mead and Company, New York.

Frohlich, D. R. 1983. On the nesting biology of Osmia ( Chenosmia ) bruneri (Hymenoptera: Mega-

chilidae). J. Kansas. Entomol. Soc., 56: 123-130.

Hurd, P. D., Jr. 1979. Superfamily Apoidea. pp. 1741-2209. In K. V. Krombein, P. D. Hurd, Jr.,

D. R. Smith & B. D. Burks (eds.). Catalog of Hymenoptera in America north of Mexico, Vol.

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Co. Pvt. Ltd., New Delhi.

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Received 18 January 1991; accepted 1 June 1990.

PAN-PACIFIC ENTOMOLOGIST

68(1): 27-37, (1992)

OBITUARY:

JAMES WILSON TILDEN (1904-1988)

Paul H. Arnaud, Jr.

Department of Entomology, California Academy of Sciences,

San Francisco, California 94118

James Wilson “Bill” Tilden, entomologist, lepidopterist, naturalist, professor

emeritus at San Jose State University, died on 27 Dec 1988, four days before his

84th birthday, in San Jose, California, of injuries resulting from a fall at his home.

Memorial services were held at the Methodist Church in Philo, California, on 27

May 1989, and his ashes were buried at Philo in the Ruddock family cemetery.

Bill Tilden is known to many through his research and publications on Lepi-

doptera and ecology, including the books Butterflies of the San Francisco Bay

Region (Tilden 1965), A Field Guide to Western Butterflies (Tilden & Smith 1986),

and California Butterflies (Garth & Tilden 1986); through teaching for 22 years

at San Jose State University; and through his membership in many societies,

including the Pacific Coast Entomological Society. He was a member of this society

for 49 years—elected to membership at the 158th meeting on 30 Sep 1939—and

served as president in 1960. Several articles have been published on Bill’s life

(Anonymous 1989; Heifer 1989; Moreno 1989; Opler & Smith 1990; Smith 1990a,

b), and this article complements information given in the others.

Bill Tilden was bom 31 Dec 1904 in a shake cabin constructed by his father

in the hills above Philo and Anderson Valley, in Mendocino County, California.

He was the second of four children (he had an older sister and two younger

brothers) of Thomas Jefferson Tilden (6 Jul 1871-3 Jun 1945) and Charlotte

Almira Tilden, nee Ruddock (13 Apr 1887-7 Apr 1966).

Bill’s grandfather, James Wilson Tilden (17 Sep 1827-2 Nov 1878), a mate on

a sailing ship, jumped ship in San Francisco to join in the Gold Rush. His

grandmother, Susan Dickerson Tilden (26 Jan 1844-6 Mar 1875), died at the

early age of 31, leaving her husband with three small children to raise. Upon the

death of Bill’s grandfather, three years later in a boating accident in Sacramento,

the orphaned children were supported by the Independent Order of the Odd

Fellows.

Bill’s father, a lumberman and carpenter, was bom in Shingle Springs, in the

foothills of El Dorado County, California. He ran away from his foster home

when he was 14 and supported himself thereafter by his own resources. He worked

near Philo lumbering in the redwood forests with oxen teams. It was here that he

met Bill’s mother, the youngest daughter of Albert G. Ruddock (1839-1895), and

Permelia Curtis Ruddock (1846-1932). Albert Ruddock was a road supervisor

and the first postmaster of Philo. Bill’s father and mother were engaged to be

married for seven years while his father acquired the land and built his cabin.

They were married on 15 Aug 1900.

Bill’s family moved from Philo in 1906, following the death, from diphtheria,

of his sister, Naomi Alice (1902-1905), just before her fourth birthday. This move

was not financially advantageous to the family but was necessitated by the grief

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Vol. 68(1)

Figures 1-5. Figure 1. Thomas Jefferson Tilden, ca. 1900. Figure 2. Charlotte Ruddock Tilden,

ca. 1900. Figure 3. Bill Tilden ten years old with two brothers—Thomas C. Tilden, four years old,

and Earl R. Tilden, two years old. Figure 4. Bill Tilden, high school graduation picture, 1922. Figure

5. Bill Tilden, San Jose State College graduation picture, 1942.

of his mother, who wanted to move elsewhere because of the loss of her daughter,

and concern over Bill’s poor health.

Bill attended public schools in Philo, Turlock, Fresno, and Hilmar. At Hilmar,

a small town located south of Turlock in Merced County, the family lived on a

20 acre ranch between Tegner and Hilmar. Bill graduated from Hilmar High

School in 1922, completing his high school requirements in three years. Bill

wanted, at the time of his graduation, to attend the University of California at

Berkeley and to major in English Literature, but no funds were available for him

to do so.

In 1923, his father purchased 86 acres of second growth redwoods in Santa

Cruz, at Route 1, box 710 (now 3363 Branciforte Drive). About 10 acres were

available to grow garden produce and to plant an orchard. In Santa Cruz, his

father also worked as a carpenter. Bill was to make the family home his head¬

quarters for the next 16 years, whenever he was not working elsewhere.

1992

ARNAUD: JAMES WILSON TILDEN

Figures 6-11. Figure 6. Thomas and Charlotte Tilden, at Santa Cruz ranch, 1942. Figure 7. Bill

Tilden and Hazel Irene Miller Tilden, wedding day, San Francisco, California, 19 Jun 1943. Figure

8. Field day, 200th meeting of the Pacific Coast Entomological Society, Rock City, Mount Diablo,

California, 18 Apr 1948. Left to right, John R. (Jack) Walker, Bill Tilden, John P. Harville, and

Richard M. Bohart (photo by Edgar A. Smith). Figure 9. Vector Control Staff of Santa Clara County

Health Department: back row, left to right, Edgar Smith, Robert Cunningham, Roy Eastwood, Tal

Lloyd, Dean Ecke, and James St. Germaine; front row, left to right, Bill Tilden, Tom Sexton, Jerry

Kraft, Bruce Eldridge, and Rocci Pisano, 1954. Figure 10. Bill Tilden with children (left to right)

James, Janice and Bruce, Christmas picture, 1954 (photo by Hazel I. Tilden). Figure 11. Bill Tilden,

while on Vector Control Staff of Santa Clara County Health Department, 1954.

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Bill’s employment at this time included working for the Southern Pacific Rail¬

road Company in San Francisco, in canneries, and as a fruit tramp with his

brothers, Tom and Earl. While following the fruit season from California to Idaho,

they managed a side trip to see Yellowstone National Park. Bill belonged to the

Musician’s Union and played trombone in a band on the Santa Cruz Boardwalk.

That also included several engagements with The Miss America pageants, and

playing in San Francisco and elsewhere. He was proud of his association with

jazz groups and he later reminisced of participating in a jam session after a Santa

Cruz concert with Jack Teagarden, the famed trombonist. With his brother, Tom,

he collected insects of all orders for L. M. McQuesten of the National Insect

Company, of Davis, California. On one occasion, McQuesten loaned Bill and

Tom his car so that they could collect in the Sierra Nevada for him. Tom recalls

they returned by way of Madera, where they also collected insects from dead

sheep. McQuesten also loaned them a 410-gauge sawed-off shotgun, so that they

could shoot wild game for food. As Tom has commented, “These were the de¬

pression years and employment was hard to come by.”

Bill became interested in natural history at an early age. He started an insect

collection when seven years old. He also enjoyed drawing birds with crayon, and

an aunt subscribed to Bird Lore for him. As was the custom for those who lived

on farms at that time, he shot robins and squirrels for food. His early correspon¬

dents included the entomological promoter, James Sinclair of San Diego, the then

Kansas lepidopterists William D. Field and Virgil F. Calkins, and the twin brothers

Arthur C. and Edgar A. Smith, of Los Banos, California, in 1933. His first exchange

was made with Hugh Gibbon of Miniota, Manitoba, and early exchanges were

made also with Lionel Paul Grey. Bill told me that in 1927 he seriously began to

study Lepidoptera, aided by the Santa Cruz resident lepidopterists Edgar A. Dodge

and John P. Strohbeen. In his first publication, “Preliminary list of the butterflies

and skippers of Santa Clara and Santa Cruz counties, California” (Tilden 1940),

Bill acknowledged the help of Dodge, Strohbeen, Art and Ed Smith, and George

S. Mansfield. Between 1940 and 1987, Bill published over 100 articles (Smith

1990b) on Lepidoptera, Coleoptera, life histories, ecology, flies of public health

importance, etc. Some of these he wrote with co-authors, including John C. Down¬

ey, Carl D. Duncan, John S. Garth, James St. Germaine, David H. Huntzinger,

George S. Mansfield, William A. Palmer, Ernest R. Schoening, Arthur C. Smith,

and Bruce A. Tilden.

In 1938, 16 years after his graduation from Hilmar High School, and when 33

years of age, Bill enrolled at San Jose State College. This was made possible, in

part, by financial support from a friend, Charles “Chas” Bowles, a retired civil

engineer who lived along Indian Creek near Philo. He was a general naturalist

who had an interest in birds. Bill had an avid interest in birds as well as insects,

and it was evident to Bowles and others how knowledgeable Bill was, in spite of

the fact that he did not have a college education. Bowles told Bill that “you simply

have to go to college.” Bill helped Bowles by being the driver on automobile trips,

including one to the deserts of southern California and Arizona. Bowles, in turn,

placed some funds in a bank account to get Bill started in college. In his freshman

year at San Jose State College, Bill roomed in an apartment at 72 South 6th Street

with his earlier entomological correspondents Art and Ed Smith, who were then

in their senior year. It was Art and Ed Smith who earlier made arrangements for

1992

ARNAUD: JAMES WILSON TILDEN

Bill to meet Carl D. Duncan on the annual field day of the Pacific Coast Ento¬

mological Society that was held at Alum Rock Park, San Jose, on 8 Apr 1938.

That was the 152nd meeting of the Society, during which year Duncan was

president. On the basis of this meeting, Bill decided to attend San Jose State

College.

Bill graduated in 1942 from San Jose State College, receiving a Bachelor of

Science degree in Biological Sciences. This was at the height of the Second World

War, and he had applied for deferment so that he might attend medical school

at Stanford University. Because he did not receive word on his request, he decided

to enlist in the U.S. Navy in August 1942, rather than be drafted into the Army.

(Bill later received his letter of deferment when he was overseas!) After six weeks

of basic training on Treasure Island in San Francisco Bay, he served as a Phar¬

macist’s Mate on the troop evacuation ship “Bloemfontaine,” in the Pacific The¬

ater, until V-J Day in 1945. His long-time friend Art Smith, who served with an

Air Force intelligence unit attached to Admiral Nimitz’s headquarters (CincPac),

recalls how he was usually able to meet and visit with Bill, whenever his ship

docked at Pearl Harbor. Bill did some entomological collecting during his military

service, but this was hampered by travel restrictions.

Bill married Hazel Irene Miller in San Francisco on 19 Jun 1943, at which time

she was teaching elementary school at Tupman, in Kern County, California. Later,

Hazel taught in Redwood City in order to be close to Stanford University when

Bill was enrolled there. They first met, in their junior year at San Jose State

College, when in an ornithology class given by Gayle Pickwell. Bill and Hazel

were two of three students who made perfect scores on one of Pickwell’s exam¬

inations.

On his return from military service, utilizing his “G. I. bill,” Bill made ar¬

rangements to commence graduate work in the Department of Biology at Stanford

University, under the guidance of Gordon Floyd Ferris. He was assigned a graduate

student office in a wing of the Leland Stanford Junior Museum that served for

many years as the Natural History Museum (housing primarily Botany, Ento¬

mology, and Ichthyology). Bill completed his Master of Arts degree in 1947, with

the thesis topic, “The comparative morphology of the larval head in Lepidoptera”

(44 pages, and 14 plates). Ferris wanted him to continue work in morphology but

Bill strongly resisted. Bill was permitted to study “The insect community on

Baccharis pilularis De Candolle,” and completed a Doctoral Dissertation (Tilden

1948) of408 pages. This was published in part in Microentomology (Tilden 1951b)

and in additional papers (see a listing in Smith 1990b: 50-55). With the completion

of his doctoral degree in 1948, Bill became a member of the staff of the Department

of Biology, at the then San Jose State College (now University). He taught about

30 different courses in the 22 years before his retirement as Emeritus Professor,

in June 1970. His primary assignment, however, was Entomology.

Starting in the early 1950s, Bill not only worked each summer for the Vector

Control program of the Santa Clara County Health Department, which was under

the management of Edgar A. Smith, but he also served as a consultant in the

identification of insects and other arthropods of public health importance through¬

out the years.

According to C. Don MacNeill, Bill pioneered in the study of our western North

American skippers. As MacNeill commented, most lepidopterists had avoided

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Vol. 68(1)

Figures 12-17. Figure 12. 12th Annual meeting of the Lepidopterists’ Society held at the Santa

Barbara Museum of Natural History. Left to right, Fred T. Thome, Charles L. Remington, John A.

Comstock, and Bill Tilden (photo Santa Barbara News Press). Figure 13. Tilden family on deck of

“Matsonia” leaving San Francisco, July, 1963. Figure 14. Left to right, Bill Tilden and Lloyd M.

Martin, Prescott, Arizona, 30 Mar 1972 (photo by Hazel I. Tilden). Figure 15. Roy O. Kendall’s Open

House, San Antonio, Texas, 22 Jun 1972. Left to right, Harry K. Clench, Charles V. Covell, Jr.,

Kenelm W. Philip, and Bill Tilden (photo by Hazel I. Tilden). Figure 16. San Antonio, Texas, June,

1972. Left to right, Mrs. Wilbur S. McAlpine, Bill Tilden, John C. Downey, and Wilbur S. McAlpine

(photo by Hazel I. Tilden). Figure 17. Lepidopterists’ dinner on river boat, San Antonio, Texas, June,

1972. Left side of table, left to right—Harry K. Clench, Hazel I. Tilden, Bill Tilden, Jerry A. Powell,

Mignon Bush, Don R. Davis, and Julian P. Donahue; right side of table, left to right—Madeleine

Field, Lloyd M. Martin, Dorothy Martin, Stanley S. Nicolay, name unknown, and W. Donald Duck¬

worth.

1992

ARNAUD: JAMES WILSON TILDEN

the skippers because “they were difficult to catch, difficult to spread, and difficult

to classify.” This only made them more interesting to Bill, and he collected these

and other butterflies extensively in California and Arizona. Because of his com¬

mitment to prepare a book on the western butterflies in the Peterson Field Guides

series, Bill took many field trips from 1969 through 1986. He covered over 200,000

miles of road travel throughout western North America from Texas and the

Mexican border east to Kansas and north to western Canada and Alaska (three

trips).

From July to August of 1963, Bill and his family spent six weeks on a sabbatical

travelling to four islands of Hawaii (Hawaii, Kauai, Maui, and Oahu). They arrived

in Honolulu aboard the “Matsonia” on 20 Jul and were met by their friend Richard

Kong and their landlord, Mr. Lum. The rent of $ 10 per day was charged for their

three bedroom house with sitting room, kitchen, washroom, two bathrooms, and

included the use of a 1953 Chevrolet. Bill was able to consult with J. Linsley

Gressitt and G. Allan Samuelson at the Bernice P. Bishop Museum. With Walter

C. Mitchell, University of Hawaii, Alan Thistle, State Department of Agriculture,

and other entomologists, he was able to observe and discuss problems concerning

termites, armyworms, cactus moth ( Cactoblastis cactorum (Berg)), southern green

stink bug ( Nezara viridula (L.)), sterilization of male fruit flies, etc. He became

acquainted with new methods of biological and vector control, information that

he incorporated in his course instruction at San Jose State University. He also

studied the Lepidoptera of the Hawaiian Islands.

In March 1978, Bill and Hazel were able to spend two weeks in Great Britain,

studying the Lepidoptera collection at the British Museum (Natural History) in

London, and visiting Scotland. They were able to meet the retired Keeper of

Entomology, Norman D. Riley, who had been away from the museum for months

because of a serious illness but was fortunately at the museum on one of the days

of their visit. This permitted them to have their copy of A Field Guide to the

Butterflies of the West Indies autographed by Riley. Richard I. Vane-Wright made

the Lepidoptera collection available for Bill’s study of certain western North

American skipper types, including those of W. H. Evans.

Bill built a superb personal collection of North American butterflies, certain

moth families, and favorite Coleoptera families (Buprestidae, Cerambycidae, and

Meloidae). He maintained, at his home, a very carefully organized and curated

collection housed in California Academy of Sciences drawers and cases, and in

Schmitt-sized boxes. The development of this major collection was undertaken

in a private manner without fanfare and came as a surprise to some of his students

when they later learned of its size, scope, and perfection.

Of the six taxa described as new by Bill, primary types of four of them are

deposited in the Academy’s collection: the holotype and allotype of Callophrys

lemberti Tilden (CAS Entomol. Type No. 7239) (Tilden 1963: 292); the holotype

and allotype of Glaucopsyche lygdamus incognita Tilden (CAS Entomol. Type

No. 12171) (Tilden 1974: 213); the holotype and allotype oiMitoura siva mans-

fieldi Tilden (CAS Entomol. Type No. 7240) (Tilden 1951b: 96); the holotype and

allotype of Euphilotes rita pallescens (Tilden & Downey) (CAS Entomol. Type

No. 6237) (Tilden & Downey 1955: 25; as Philotes). The holotype of Tharsalea

arota schellbachi (Tilden) is deposited in the National Museum of Natural History,

Smithsonian Institution, Washington, D.C. (Tilden 1955: 72; as Lycaena ), and

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Vol. 68(1)

Figures 18-22. Figure 18. Bill Tilden’s 75th Birthday party at his home in San Jose, California,

31 Dec 1979. Left to right, Paul H. Amaud, Jr., Hugh B. Leech, Dick Mewaldt, and Bill Tilden (photo

by Hazel I. Tilden). Figure 19. Bill Tilden, with drawer of Lepidoptera at his home in San Jose,

California, ca. 1980 (photo by Hazel I. Tilden). Figure 20. Bill Tilden with net, Palo Alto Bay lands,

California, ca. 1980 (photo by Hazel I. Tilden). Figure 21. Book signing at Book Cafe, Capitola,

California, 12 Oct 1986. Left to right, Arthur C. Smith and Bill Tilden (photo by Hazel I. Tilden).

Figure 22. Book signing at Willow Glenn Tattler, San Jose, California, 22 Nov 1986. Left to right,

Madeline M. Amaud, Paul H. Amaud, Jr., Arthur C. Smith, Bill Tilden, John S. Garth (phofo by

Hazel I. Tilden).

1992

ARNAUD: JAMES WILSON TILDEN

the holotype of Philotiella speciosa bohartorum (Tilden) is deposited in the Los

Angeles County Museum of Natural History, Los Angeles (Tilden 1968: 281; as

Philotes).

The paratypes of butterflies contained in Bill’s collection, when received by the

Academy in 1989, totaled 85 specimens representing 30 species, as follows: Family

Hesperiidae: 2, Agathymus dawsoni Harbison; 3, Erynnis telemachus Bums; 1,

Hesperia miriamae MacNeill; 2, Hesperia pahaska martini MacNeill; 2, Hesperia

comma tildeni H. A. Freeman; 2, Hesperia uncas macswaini MacNeill; 2, Mega¬

thymus coloradensis louiseaeW. A. Freeman; 2, Megathymus coloradensis reinthali

H. A. Freeman; 3, Pholisora gracielae MacNeill; and 2, Stallingsia maculosa (H.

A. Freeman). Family Papilionidae: 1, Papilio cresphontes pennsylvanicus F. & R.

Chermock. Family Pieridae: 2, Colias gigantea mayi F. & R. Chermock. Family

Lycaenidae: 3, Callophrys lemberti Tilden; 2, Epidemia dorcas megaloceras Ferris;

2, Euristrymon Ontario violae (D. Stallings & Turner); 13, Everes comyntas texanus

R. Chermock; 2, Glaucopsyche lygdamus jacki D. Stallings & Turner; 4, Mitoura

siva mansfieldi Tilden; 2, Mitoura thornei J. W. Brown; 4, Philotes enoptes bayensis

(Langston); 1, Philotes enoptes tildeni (Langston); 11, Euphilotes pallescens (Tilden

& Downey); 1, Philotiella speciosa bohartorum (Tilden); and 2, Tharsalea arota

schellbachi (Tilden). Family Nymphalidae: 2, Basilarchia archippus lahontani

(Herlan); 2, Phyciodes orseis herlani Bauer; 2, Speyeria aphrodite manitoba (F. &

R. Chermock); 1, Speyeria cybele pseudocarpenteri (F. & R. Chermock); and 1,

Vanessa anabella (Field). Family Satyridae: 6, Enodia portlandia missarkae (J.

R. Heitzman & dos Passos). The collection also contained 12 specimens labelled

as paratypes in the genera Boloria, Euphyes, and Plebejus that appear to be manu¬

script names, as they cannot be confirmed in current Lepidoptera checklists.

As commented by Heifer (1989), Bill was the first lepidopterist to investigate

the occurrence of the Lotis blue {Lycaeides idas lotis (Lintner)) colony that was

discovered south of Caspar, in Mendocino County. Bill also published on San

Francisco’s vanishing butterflies (1956). He knew the locality, in the Santa Cmz

Mountains, where the now possibly extinct Strohbeen’s pamassian {Parnassius

clodius strohbeeni Stemitzky) occurred and encountered his first specimen there

on 12 Jun 1933 (Tilden 1941). He collected only a few specimens and the two

given to the Academy bear his collection dates of 21 Jun 1936 and 1 Jul 1958.

The latter may be one of the last collected specimens. The Tilden bequest con¬

tained four specimens of this taxon, which Bill had earlier emphasized to me that

he wanted deposited in the Academy collection (Bill had recently declined an offer

of $3,000 for these four rare specimens.)

Bill Tilden had a long association with the Department of Entomology of the

California Academy of Sciences. Letters indicate that Hartford H. Keifer and

Edward P. Van Duzee identified Microlepidoptera and larger moths (noctuids

and geometrids) for him in 1933, and in 1936 beetles were identified by Edwin

C. Van Dyke. Before his entry into military service in 1942, Bill had made an

agreement with the Academy Director, Robert C. Miller, and Entomology Curator,

Edward S. Ross (Tilden letters: 4 and 31 Mar 1942), for the presentation of his

collection to the Academy, but there appeared to be a problem of delivery: Bill

did not have transportation, Ross was about to leave for his military duty, and

there was strict gasoline rationing for civilians during the war. The collection was

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

then stored in Santa Cruz, but unfortunately was somewhat damaged while being

fumigated during his absence.

After World War II, from 1946 through 1988, Bill donated portions of his

collection, totaling 35,607 specimens (27,087 Lepidoptera, 7091 Coleoptera, and

1429 miscellaneous insects and arachnids), to the Academy. Following his death,

the receipt of 20,274 additional specimens of Lepidoptera (16,460 pinned and

spread; 3814 papered) gives a total of 47,361 Lepidoptera received through 1991.

In addition, with nearly 8000 specimens of Coleoptera still to be accessioned, the

final Tilden donations will total approximately 64,000 specimens.

The California Academy of Sciences has been indeed fortunate to have such

loyal support of Bill and his family for nearly half a century. Bill became a member

of the California Academy of Sciences on 13 Apr 1948, at a time when there were

fewer than fifteen hundred active members. He was elected a Fellow of the Acad¬

emy in 1968. Bill’s reprint library on Lepidoptera was donated to the Department

of Entomology, California Academy of Sciences, with the stipulation that if papers

were duplicate to Academy holdings they would be sent to the Department of

Entomology, Bernice P. Bishop Museum. Bill’s correspondence files are also stored

in the Archives section in the Special Collections of the Academy’s Library.

Prior to his retirement, Bill also donated about 100 drawers of Lepidoptera and

miscellaneous insects, including Odonata, Diptera, and Coleoptera, to the En¬

tomology Museum at San Jose State University (a collection that now houses

about 600,000 specimens in the Carl D. Duncan Hall of Science).

Bill Tilden lived a full and active life devoted to family, friends, and institutions.

He overcame the limitations of the depression years and a delayed higher edu¬

cation to fully dedicate his exceptional talents to teaching, researching, publishing,

and collection making. Shortly after his retirement, Bill faced heart problems that

included open heart surgery, valve replacement; eventually he had a pacemaker

installed. Yet he did most of his extensive field collecting program for the Guides

following this surgery. The development of his collection and its donation to the

Academy places a responsibility on the Academy to oversee its proper storage,

make it available to the scientific community, and to preserve it for use by future

generations.

Bill Tilden is survived by his wife, Hazel Irene Tilden, of San Jose, California;

two sons, Bruce Allen Tilden of San Jose, California, and James Wilson Tilden,

Jr. of Phuket, Thailand; one daughter, Janice Elaine Tilden of Denver, Colorado;

and two brothers, Thomas C. Tilden of Santa Cruz, California, and Earl R. Tilden

of Dillingham, Alaska.

Acknowledgment

For information, loan of photographs, and review of this manuscript I am

especially grateful to Hazel I. Tilden of San Jose, California; Thomas C. Tilden

of Santa Cruz, California; and Arthur C. Smith of Watsonville, California; Donald

J. Burdick, California State University, Fresno, California; Helen K. Court, Vin¬

cent F. Lee, C. Don MacNeill, and Keve J. Ribardo of the Department of En¬

tomology, California Academy of Sciences. I would also like to thank Caroline

Kopp and Charlotte Fiorito of the Photography Department, California Academy

of Sciences, for their photographic assistance with the illustrations.

1992 ARNAUD: JAMES WILSON TILDEN 37

Literature Cited

Anonymous. 1989. [In Memoriam.] Professor Emeritus James W. Tilden. San Jose State University

Digest, Summer 1989: p. 4.

Garth, J. S. & J. W. Tilden. 1986. California butterflies. California Natural History Guides, 51.

University of California Press, Berkeley.

Heifer, J. 1989. [Death of J. W. Tilden.] Mendocino Beacon, February 16, 1989: p. 5.

Moreno, E. M. 1989. Entomology professor dies, colleague calls him ‘genius’. Spartan Daily, January

31, 1989.

Opler, P. A. & A. C. Smith. 1990. James Wilson Tilden, 1904-1988. Bull. Entomol. Soc. Am. (1989),

35(4): 47-48.

Smith, A. C. 1990a. [Metamorphosis]. J. W. Tilden. News Lepidop. Soc., 1989(2): 38.

Smith, A. C. 1990b. [Obituary]. James Wilson Tilden (1904-1988), a remembrance. J. Lepidop. Soc.,

44: 45-55.

Tilden, J. W. 1940. Preliminary list of the butterflies and skippers of Santa Clara and Santa Cruz

Counties, California. Natural Science Department, San Jose State College, San Jose, California.

Tilden, J. W. 1941. Collectors’ calendar for Santa Clara and Santa Cruz counties, California, and

preliminary list of the butterflies and skippers of Santa Clara and Santa Cruz counties. Ento¬

mologists’ Exchange News, 6(4): 1-6.

Tilden, J. W. 1948. The insect community on Baccharispilularis De Candolle. Ph.D. Thesis, Stanford

University.

Tilden, J. W. 1951a. A new subspecies of Mitoura siva Edwards. Bull. So. Calif. Acad. Sci., 50(2):

96-98.

Tilden, J. W. 1951b. The insect associates of Baccharis pilularis De Candolle. Microentomology,

16(1): 149-188.

Tilden, J. W. 1951. The insect associates of Baccharis pilularis De Candolle. Microentomology,

16(1): 149-188.

Tilden, J. W. 1955. A revision of Tharsalea Scud. (s. str.) with description of a new subspecies

(Lepid., Lyc.). Bull. So. Calif. Acad. Sci., 54(2): 67-77.

Tilden, J. W. 1956. San Francisco’s vanishing butterflies. Lepidop. News, 10(3/4): 113-115.

Tilden, J. W. 1963. An analysis of the North American species of the genus Callophrys. J. Res.

Lepidop., 1: 281-300.

Tilden, J. W. 1965. Butterflies of the San Francisco Bay region. California Natural History Guides,

12. University of California Press, Berkeley.

Tilden, J. W. 1968. A previously unrecognized subspecies of Philotes speciosa. J. Res. Lepidop.

(1967), 6: 281-284.

Tilden, J. W. 1974. A name for Glaucopsyche lygdamus behrii auct., not Edwards 1862. J. Res.

Lepidop. (1973), 12: 213-215.

Tilden, J. W. & J. C. Downey. 1955. A new species of Philotes from Utah. Bull. So. Calif. Acad.

Sci., 54(1): 25-29.

Tilden, J. W. & A. C. Smith. 1986. A field guide to western butterflies. The Peterson field guide

series. Houghton Mifflin Company, Boston.

Received 18 April 1991; accepted 24 May 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 38-45, (1992)

PREVALENCE OF TWO BACILLUS POPILLIAE DUTKY

MORPHOTYPES AND BLUE DISEASE IN

CYCLOCEPHALA HIRTA LECONTE

(COLEOPTERA: SCARABAEIDAE)

POPULATIONS IN CALIFORNIA

Harry K. Kaya, 1 Michael G. Klein, 2 T. M. Burlando, 1

Robert E. Harrison, 3 and Lawrence A. Lacey 4

department of Nematology, University of California, Davis, California 95616

2 USDA, Agricultural Research Service, Application Technology Research Unit,

Wooster, Ohio 44691

Cooperative Agricultural Research Program, Tennessee State University,

Nashville, Tennessee 37209

4 USDA, Agricultural Research Service, APO, New York 09406

Abstract. — The milky disease bacterium, Bacillus popilliae Dutky, and the blue disease rickettsial

organism, Rickettsiella popilliae (Dutky & Gooden), were isolated from larval populations of

Cyclocephala hirta LeConte in California. Two morphological types of B. popilliae, characterized

by the size of sporangia, spores, and parasporal bodies, were recovered. The typical Cyclocephala

strain had a sporangium averaging 5.5 x 2.1 pm, a spore averaging 2.1 x 0.9 pm, and a primary

parasporal body averaging 1.9 x 0.7 pm. Multiple parasporal bodies occurred in up to 50% of

this strain. The smaller atypical Cyclocephala strain had a sporangium averaging 4.4 x 1.3 jum,

a spore averaging 1.9 x 0.8 pm, and a parasporal body averaging 1.0 x 0.7 pm. Bacillus popilliae

was recovered from seven of eight sample sites located in southern and northern California. The

typical Cyclocephala strain of B. popilliae was predominant in larvae at two sites from southern

and one site from northern California, whereas the atypical strain was predominant in larvae at

five sites from northern California. Rickettsiella popilliae was recovered from larvae at two sites

in northern California.

Key Words. — Insecta, Scarabaeidae, Cyclocephala hirta, Bacillus popilliae, Rickettsiella popilliae,

milky disease, blue disease, turfgrass

Scarabaeid larvae are the most serious pests of turfgrass throughout much of

the United States (Tashiro 1987). In the northeastern United States, introduced

scarabaeid species, notably the Japanese beetle, Popillia japonica Newman, the

oriental beetle, Anomala orientalis Waterhouse, and the Asiatic garden beetle,

Maladera castanea (Arrow), cause severe damage to turfgrass. In the midwestem

states, the Japanese beetle and native species of May and June beetles, Phyllophaga

spp., and masked chafers, Cyclocephala spp., are more prevalent. In California,

a complex of Cyclocephala species occurs (Saylor 1945, Endrodi 1985), some of

which cause extensive turf damage. These pestiferous species include Cyclocephala

hirta LeConte, C. lurida Bland (= immaculata (Olivier)), C. longula LeConte, C.

melanocephala (Fabr.), and C. pasadenae Casey (Ali 1989). In 1988, we observed

that scarabaeid larvae caused considerable damage to turfgrass at a golf course in

northern California. Moreover, we detected larvae infected with the bacterium

Bacillus popilliae Dutky, the causal agent of milky disease, and Rickettsiella po¬

pilliae (Dutky & Gooden), the causal agent of blue disease. Because of the bio¬

logical control potential of these microorganisms, in particular the milky disease

1992

KAY A ET AL.: BACTERIAL DISEASE IN CYCLOCEPHALA

bacterium (Klein 1988), we determined their prevalence in scarabaeid populations

in California.

Materials and Methods

Second- and third-instar scarabaeids were sampled during the spring and fall

of 1989 and 1990 from northern and southern California. In the towns of Fairfax,

Moraga, San Rafael, and Santa Rosa, larval density was determined by taking at

least three 18x18x10 cm soil samples and counting the number of larvae in

each. Counts were averaged and the number of larvae per 0.1m 2 was determined.

In the town of Walnut Creek, only one larval density was taken because of the

small area of infestation. In the town of Chino, larval density was determined by

taking ten 0.1 m 2 samples or by estimating the number of larvae per 0.1 m 2 .

Larval density was estimated because it was too low and a wide area was sampled

to collect sufficient larvae for examination.

Larvae were collected from areas of golf courses and parks where no chemical

insecticides had been applied. Larvae were held either individually in 35 ml plastic

cups or in groups of eight to 10 larvae with field soil in a 210 ml plastic cup.

Groups of larvae were placed in field soil and were held for three to 10 days in

cold storage (10° C) before they were examined. Individually held larvae were

examined immediately upon return to the laboratory or placed in 35 ml cups

with 31 grams of sterilized dry soil (75% sand, 18% silt, 7% clay, pH 6.9, organic

matter 0.3%), 4.75 ml of distilled water, and perennial rye grass seeds. These

larvae were held for 19 days at 25 ± 2° C to allow further development and

expression of disease symptoms before they were examined.

Each larva was bled by inserting a sterilized size-2 insect pin through the

integument behind the head and a drop of hemolymph was placed on a microscope

slide. The hemolymph was examined for bacterial or rickettsial infection using

phase contrast microscopy at 400 x. Measurements of B. popilliae sporangia,

spores, and parasporal bodies at their longest and widest points were made with

an interference contrast microscope system at 9000 x and computer software

image analyzer. Data for measurements of B. popilliae (lengths and widths of the

sporangia, spores, and parasporal bodies) were analyzed using SAS program (t-

test) (SAS Institute 1988).

Two steps were used in identifying the scarabaeids. Raster patterns were used

to identify larvae to genus, and a subset of larvae was reared to adults. Adults

were identified to species by examination of the male genitalia (Saylor 1945,

Endrodi 1985).

Results and Discussion

Only one scarabaeid genus, Cyclocephala, was recovered during this study and

adult male specimens from each site were identified as C. hirta. Distribution of

larvae in turfgrass was patchy with infested areas ranging from <1 to 38 larvae/

0.1 m 2 (Table 1).

Bacillus popilliae was recovered from C. hirta in seven out of eight sites in

California and the prevalence of infection ranged from none to 71% (Table 1).

Even within a site, the prevalence of infection varied greatly indicating the uneven

distribution of spores in the soil. Thus, in Santa Rosa, we sampled two adjacent

areas (Santa Rosa 1A and IB) which were separated by about 50 m. Cyclocephala

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

Table 1. Prevalence of Bacillus popilliae and Rickettsiella popilliae from Cyclocephala hirta in

California.

Collection

% larvae infected with

Larval density/0.1

m 2 ± SD

Site 1

Date

No. larvae

Bacillus

Rickettsiella

Chino 1

5 May 89

61.9

3 Oct 89

199

28.1

19 ± 5.1

23 Apr 90

41.7

Chino 2

5 May 89

25.0

Fairfax

3 Oct 90

2.6

Moraga A

25 Sep 89

511

17.8

4.7

22 ± 15.5

2 Oct 89 b

11.1

22.2

ND C

Moraga B

24 Sep 90

3.3

38 ± 4.0

Moraga C

24 Sep 90

94.7

San Rafael

19 Sep 90

17.5

9 ± 3.0

Santa Rosa 1A

19 Sep 90

4 ± 1.7

Santa Rosa IB

19 Sep 90

71.0

6 ± 5.1

Santa Rosa 2

4 Oct 90

15.8

10 ± 4.6

Walnut Creek

24 Sep 90

16.2

18 d

a Moraga A, B, and C are samples from the same golf course. Larvae were collected from areas

separated by more than 300 m. Santa Rosa 1A and IB sites were located in the rough of a golf course

no more than 50 m from each other. Site 1A was on a slope and Site IB was a level area.

b Only larvae showing signs of a frank infection were bled and examined for B. popilliae or R.

popilliae infection. The remaining 54 larvae were held individually for 17 days before they were

examined (see text for data).

c ND = no data.

d Only one larval density measurement taken. Larvae were concentrated in a small area.

hirta larvae from one area had a high prevalence (71%) of B. popilliae infection,

whereas larvae from the other area were apparently free from infection (Table 1).

Similarly in Moraga, the prevalence of infection ranged from none to 17.8% at

three different areas within the same golf course.

Although milky disease bacteria have been isolated from other Cyclocephala

species on a number of occasions (White 1948, Harris 1959, Warren & Potter

1983, Boucias et al. 1986, Hanula & Andreadis 1988, Cherry & Boucias 1989),

we believe that this is the first report of milky disease bacteria in C. hirta pop¬

ulations. We also recovered two morphological types of B. popilliae from diseased

larvae (Fig. 1). In Chino (southern California) and Moraga (northern California),

the spores and parasporal bodies are primarily typical of those isolated from other

Cyclocephala species (Klein 1981). They have large and often multiple parasporal

bodies located adjacent to or overlapping the spore (Figs. IB, 1C). When multiple

parasporal bodies occurred, there was a primary parasporal body with a maximum

of three secondary bodies observed (Table 2). Multiple parasporal bodies were

found in up to 50% of these milky disease bacteria. A second type (Fig. 1A),

isolated primarily from northern California, had a single parasporal body which

was significantly smaller than the typical Cyclocephala strain (Table 2) and re¬

sembled B. popilliae from the Japanese beetle (Dutky 1940). We refer to this

isolate as the atypical Cyclocephala strain of B. popilliae and, as far as we are

aware, it is the first report of this morphological variant from a Cyclocephala

species.

*»

Figure 1. Bacillus popilliae spores from Cyclocephala hirta. A. Atypical Cyclocephala strain with

single parasporal body (p) and spore (s). B. Typical Cyclocephala strain with a mixture of single (p)

and multiple parasporal bodies (mp) and a single spore (s). Bar for A and B indicate 10 jim. C. Typical

Cyclocephala strain with multiple parasporal bodies (mp) distributed within a sporangium (sg). Bar

indicates 5 jim.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(1)

Table 2. Mean measurements in micrometers (±SD) of the sporangium, spore, and parasporal

body from the typical and atypical Cyclocephala strain of Bacillus popilliae from Cyclocephala hirta

in California.

B. popilliae strain 3

Typical

Atypical

Length

Width

Length

Width

Sporangium (^m)

5.5 (± 0.6)A

2.1 (± 0.2)a

4.4 (± 0.02)B

1.3 (± 0.1)b

Spore (nm)

Parasporal body Oum)

2.1 (± 0.3)A

0.9 (± 0.1)a

1.9 (± 0.05)B

0.8 (± 0.1)a

Primary

1.9 (± 0.1)A

0.7 (± 0.1)a

1.0 (± 0.1)B

-j

Secondary

1.0 (± 0.5)

0.5 (± 0.2)

—

a Means followed by different upper or lower case letters within a row are significantly different

(P < 0.001). Comparison was made only with the primary parasporal body.

The sporangium length (. F = 1015.5; df = 24; P = 0.0001) and width (F = 4.05;

df = 24; P = 0.0001), the spore length {F = 35.86; df = 24; P = 0.0017), and the

primary parasporal body length ( F = 19.47; df = 24; P = 0.0001) and width (F

= 2.61; df = 24; P = 0.0001) of the typical B. popilliae strain were significantly

longer or wider than the atypical B. popilliae strain. Only the width of the spores

was not significantly different.

The isolation of different morphological bacterial spores responsible for milky

disease from the same insect species is not unusual (Milner 1981), but the differ¬

ence relates to the presence or absence of the parasporal body (Dutky 1940). For

example, Boucias et al. (1986) and Hanula & Andreadis (1988) isolated milky

disease bacteria with and without a parasporal body from the same species of

scarabaeid larvae. In our study, we did not observe milky disease bacteria without

parasporal bodies. We isolated milky disease bacteria which were different in size

and in the number of parasporal bodies. The reason for the differences between

the two morphological spore/parasporal body complexes isolated from C. hirta

is unknown. Although the majority of sites contained either the typical or atypical

strain, we did observe an occasional larva with the typical strain at a site which

had the atypical strain and vice versa (HKK, unpublished data). A caveat is that

the typical Cyclocephala strain appears very much like the atypical strain during

early stages of sporogenesis, and when classifying the spore/parasporal body com¬

plex, sporogenesis must be complete.

Rickettsiella popilliae was only recovered from C. hirta larvae in Walnut Creek

and the three sample areas at Moraga (Table 1). This restricted occurrence of

Rickettsiella infection confirms earlier observations by other workers who also

recovered the disease from larvae at only a small proportion of sites examined

(Dutky & Gooden 1952, Hanula & Andreadis 1988). Larvae infected with Rick¬

ettsiella were easily detected by their bluish color, and upon examination of

dissected tissues with phase contrast microscopy, by the presence of crystals and

Brownian movement of the Rickettsiella cells in the fat body and hemolymph.

Moreover, we confirmed the Rickettsiella infection by transmission electron mi¬

croscopy (Fig. 2).

Some larvae infected with Rickettsiella did not appear blue during the early

stages of infection, but Brownian movement of particles was evident in the he-

1992

KAYA ET AL.: BACTERIAL DISEASE IN CYCLOCEPHALA

Figure 2. Transmission electron micrograph of Rickettsiella popilliae (kidney-shaped particles)

from Cyclocephala hirta and crystals (arrow) which are associate with the infection. 24,000 x.

molymph. Larvae collected in the field in an advanced stage of Rickettsiella

infection appeared milky due to the turbidity of the hemolymph (Dutky & Gooden

1952). Without a microscopic examination, the larvae in early and late stages of

infection may be erroneously diagnosed as healthy or as B. popilliae infection,

respectively.

The importance of examining the hemolymph for bacterial and rickettsial in¬

fection was shown when 81 C. hirta larvae collected in Moraga A on 2 Oct 1989

were separated visually into “healthy,” “milky” and “blue” (Table 1). Those

diagnosed as “milky” or “blue” were bled and examined to confirm the disease

and in all cases were infected with B. popilliae or R. popilliae. Fifty-four larvae

classified as “healthy” were held individually for 17 days in sterilized soil before

they were bled and examined for pathogens. Of the 54 larvae, four (7.4%) were

infected with B. popilliae, 11 (20.3%) were infected with Rickettsiella, and eight

(14.8%) died from unknown causes. Thus, the pooled data for the 2 Oct collection

in = 81) should be higher for B. popilliae (11.1% on 2 Oct vs 16.0% on 19 Oct)

and R. popilliae (22.2% on 2 Oct vs 35.8% on 19 Oct). In another example, only

5.5% of the larvae collected at Chino 1 on 3 Oct 1989 had apparent milky disease

symptoms. Microscopic examination of the hemolymph revealed the true rate of

infection to be 28% (Table 1).

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

We detected both milky and blue diseases in C. hirta populations, but the most

prevalent disease throughout California was milky disease. Although the preva¬

lence of infection of milky disease was not related to larval density, a long term

study may show density dependent relationships. The isolation of two morpho-

types of B. popilliae containing parasporal bodies (either multiple or single) was

a unique feature of our study. The natural occurrence of milky disease and its

high prevalence in some C. hirta populations indicate that these bacteria (i.e., the

typical and atypical Cyclocephala strains of B. popilliae ) are important in sup¬

pressing localized populations. Our data suggest that the milky disease organisms

may offer potential as effective biological control agents if they are introduced

into areas where they do not occur.

Acknowledgment

We thank Jean Adams for the electron micrograph, Bruce Jaffe for the inter¬

ference contrast photomicrographs, Kirk Smith for the species identification of

Cyclocephala hirta, Graham Thurston for the statistical analysis, and the golf

superintendents for their assistance in this study. This study was funded, in part,

by the California Statewide Integrated Pest Management Project.

Literature Cited

Ali, A. D. 1989. White grubs: the most troublesome turfgrass pest? Lawn & Landscape Maintenance,

July (1989): 48-51.

Boucias, D. G., R. H. Cherry & D. L. Anderson. 1986. Incidence of Bacillus popilliae in Ligyrus

subtropicus and Cyclocephala parallela (Coleoptera: Scarabaeidae) in Florida sugarcane fields.

Environ. Entomol., 15: 703-705.

Cherry, R. H. & D. G. Boucias. 1989. Incidence of Bacillus popilliae in different life stages of Florida

sugarcane grubs (Coleoptera: Scarabaeidae). J. Entomol. Sci., 24: 526-530.

Dutky, S. R. 1940. Two new spore-forming bacteria causing milky disease of Japanese beetle larvae.

J. Agric. Res., 61: 57-68.

Dutky, S. R. & E. L. Gooden, 1952. Coxiella popilliae, n. sp., a rickettsia causing blue disease of

Japanese beetle larvae. J. Bacteriol., 63: 743-750.

Endrodi, S. 1985. The Dynastinae of the world. Dr. W. Junk, The Hague, The Netherlands and

Akademiai Kiado, Budapest, Hungary.

Hanula, J. L. & T. G. Andreadis. 1988. Parasitic microorganisms of Japanese beetle (Coleoptera:

Scarabaeidae) and associated scarab larvae in Connecticut soils. Environ. Entomol., 17: 709-

714.

Harris, E. D. 1959. Observations of the occurrence of a milky disease among larvae of the northern

masked chafer, Cyclocephala borealis Arrow. Fla. Entomol., 42: 81-83.

Klein, M. G. 1981. Advances in the use of Bacillus popilliae. pp. 181-192. In Burges, H. D. (ed.).

Microbial control of pests and plant diseases 1970-1980. Academic Press, New York.

Klein, M. G. 1988. Pest management of soil-inhabiting insects with microorganisms. Agric. Ecosys.

Environ., 24: 337-349.

Milner, R. J. 1981. Identification of the Bacillus popilliae group of insect pathogens, pp. 45-60. In

Burges, H. D. (ed.). Microbial control of pests and plant diseases 1970-1980. Academic Press,

New York.

Saylor, L. W. 1945. Synoptic revision of the United States scarab beetles of the subfamily Dynastinae,

no. 1: tribe Cyclocephalini. J. Wash. Acad. Sci., 35: 378-386.

SAS Institute. 1988. SAS/STAT user guide. Release 6.03 ed. Cary, North Carolina.

Tashiro, H. 1987. Turfgrass insects of the United States and Canada. Cornell University, Ithaca,*

New York.

Warren, G. W. & D. A. Potter. 1983. Pathogenicity of Bacillus popilliae (Cyclocephala strain) and

1992

KAYA ET AL.: BACTERIAL DISEASE IN CYCLOCEPHALA

other milky disease bacteria in grubs of the southern masked chafer (Coleoptera: Scarabaeidae).

J. Econ. Entomol., 76: 69-73.

White, R. T. 1948. Milky disease infecting Cyclocephala larvae in the field. J. Econ. Entomol., 40:

912-913.

Received 29 May 1991; accepted 31 July 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 46-51, (1992)

THE GENETIC RELATIONSHIP BETWEEN

BOMBUS FRANKLINI (FRISON) AND

OTHER TAXA OF THE

SUBGENUS BOMBUS S.STR. (HYMENOPTERA: APIDAE)

A. Scholl, 1 R. W. Thorp, 2 and E. Obrecht 1

^oologisches Institut, Universitat Bern,

Baltzerstr. 3, CH-3012 Bern, Switzerland

department of Entomology, University of California, Davis, California 95616

Abstract.— Bombus franklini (Frison) has either been regarded as a distinct species or has been

synonymized with B. occidentalis Greene. We surveyed 21 enzymes by vertical starch-gel elec¬

trophoresis and compared B. franklini with B. occidentalis and nine other species of the subgenus

Bombus sensu stricto. We found that B. franklini differs from B. occidentalis at three enzyme

loci and there was no evidence of intergradation in areas of sympatry. According to the electro¬

phoretic data B. franklini is close to two species groups which comprise: (a) B. cryptarum (Fabr.),

B. magnus Vogt, B. moderatus Cresson and B. hypocrita Perez; and (b) B. occidentalis, B. terricola

Kirby and B. lucorum (L.), whi l e B. terrestris auct., B. affmis Cresson and B. sporadieus Nylander

are more distant.

Key Words. — Insecta, Apidae, enzyme electrophoresis, bumble bee genetic relationships

Among the North American bumble bee species, Bombus franklini (Frison) has

by far the smallest area of geographical distribution. All recent records have been

taken within a 60 mile radius of Grants Pass, Oregon (Thorp 1970). Bombus

franklini has puzzled entomologists for a long while. It was described from the

Oslar collection (Frison 1921) and the holotype was apparently erroneously re¬

corded from Nogales, Arizona, as discussed by Thorp (1970), who proposed Gold

Hill, Jackson County, Oregon, as the new type locality. Milliron (1971) considered

B. franklini conspecific with B. occidentalis Greene. Thorp et al. (1983), however,

had collected B. franklini at several localities sympatrically with B. occidentalis

and did not find intergrades between them. Plowright & Stephen (1980), working

on a multivariate analysis of wing venation data taken from queens, were able to

show a clear separation of B. franklini from other species within the subgenus.

They furthermore indicated that the male genitalia of B. franklini are markedly

different from those of B. occidentalis and advocated retention of specific status

for franklini, they concluded (Plowright & Stephen 1980: 479): “The origin of B.

franklini is mysterious. The results from the present study give no indication that

it is closely related to any of other nearctic representatives of its subgenus.”

In a recent study, Scholl et al. (1990) investigated the genetic relationships of

Nearctic and Palaearctic representatives of the subgenus Bombus s.str. by enzyme

electrophoretic data with special reference to B. moderatus Cresson, another prob¬

lematical taxon in this group. Bombus franklini unfortunately could not be in¬

cluded in this analysis because it was not available at that time. We now have

been able to collect B. franklini in northern California and in Oregon and we have

compared it electrophoretically with the previously studied representatives of the

subgenus Bombus s.str.

1992

SCHOLL ET AL.: BOMBUS GENETICS

Material and Methods

Specimens Analyzed.— Bombus franklini : CALIFORNIA. SISKIYOU Co.: Hilt,

29 Jul 1989, 12 workers; 6.4 km (4 mi) E of Yreka, 29 Jul 1989, 2 workers.

OREGON. JACKSON Co.: Ashland, 29 May 1990, 4 queens and 1 worker; Ruch,

29 May 1990, 12 workers; Central Point, 30 May 1990, 2 workers; Gold Hill, 30

May 1990, 12 workers.

Frozen homogenates of previously studied material [B. affinis Cresson, B. cryp-

tarum (Fabr.), B. hypocrita Perez, B. lucorum (L.), B. moderatus, B. occidentalis,

B. terrestris auct., B. terricola Kirby and B. sporadieus Nylander] (Scholl et al.

1990) and additional specimens, including one queen, nine workers and one male

of B. occidentalis from the same localities where B. franklini was collected and

10 queens, 38 workers and two males of B. occidentalis from eight other localities

in northern California have been used for electrophoretic comparison. This ma¬

terial is summarized in Table 1. Californian B. occidentalis included the nominate

subspecies and B. o. nigroscutatus Franklin along with their intergrades, these are

not listed separately because the electrophoretic data did not indicate any differ¬

ence.

Electrophoresis. — We have used the same methods (vertical starch-gel electro¬

phoresis) and enzymes (21 loci) as Scholl et al. (1990). These enzymes are:

Aconitase, 2 loci: Acon-1 and Acon-2; Arginine kinase, Apk; Hydroxybutyric

dehydrogenase, Bdh; Esterase, Est-1; a-Glycerophosphate dehydrogenase, 2 loci:

a-Gpd-2 and a-Gpd-3; Glutamic-oxaloacetic transaminase, Got-2; Glutamic-py¬

ruvic transaminase, Gpt; Hexokinase, 2 loci: Hk-1 and Hk-3; Isocitrate dehy¬

drogenase, 2 loci: Idh (NAD) and Idh (NADP); Leucine aminopeptidase, Lap;

Malate dehydrogenase, 2 loci: Mdh-1 and Mdh-2; Malic enzyme, Mod; Peptidase,

Pep; Phosphoglucose isomerase, Pgi; Phosphoglucomutase, Pgm; Superoxide dis-

mutase, Sod (for details see Scholl et al. 1990). A phenogram of the genetic

relationships of the species investigated was constructed by average linkage cluster

analysis (UPGMA) (Nei 1987) using Nei’s (1972) standard coefficient of genetic

identity (I).

Results and Discussion

Ten of the 21 loci scored were found invariant in all species surveyed. These

loci are: Acon-2, Apk, Bdh, a-Gpd-l, a-Gpd-2, Idh (NADP), Lap, Mdh-2, Mod,

and Sod. Eleven loci showed interspecific variation. The zymograms observed

are schematically shown in Fig. 1, where the designation of electromorphs is based

on mobilities (in mm) relative to the electromorph of B. lucorum (= index 100),

as in previous electrophoretic studies on bumble bees (e.g., Scholl & Obrecht

1983, Scholl et al. 1990).

Bombus franklini was monomorphic in all loci scored, except Pep, where one

worker was heterozygous. Minor polymorphisms were observed in some loci of

other species. These are: Acon-1 in B. lucorum, B. terrestris and B. spor adieus]

Est-1 in B. lucorum and B. terricola, Got-2 in B. cryptarum, B. magnus, B.

lucorum, B. occidentalis, B. terrestris and B. sporadicus] Hk-1 in B. occidentalis]

Mdh-1 in B. terrestris. The level of polymorphism, however, was usually very

low (H < 0.05), as also observed previously in other bumble bee species (Obrecht

& Scholl 1981, Pamilo et al. 1984), except in Got-2 of B. occidentalis, where the

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

ACON-1

EST-1

GOT-2

GPT

HK-1

HK-3

IDH (NAD)

MDH-1

PEP

PGI

PGM

Figure 1. Schematic illustration of enzyme phenotypes in Bombus s.str. species. (Note: B. lucorum

is the reference, assigned mobility index =100 for each enzyme.)

frequency of a minor allele Got-2 105 ranged between 0.05 and 0.25 at three sam¬

pling sites in Alberta, Calgary, Barrier Lake and Fortress Mountain respectively,

while allele Got-2 100 was fixed in samples from other areas (Table 1).

We have not found an electromorph that is unique to B. franklini. However,

Table 1. Specimens analyzed and origin of material.

California

Oregon

Alaska

British

Columbia

Alberta

Ontario

Japan

N-Europe

E-Europe

C-Europe

S-Europe

Total

B. franklini

—

(.n = 45)

B. moderatus

—

(n = 9)

—

B. crypt arum

—

(.n = 22)

—

B. magnus

—

(«= 17)

—

B. hypocrita

—

( n = 8)

B. lucorum

—

(n = 57)

—

B. occidentalis

—

101

(n= 110)

—

B. terricola

—

(n= 18)

—

B. terrestris

—

(n = 40)

B. affinis

—

(n= 11)

B. sporadicus

—

(n= 15)

N-Europe = Finland, Norway, Scotland, England; E-Europe = Hungary; C-Europe = Switzerland, France, Belgium; S-Europe = Spain, Italy.

4 ^

SCHOLL ET AL.: BOMB US GENETICS

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

- franklini

r moderatus

-cryptarum

| magnus

- hypocrita

- lucorum

- occidentalis

- terricola

- terrestris

i - affinis

- sporadicus

i-1-1-1-1-1

0.5 0.6 0.7 0.8 0.9 1.0

Nei coefficient!

Figure 2. Phenogram showing the genetic relationship between B. franklini and other taxa of the

subgenus Bombus s.str., as revealed by the electrophoretic data.

it is the electrophoretic pattern of enzymes that is unique to B. franklini (Fig. 1).

Thus, B. franklini differs from B. moderatus in GOT-2 and IDH (NAD); B.

terrestris is identical with B. franklini in GOT-2 and IDH (NAD), but these species

differ in ACON-1, HK-1 and PGM, etc. Bombus franklini differs from B. occi¬

dentalis in GOT-2, PEP and PGM; the Got-2 locus, however, was found weakly

polymorphic in three B. occidentalis samples from Southern Alberta, as mentioned

above. In our material from California and Oregon, there was always a clear

separation on the basis of these three enzymes and the electrophoretic data did

not provide any evidence of intergradation of B. franklini and B. occidentalis.

The genetic relationships of B. franklini and other Nearctic and Palaearctic

representatives of its subgenus, as revealed by the enzyme electrophoretic studies,

are presented in Fig. 2 as a phenogram that is based on a similarity matrix (Nei

coefficient I) calculated in pairwise species comparisons from the 21 loci surveyed.

According to these data, B. franklini is close to two species groups that comprise:

(a) the European B. cryptarum and B. magnus, the North American B. moderatus

and the Japanese B. hypocrita ; and (b) the European B. lucorum and the North

American B. terricola and B. occidentalis, while the European B. terrestris and 3.

sporadicus and the North American B. affinis are more distant. This new infor¬

mation adds to, but does not alter the basic genetic relationships among species

of the subgenus Bombus as determined by Scholl et al. (1990).

The narrow endemism of B. franklini is intriguing. As Thorp (1970) pointed

out, all recent records have been taken within a 60 mile radius of Grants Pass,

Oregon. Bombus franklini is keyed out from sympatric B. o. occidentalis by its

1992

SCHOLL ET AL.: BOMB US GENETICS

coat color. But Stephen (1957) found that separation is often difficult. The north¬

western coast is an area where several bumble bee species, including B. occiden-

talis, show gradation from one color form to another resulting in color convergence

toward local Mullerian mimicry groups (Plowright & Owen 1980, Thorp et al.

1983). However, B. occidentalis females do not have yellow anterolaterally on

the scutum, extending back beyond the tegulae, and B. franklini females are

uniform in color throughout the known range (Thorp et al. 1983). One might

speculate that B. franklini has in fact a more widespread distribution, but becomes

hidden within the color variation of B. occidentalis. The electromorphetic data

presented here provide an opportunity to test this hypothesis.

Acknowledgment

The competent assistance of Mrs. V. Siegfried and Mrs. L. Frauchiger in the

electrophoretic studies is gratefully acknowledged.

Literature Cited

Frison, T. H. 1921. New distribution records for North American Bremidae, with the description

of a new species (Hym.). Entomol. News, 32: 144-148.

Milliron, H. E. 1971. A monograph of the western hemisphere bumblebees (Hymenoptera: Apidae;

Bombinae). The genera Bombus and Megabombus subgenus Bombias. Mem. Entomol. Soc.

Can., 82.

Nei, M. 1972. Genetic distance between populations. Am. Nat., 106: 283-292.

Nei, M. 1987. Molecular evolutionary genetics. Columbia University Press, New York.

Obrecht, E. & A. Scholl. 1981. Enzymelektrophoretische Untersuchungen zur Analyse der Ver-

wandtschaftsgrade zwischen Hummel- und Schmarotzerhummelarten (Apidae, Bombus).

Apidologie, 12: 257-268.

Pamilo, P., S.-L. Varvio-Aho & A. Pekkarinen. 1984. Genetic variation in bumblebees {Bombus,

Psithyrus) and putative sibling species of Bombus lucorum. Hereditas, 101: 245-251.

Plowright, R. C. & R. E. Owen. 1980. The evolutionary significance of bumble bee color patterns:

a mimetic interpretation. Evolution, 34: 622-637.

Plowright, R. C. & W. P. Stephen. 1980. The taxonomic status of Bombus franklini (Hymenoptera:

Apidae). Can. Entomol., 112: 475-479.

Scholl, A. & E. Obrecht. 1983. Enzymelektrophoretische Untersuchungen zur Artabgrenzung im

Bombus lucorum- Komplex (Apidae, Bombini). Apidologie, 14: 65-78.

Scholl, A., E. Obrecht & R. E. Owen. 1990. The genetic relationship between Bombus moderatus

Cresson and the Bombus lucorum auct. species complex (Hymenoptera: Apidae). Can. J. Zool.,

68: 2264-2268.

Stephen, W. P. 1957. Bumble bees of western America (Hymenoptera: Apoidea). Tech. Bull. Oreg.

Agric. Exp. Stn., 40.

Thorp, R. W. 1970. The type locality of Bombus franklini and notes on putative Arizona records

of other Bombini. Pan-Pacific Entomol., 46: 177-180.

Thorp, R. W., D. S. Homing, Jr. & L. L. Dunning. 1983. Bumble bees and cuckoo bumble bees of

California. Bull. Calif. Insect Surv., 23.

Received 25 June 1991; accepted 13 August 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 52-61, (1992)

OCCURRENCE OF DIURAPHIS ( HOLCAPHIS) FREQUENS

(WALKER) (HOMOPTERA: APHIDIDAE) ON

WHEAT, NEW TO IDAHO, AND A KEY TO

NORTH AMERICAN DIURAPHIS

Susan E. Halbert, B. June Connelly and Ming-Guang Feng

Southwest Idaho Research and Extension Center,

Department of Plant, Soil and Entomological Sciences,

University of Idaho, Parma, Idaho 83660-6699

Abstract.— Colonies of a species of Diuraphis (Holcaphis ) were found on wheat near Parma,

Idaho in 1986. Morphologically, the species best fits the description of Diur aphis (. Holcaphis)

frequens (Walker), though the process terminalis is longer with respect to the base of the sixth

antennal segment than is reported for European D. frequens. The host range of the Idaho Diur aphis

frequens is also consistent with that reported for D. frequens in Europe, except that the populations

found in Idaho multiply much more quickly on wheat than on Elytrigia repens (L.) Beauvois.

In spite of these differences, we think this Diuraphis sp. is D. frequens. We do not expect that it

will become a serious pest in Idaho.

Key Words. — Insecta, Aphididae, Diuraphis frequens, Diuraphis noxia

In surveys of wheat ( Triticum aestivum L.) in 1986, colonies of an unusual

Diuraphis (.Holcaphis) were found near Parma, Idaho. Isolated plants were severely

stunted and contorted, their rolled and twisted leaves containing hundreds of wax-

covered aphids. The same species was collected in suction trap samples from

Parma, Rockland Valley and Arbon in 1986, from Parma in 1988 and 1989, and

from Kimberly and Moscow in 1990. We think this aphid is Diuraphis (Holcaphis)

frequens (Walker). We discuss its identity, based upon morphology and host range.

We also compare its ability to colonize the plants with that of Diuraphis (Diura¬

phis) noxia (Mordvilko), which is the only other species of Diuraphis known to

occur in Idaho, and with Rhopalosiphum padi (L.) and Schizaphis graminum

(Rondani), which are both common pests in the U.S. with wide host ranges among

the Gramineae.

Methods and Materials

Twenty apterous viviparae from a colony of D. frequens, collected in Canyon

County, Idaho on T. aestivum propagated on T. aestivum cv. ID0232, were mount¬

ed and measured using a Zeiss compound binocular microscope equipped with

an eyepiece micrometer. Two attempts were made to obtain additional specimens

from each of two accessions of Elytrigia repens (L.) Beauvois, but colonies grew

inadequately to produce sufficient adults.

Host plants for each of the described Diuraphis (Holcaphis) spp. were obtained

including Calamagrostis sp., Agrostis alba L., Agrostis palustris (Hudson) Persoon,

Agrostis tenuis Sibthorp, Apera interrupta (L.) Beauvois (formerly in Agrostis),

Holcus lanatus L., Bromus inermis Leys, Bromus tectorum L., Elytrigia repens

(formerly in Agropyron), Agropyron cristatum (L.) Gaertner, Thinopyrum ponti-

cum (Podperae) Barkworth & D. R. Dewey (second time only) and T. aestivum.

1992

HALBERT ET AL.: DIURAPHIS FREQUENS IN IDAHO

The plants were started from seed in the autumn of 1987, except for Cala-

magrostis sp., E. repens, B. inermis and B. tectorum, which were transplanted

from the field, trimmed and allowed to regrow new shoots that were solely used.

Plants were infested 30 Nov-3 Dec 1987 using three pots of each plant species

for each of the three species of aphids, including the Idaho D. frequens, D. noxia

and R. padi. The infestation rate was ten aphids per plant. On 14 Dec 1987, plants

were scored for colonization using the following scale: 1—no aphids, 2—a few

solitary aphids, 3—small colonies, 4—plant heavily colonized. After the readings,

the plants were cut back and sprayed with Bifenthren (Capture 2EC) (1.26 g a.i./

liter), if infestations were found.

On 18-19 Jan 1988, the same plants, with exceptions that A. interrupta and

the Minnesota accession of B. inermis were omitted and T. ponticum was added,

were infested with 20 aphids per pot in the same manner described above. On 3

Feb 1988, readings were taken as before.

In order to quantitatively determine relative colonization ability, the plants

were reinfested in 1989. The same plants (by then more than one year old) were

used, except the three Bromus accessions and A. cristatum (omitted because they

had died in the interim), an accession of E. repens from Moscow, Idaho (added),

and the wheat (about six weeks old). We used three pots of each plant species for

each of four species of aphids, including the Idaho D. frequens, D. noxia, R. padi

and S. graminum. Plants were infested with 20 adult aphids per plant on 30-31

Jan 1989. On 14-15 Feb 1989, the parts of the plants above ground were clipped

and placed in Berlese funnels until they dried completely. Aphids were counted

using a dissecting microscope. In the case of the wheat, 10% subsamples were

counted.

The data were analyzed using the ANOVA and LSD mean separation procedures

(0.05 significance level) in SAS software (SAS Institute 1985). Because plant and

aphid species interactions were significant, the species were analyzed separately.

The data were analyzed using a transformation to normalize the variance (Y =

Vcount + 0.5).

Results and Discussion

Morphology. — The subgenus Holcaphis is distinguished from Diuraphis s. str.

by lack of a supracaudal process, and currently includes six described species:

Diur aphis tritici (Gillette) * 1 (native to North America), Diur aphis agrostidis (Mud-

dathir), Diuraphis bromicola (Hille Ris Lambers), Diuraphis calamagrostis (Os-

siannilsson), Diuraphis frequens (Walker) and Diuraphis hold (Hille Ris Lambers)

(Eastop & Hille Ris Lambers 1976). In addition to D. tritici, only two species, D.

frequens and D. hold, have been reported from North America (Smith & Parron

1978). The ultimate rostral segment of the Idaho D. frequens is 0.069 mm long,

as compared with 0.12 mm for D. tritici, thus ruling out the possibility that our

aphid is D. tritici. Hille Ris Lambers (1939) separates European D. hold from D.

1 After this article went to press, Zhang et al. (1991) published a review of Diuraphis that treats D.

tritici as a subspecies of D. frequens. We prefer to retain these as distinct species until their change

can be confirmed by hybridization experiments. Two new Diuraphis from China, which are described

by Zhang et al. (1991), are not discussed here.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(1)

frequens using relative lengths of antennal segments III and IV + V, and the ratio

of the base of antennal segment VI to the process terminalis. The D. frequens

found in Idaho will key to D. frequens in Hille Ris Lambers (1939), using the

former character, and to D. hold, using the latter; however, the same was true of

five specimens of D. frequens collected in Enfield, England on 12 Jul 1987, in¬

dicating that the antennal segment VI character is not consistently reliable, even

in Europe.

The key by Muddathir (1965) separates the two species using siphuncular place¬

ment, and presence or absence of intersegmental muscle insertions and sclerites

on abdominal segment VI. Most Idaho material will key to D. frequens using

Muddathir’s key, but the siphunculi on some specimens are slightly closer to the

sixth than to the seventh abdominal spiracles, a character given there for D. hold.

Of the species in the subgenus not yet reported in North America, Diuraphis

bromicola was described from Bromus inermis (Hille Ris Lambers 1959), and can

be separated from D. hold and D. frequens by the absence of sclerotic areas on

the abdomen, other than on abdominal segment VIII. The Idaho specimens have

an obvious sclerotic bar on abdominal segment VII, which would appear to rule

out D. bromicola. Diuraphis agrostidis and D. calamagrostis have pore-like si¬

phunculi with the siphuncular aperture facing upward, while D. hold and D.

frequens have longer siphunculi, shaped such that the aperture faces posteriorly

(Muddathir 1965). Diuraphis agrostidis has In = 12 chromosomes and D. frequens

has In = 14 (Blackman 1980). The specimens from Idaho have siphunculi that

fit the description of D. frequens better than those of D. agrostidis and D. cala-

matrostis, and have 2n = 14 chromosomes (R. L. Blackman, personal commu¬

nication).

Occasional specimens of D. frequens collected in Idaho have a posteromedial

extension on abdominal tergite VIII that suggests a supracaudal process. These

specimens can be separated from Diuraphis {Diuraphis) nodulus (Richards) and

Diuraphis {Diuraphis) mexicana (Baker) by the position of the siphunculi in re¬

lation to the sixth and seventh abdominal spiracles. Siphunculi of D. frequens are

clearly between the sixth and seventh spiracles (Muddathir 1965), whereas si¬

phunculi of D. nodulus and D. mexicana are anterior to the sixth pair of spiracles.

Morphological differences among species found in North America can be sum¬

marized by the following key:

Key to Species of Diuraphis Reported in

North America

1. Supracaudal process present on apterous viviparae; siphunculi anterior

to sixth pair of abdominal spiracles (Fig. 1; also see Fig. 3) [Diuraphis

(. Diuraphis )]. 2

T. Supracaudal process on apterous viviparae absent or barely indicated;

siphunculi between sixth and seventh pair of abdominal spiracles (Fig.

2) [Diuraphis {Holcaphis)] . 3

2(1). Process terminalis of viviparae at least 2.0 x as long as base of antennal

segment VI; supracaudal process on apterous viviparae fingerlike and

at least 1.5 x as long as width at the middle (Fig. 1) . D. noxia

1992

HALBERT ET AL.: DIURAPHIS FREQUENS IN IDAHO

Figure 1. Diuraphis ( Diuraphis ) noxia (Mordvilko) apterous vivipara, showing supracaudal process

and position of siphunculi with respect to sixth and seventh pairs of abdominal spiracles.

Figure 2. Diuraphis (Ho leap his) frequens (Walker) apterous vivipara, showing supracaudal process

and position of siphunculi with respect to sixth and seventh pairs of abdominal spiracles.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

Figure 3. Diuraphis (Diuraphis) mexicana (Baker) apterous vivipara, showing supracaudal process,

position of siphunculi and length of process terminalis.

2'. Process terminalis of apterous viviparae less than 1.5 x as long as base

of antennal segment VI, of alate viviparae less than twice as long as

base of VI; supracaudal process on apterous viviparae broad and wider

than long, sometimes with a short projection in the center (Fig. 3) ..

. D. mexicana and D. nodulus 2

3(1). Ultimate rostral segment of all forms 0.12 mm long and nearly 3.0 x

as long as wide (Fig. 4). D. tritici

3'. Ultimate rostral segment of all forms 0.07 mm long and 2.0 x as long

as wide (Fig. 5). 4

4(3). Clear markings on abdominal segment VI of all viviparae; antennal

segment III on apterous viviparae longer than segments IV and V

combined; specific to Holcus (Fig. 6) . D. hold

4'. No markings on abdominal segment VI of viviparae; antennal segment

III on apterous viviparae shorter than segments IV and V combined;

on Agropyron, Triticum and Elytigia, but not on Holcus (Fig. 7) ...

. D. frequens

2 Resolution of D. nodulus and D. mexicana requires further taxonomic study. A revision of Di¬

ur aphis is pending (Manya B. Stoetzel, personal communication).

1992

HALBERT ET AL.: DIURAPHIS FREQUENS IN IDAHO

Figure 4. Diuraphis ( Holcaphis ) tritici (Gillette) apterous vivipara, showing ultimate rostral seg¬

ment. Scale: 1 cm = 0.05 mm actual size.

Figure 5. Diuraphis ( Holcaphis ) frequens (Walker) apterous vivipara, showing ultimate rostral

segment. Scale: 1 cm = 0.05 mm actual size.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

Figure 6. Diuraphis ( Holcaphis ) hold (Hille Ris Lambers) apterous vivipara, showing markings

on abdominal segments VI, VII and VIII and relative lengths of antennal segments.

Host Range.— In the first experiment, the Idaho D. frequens colonized only

wheat. In the second experiment, it also heavily colonized T. ponticum (not

included in the first experiment). No colonies were found on E. repens in the first

two experiments. In the final experiment, there was heavy colonization on wheat,

and smaller colonies were found in A interrupta, E. repens and T. ponticum (Table

1). This host range fits D. frequens, but not D. hold or any other Diur aphis

(Holcaphis ) sp. except D. tritici, which can be ruled out due to its much longer

ultimate rostral segment. We observed that Idaho D. frequens placed on Holcus

lanatus exhibited a toxic reaction; within an hour, the aphids fell off the plants

and were unable to move in a coordinated manner. We observed no endophytic

fungus in our H. lanatus leaves when preparations were made according to the

procedure by Saha et al. (1988). Thus, H. lanatus is not a host of the Idaho

Diur aphis sp., ruling out any possibility that it is D. hold.

It is, in fact, doubtful that D. hold occurs in North America in spite of the

records cited by Smith & Parron (1978). There are none in the Canadian national

collection in 1989 (Robert Foottit, personal communication), nor in the collection

at the Illinois State Natural History Survey in 1990 (David Voegtlin, personal

communication), nor in the collections at the California Dept, of Food & Agri-

1992

HALBERT ET AL.: DIURAPHIS FREQUENS IN IDAHO

Figure 7. Diuraphis (Holcaphis)frequens (Walker) apterous vivipara, showing pattern of abdominal

markings and relative lengths of antennal segments.

culture, Sacramento and the University of California, Berkeley, California in 1990

(John Sorensen, personal communication). Hille Ris Lambers (1939) indicated

that D. hold is restricted to plants in the genus Holcus. None of the North Amer¬

ican specimens identified as D. hold in collections at the United States National

Museum or the Palmer Collection at Colorado State University are specified to

have been collected from Holcus. About half of the slides have inadequate or no

host information, but specified host plants include petunia, quackgrass and Agro-

pyron glaucum, all unlikely hosts of D. hold. Forbes & Chan (1989) reported an

extensive survey of aphids in British Columbia where Holcus lanatus is abundant.

They report Hyalopteroides humilis (Walker) and Sitobion fragariae (Walker) on

H. lanatus, but no Diuraphis spp. were found. Thus, we are not aware of evidence

that any Diuraphis sp. colonizing Holcus occurs in North America.

Triticum aestivum was a better host for the Idaho D. frequens than any other

plant tested, including E. repens, the host reported to be preferred by D. frequens

in Europe (Hille Ris Lambers 1939) (Table 1); however, colonies on wheat were

not as large as those produced by the three pest species. We have not found D.

frequens on E. repens in Idaho, although a number of other aphid species have

been found, including Sip ha elegans del Guercio, D. noxia, Metopolophium dir-

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

Table 1. Mean numbers of aphids on various plants after two weeks of colonization, Parma, Idaho,

January-February, 1989. Means followed by the same letter (a, b, c) are not significantly different

from others in the same column using the LSD method of means separation.

Host

D. frequens

D. noxia

S. graminum

R. padi

Triticum aestivum L.

119.0 a

1816.7 a

2793.3 a

1450.0 a

Apera interrupta (L.) Beauvois

31.3 b

42.7 be

304.3 b

112.3 b

Elytrigia repens (L.) Beauvois (Moscow, Idaho)

6.3 c

94.0 b

57.7 be

7.3 c

Elytrigia repens (L.) Beauvois (Caldwell, Idaho)

9.0 c

31.0 be

36.0 be

Agrostis alba L.

0 c

0.3 c

12.0 be

118.7 b

Agrostis tenuis Sibthorp

0 c

0.3 c

38.0 be

3.0 c

Agrostis palustris (Hudson) Persoon

0 c

4.3 c

3.7 c

Holcus lanatus L.

0 c

5.0 c

7.0 c

Thinopyrum ponticum (Podperae)

Barkworth & D. R. Dewey

2.3 c

0 c

2.7 c

0 c

Calamagrostis sp.

0 c

71.7 be

0 c

hodum (Walker), Sitobion avenae (Fabr.), S. graminum and Forda marginata

Koch (Gittins et al. 1976; SEH, unpublished data).

The Idaho D. frequens, D. noxia and R. padi all colonized T. ponticum, a species

used in conservation plantings, much more heavily when the plants were young

than they did one year later. This observation suggests that some perennial grasses

may become less palatable to aphids over time. If so, mature conservation plant¬

ings pose fewer problems as reservoirs of aphid pests than young stands. This

question should be examined further.

Apera interrupta was usually colonized more heavily than Agrostis spp. This

supports recent botanical evidence that A. interrupta should not be placed in the

genus Agrostis as it has sometimes been in the past (McNeill 1981, Hitchcock &

Cronquist 1973).

Based on morphology and host range analysis, we think the Diuraphis sp. found

on Idaho wheat is D. frequens, although slight differences in morphology and host

preference remain to be resolved. These differences could be due to founder effect,

because it is likely that very few individuals were originally introduced into North

America.

Other species that have been introduced into North America have host ranges

that differ from their parent populations. Probably the most famous example is

Therioaphis trifolii (Monell). According to Blackman (1981), the original North

American population fed on Trifolium. About 70 years later, T. trifolii forma

“maculata ” (Buckton) appeared on alfalfa. This population has several traits that

are not typical of the parent population in the Old World. Evidence suggests that

the North American alfalfa population resulted from the introduction of a single

clone (Blackman 1981).

In the case of D. frequens, however, introduction of a single clone from Europe

may not explain the marked preference for wheat in Idaho, because wheat is not

considered to be a host of D. frequens in Europe. Another possible explanation

is that Idaho D. frequens came from Asia, across the Bering Strait, rather than

from Europe, and thus has a host range differing from European populations;

however, wheat is not listed as a host in western Siberia (Ivanovskaya 1977).

We have observed wheat plants colonized by this species each year. Infested

plants are usually severely damaged, but damage is restricted to isolated plants.

We have observed that the most common situation is to find several infested

1992

HALBERT ET AL.: DIURAPHIS FREQUENS IN IDAHO

plants near the edge of a late maturing field of spring wheat. Laboratory cultures

of the Idaho D. frequens do not produce many alatae in comparison with D. noxia

and D. tritici. If the same is true for field populations, this could explain its

restricted distribution. Theoretically, given the right conditions (e.g., when alate

production is greatly increased), outbreaks could occur, but it is unlikely that the

species will become a serious pest.

Acknowledgment

We thank Roger Blackman of the British Museum of Natural History for de¬

termining the chromosome number of our aphids and for loaning specimens. We

thank Manya Stoetzel (U.S. National Museum), Robert Foottit (Canadian Na¬

tional Museum), Frank Peairs, B. C. Kondratieff and Robert Hammon (Colorado

State University), Paul Brown (British Museum of Natural History), David Voegt-

lin (Illinois Natural History Survey) and John Sorensen (Calif. Dept. Food &

Agriculture) for providing information and/or specimens from collections at their

respective institutions. We thank Leslie Boydston and Keith Pike for photographs

in Figs. 3, 6 and 7. We thank Richard Old, Edward Northam, Anna Zeigler, Mark

Longstroth and Carol Scinocca for providing plants and seeds. We also thank

Thomas Mowry and Keith Dorschner for help with statistics, Jane Breen for

advice on endophyte analysis and Keith Dorschner for reviewing the manuscript.

This is Idaho Agricultural Experiment Station Scientific Paper number 91732.

Literature Cited

Blackman, R. L. 1980. Chromosome numbers in the Aphididae and their taxonomic significance.

Syst. Entomol., 5: 7-25.

Blackman, R. L. 1981. Species, sex and parthenogenesis in aphids, pp. 75-85. In Greenwood, P. H.

& P. L. Forey (eds.). The evolving biosphere. British Museum of Natural History, London.

Eastop, V. F. & D. Hille Ris Lambers. 1976. Survey of the world’s aphids. Junk, The Hague.

Forbes, A. R. & C. K. Chan. 1989. Aphids of British Columbia. Agriculture Canada Technical

Bulletin, 1989-IE.

Gittins, A. R., G. W. Bishop, G. F. Knowlton & E. J. Parker. 1976. An annotated list of the aphids

of Idaho. Idaho Agr. Exp. Stat. Res. Bull., 95.

Hille Ris Lambers, D. 1939. On some western European aphids. Zool. Med. Mus. Leiden, 22: 79-

199.

Hille Ris Lambers, D. 1959. Notes on European aphids with descriptions of new genera and species

(Homoptera, Aphididae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 32:

271-286.

Hitchcock, C. L. & A. Cronquist. 1973. Flora of the Pacific northwest. University of Washington

Press, Seattle, Washington.

Ivanovskaya, O. I. 1977. Aphids of western Siberia, Volume II. USSR Academy of Science, “Science”

Publishers, Novosibirsk.

McNeill, J. 1981. Apera, silky-bent or windgrass, an important seed genus recently discovered in

Ontario, Canada. Can. J. Plant Sci., 61: 479-485.

Muddathir, K. 1965. A new species of Holcaphis (Homoptera: Aphididae) together with a key to

the British Species. Ann. Mag. Nat. Hist., 8: 477-485.

Saha, D. C., M. A. Jackson & J. M. Johnson-Cicalese. 1988. A rapid straining method for detection

of endophytic fungi in turf and forage grasses. Phytopath., 78: 237-239.

SAS Institute. 1985. SAS user’s guide: statistics. SAS Institute, Cary, N.C.

Smith, C. F. & C. S. Parron. 1978. An annotated list of Aphididae (Homoptera) of North America.

North Carolina Agricultural Experiment Station Technical Bulletin, 255.

Zhang, G.-X., W.-Y. Zhang & T.-S. Zhong. 1991. A review of Diuraphis Aizenberg with descriptions

of two new species (Homoptera: Aphidoidea). Scientific Treatise on Systematic and Evolutionary

Zoology, 1: 121-133.

Received 22 April 1991; accepted 13 September 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 62-63, (1992)

Scientific Note

NEW HOST, BAUHINIA VARIEGATA L., AND

NEW LOCALITY RECORDS FOR

CARYEDON SERRATUS (OLIVIER) IN THE NEW WORLD

(COLEOPTERA: BRUCHIDAE: PACHYMERINAE)

The peanut (groundnut) bruchid, Careydon serratus (Olivier), is native to the

tropics and subtropics of the Old World and attacks seeds of tamarind, Tamarin-

dus indica L. and peanut, Arachis hypogaea L. (Davey, P. M. 1958. Bull. Entomol.

Res., 49: 385-404). This species has been introduced into the New World and

may become a serious pest if it becomes well established. In Africa, the species

is a serious pest of peanuts (Davey 1958; Prevett, P. F. 1967. J. Stored Prod. Res.,

3: 267-268), but in the New World, so far it has been collected only on its primary

host, tamarind. However, Velez Angel, in the article “Tres plagas insectiles re-

cientemente detectadas en Antioquia. 1. El gorgojo del tamarindo, Caryedon ser¬

ratus (Olivier),” (Velez Angel, R. 1972. Rev. Facultad Nal. de Agronomia, Med¬

ellin, Colombia, 27: 71—74) and Johnson reported that this species is already well

established in South America and is a potential threat to stored peanuts there

(Johnson, C. D. 1986. Coleopt. Bull., 40: 264). Although tamarind is becoming

an increasingly important crop for juice production in the tropics, the occurrence

of Caryedon serratus on this plant is not yet considered serious. Johnson reported

Careydon gonagra [= Caryedon serratus] from Mexico (Johnson, C. D. 1966.

Pan-Pacif. Entom., 42: 36), and later (Johnson 1986) reported it from additional

localities in Mexico and from Venezuela, but did not give any collection localities

for Venezuela. Caryedon serratus has been established in Hawaii for a long time

(Bridwell, J. C. 1920. Proc. Hawaiian Entomol. Soc., 4: 403-409).

Several new locality records and a new host record, Bauhinia variegata L., in

Mexico are reported here. It is the first time this species has been collected from

a host other than tamarind in the New World, including Hawaii. The new host

could be important for the successful dispersal of this introduced seed beetle, and

an indication that the species is becoming well established in the New World. It

could eventually become a serious pest on both tamarind and peanuts. Because

B. variegata is a commonly cultivated ornamental plant in the New World tropics,

introduced from Asia, this plant could function for dispersal of the insect.

New Host and Locality Records. — (Records from Hawaii are not included, depositories follow in

parentheses). MEXICO. BAJA CALIFORNIA SUR: La Paz, 4-9 Apr 1967, J. Chemsak & M. Mich-

elbacher (C. D. Johnson). DURANGO: 3 km W of El Palmito, 2-3 Oct 1976, E. Giesbert (Nat. Hist.

Mus. Los Angeles). NAYARIT: San Bias, 14 Jul 1960, R. B. Loomis & J. Maris (LBS). OAXACA: 47

km W of Tapanatepec, 27 May 1983, L. O’Brien & G. B. Marshall (CWO). SAN LUIS POTOSI:

Tamazunchale, ex. seeds of Bauhinia variegata, 22 May 1990 (emerged 10 Jul 1990), Jan A. Nilsson

(J. A. Nilsson). HAITI. Port-au-Prince, 15-23 Jan 1923 (Am. Mus. Nat. Hist.). ST. CROIX. H. A.

Beatty (Mus. Comp. Zool.). VENEZUELA. Puerta Negra, Tocoron, 8 Mar 1978, ex. tamarindo (Univ.

Centr. Venez.). ARAGUA: Maracay, El Limon, 1 May 1983, ex. tamarindo (Bordon); 14 May 1976,

1992

SCIENTIFIC NOTE

11 Feb 1977, 6 Feb 1985, G. Yepez, ex. tamarindo (Univ. Centr. Venez.). CARABOFO: Maziata, 25

Feb 1965, D. Gonzales, ex. tamarindo (Univ. Centr. Venez.). ZULIA: Maracaibo, 28 Jan 1979, B.

Tambrano, ex. tamarindo (Univ. Centr. Venez.).

Jan A. Nilsson and Clarence Dan Johnson, Department of Biological Sciences,

Northern Arizona University, Flagstaff, Arizona 86011-5640.

Received 26 February 1991; accepted 2 June 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 63, (1992)

Scientific Note

HEIM BRA OP AC A (ASHMEAD)

(HYMENOPTERA: EURYTOMIDAE)

DISCOVERED IN WASHINGTON 1

During 1990, a specimen of Heimbra opaca (Ashmead) was collected in south¬

eastern Washington. The site is a grassy slope with scattered forbs and shrubs

along the north side of the Snake River. This is the first report of H. opaca and,

thus, the subfamily Heimbrinae, from Washington. The specimen was identified

by the authors using key criteria (Burks, B. D. 1971. Trans. Am. Entomol. Soc.,

97: 1-87) and comparison with previously determined specimens.

This species was previously reported from California, Arizona, New Mexico,

Utah, Colorado, Kansas, Montana (Burks, B. D. 1979. In Krombein, K. V. et al.

Cat. Hymen. Am. N. of Mex., Vol. 1: 846), and Idaho (Johnson, J. B. & T. D.

Miller. 1987. Pan-Pacif. Entomol. 63: 324). Thus, H. opaca is widespread in arid

and semi-arid areas of the western U.S. However, it remains rarely collected and

biologically virtually unknown. The four collection sites in Idaho and Washington

known to the authors are all mixed grasslands, as described above, on sandy or

loess soils.

Material Examined. —WASHINGTON. WHITMAN Co.: 11.2 km (7 mi) E of Wawawai, 26 Jul

1990, sweeping vegetation, L. M. Wilson collector (specimen deposited: Maurice T. James Collection,

Department of Entomology, Washington State University, Pullman, Washington 99164-6432).

James B. Johnson and Linda M. Wilson, Department of Plant, Soil and En¬

tomological Sciences, University of Idaho, Moscow, Idaho 83843-4196.

Received 28 February 1991; accepted 8 May 1991.

1 Published with the approval of the director of the Idaho Agricultural Experiment Station as

Research Paper 91714.

PAN-PACIFIC ENTOMOLOGIST

68(1): 64-65, (1992)

Scientific Note

HEMIHYALEA EDWARDSII (PACKARD)

(LEPIDOPTERA: ARCTIIDAE) IS THE HOST OF

PARADEJEANIA RUTILIOIDES (JAENNICKE)

(DIPTERA: TACHINIDAE) IN

CENTRAL COASTAL CALIFORNIA

Paradejeania rutilioides (Jaennicke) is a widespread species in the western Ne-

arctic, including Mexico. West coast populations, which range from Vancouver

Island to southern California, have more extensive black markings on the ochreous

abdomen and were designated (Amaud, P. 1951. Canad. Entomol., 83: 332) as

a subspecies, nigrescens Amaud. In California, this race occurs along the coast

and Coast Ranges from Humboldt to Monterey Counties, on the west slope of

the Sierra Nevada, and in the Transverse and Peninsular Ranges, to elevations

of 1900 m; nearly all collection records are from late August to November (Amaud

1951; University of California, Berkeley, specimens).

Paradejeania rutilioides nigrescens is the largest and most conspicuous tachinid

in California, yet its biology is poorly known. General and perhaps presumed host

records date back to the early 1900s (Essig, E. O. 1913. Injurious and beneficial

insects of California. Calif. State Comm. Hort. Mo. Bull. 2: 261; 1915, ibid, 2nd

ed. Suppl. 4: 330), when Paradejeania was stated to feed on “caterpillars” and

“caterpillars of various species,” respectively, without specific taxa given.

Amaud reported a host of this fly to be an arctiid, Hemihyalea sp., based on a

specimen from Los Gatos, Santa Clara Co., California, reared by H. P. Allmendiger

and R. Maddux (Amaud, P. 1974. Pan-Pacif. Entomol., 50: 93). Because there

is only one species of Hemihyalea known in the area, H. edwardsii (Packard), this

is presumbly Amaud’s host record. The following observations on a rearing lot

that produced three examples of the fly and one of the moth agree with this

assumption. The only other reared specimen of P. r. nigrescens that we have seen

is one that emerged from leaf litter collected beneath Quercus agrifolia Nee in

Golden Gate Park, San Francisco, by P. A. Opler (JAP 67G16). The duff was

procured 21 Jul 1967, evidently containing a puparium, and the fly eclosed 6 Sep

1967 (UCB).

During a Lepidoptera survey trip to Big Creek Reserve in coastal Monterery

Co., California, 5 Jun 1990, we noticed several large woolybears climing the main

tmnk of a mature Q. agrifolia at dusk. Six were collected from heights of 2.4 to

3.0 m and others could been seen further up. There was a large tree hole in this

tmnk that contained arctiid exuvia and copious frass. Apparently, the Hemihyalea

larvae migrate up to the foliage to feed at night and retreat to a shelter during the

day. This explains why we have never found larvae of this arctiid on Q. agrifolia

during diurnal survey for lepidopterous larvae at numerous localities in the region.

We also found one larva of H. edwardsii at night on Q. agrifolia at a higher

elevation site at Big Creek in April (JAP 90D64).

1992

SCIENTIFIC NOTE

The six larvae collected in June were transported to Berkeley for rearing. Five

produced cocoons; one larva died prior to, and one following cocoon construction.

A large tachinid puparium appeared in each of three of the cocoons in late June

and July after emergence of the maggots. Three specimens of P. rutilioides ni-

grescens emerged 12 Jul to 30 Aug and one H. edwardsii on 17 Sep.

It is clear that Hemihylea serves as a host for Paradejeania, and the coincidental

autumnal flight periods of the two insects suggest that this moth is the only host

in this region, because we do not have any other large fall-flying, univoltine moth

in the coastal areas inhabited by P. rutilioides. Although we still do not know how

this tachinid infects its host, some observations provide clues for a possible answer.

Amaud observed females of this fly that bore dozens of active maggots in uturus,

in December, 1966, in San Francisco at the same site where Opler’s Paradejeania

was reared the following September (Amaud, P. 1968. Pan-Pacif. Entomol., 44:

85). This suggests that this tachinid, at least the females, can survive for a long

time, into mid-winter when H. edwardsii presumably has entered early larval

stages. Considering that the tachinid is larviparous and diurnal and the arctiid

larvae are nocturnal, it seems difficult to explain the high parasitism rate of larvae

we encountered. Hemihyalea larvae likely are widely dispersed during feeding in

the expansive canopy of Q. agrifolia. The female tachinid produces hundreds, or

even thousands, of larvae and the first instars may be distributed generally on the

foliage, where they would have to be capable of surviving until arrival of the host

caterpillars. Alternatively, P. rutilioides may be able to locate resting groups of

larvae and larviposit at the retreat either near, or on, the arctiid larvae.

The moths are abundant in late September and October, often 20-30 per black-

light trap sample, with a few worn examples persisting into November. The

conspicuous fly visits flowers of Eriogonum, Haplopappus, Aster, etc. (numerous

on introduced ivy, Hedera, growing at the Gatehouse), in late October and No¬

vember. None was observed during late September and early October when adults

of the arctiid are most numerous.

Records. —All the following data are from: CALIFORNIA. MONTEREY Co.: Big Creek Reserve:

H. edwardsii : HQ area 0-10 m, coastal scrub, 3^1 Oct 1985, 27-28 Sep 1987, 29 Oct 1989, J. W.

Brown, J. A. Powell; Gatehouse, 18 Oct 1988, J. Smiley; Trail to Redwood Camp 80 m, 27 Sep 1987,

J. A. Powell; Devils Creek Flat 120 m, 27 Sep 1987, J. W. Brown; South Ridge Rd, 220 m, 7 Nov

1988, 29-30 Oct 1989, and larvae 5 Jun 1990, r.f. Quercus agrifolia, emgd. 17 Sep 1990 (JAP 90F12)

Y.-F. Hsu, J. A. Powell; South Ridge Rd, 450-500 m, 29-30 Oct 1989; South Highlands, 675 m,

larva 27 Apr 1990, r.f. Q. agrifolia, emgd. 12 Sep 1990 (JAP 90D64) Y.-F. Hsu.

P. rutilioides nigrescens: HQ area 0-10 m, coastal scrub, 30 Oct 1989; grassland above Brunette

Crk, 180-240 m, 31 Oct 1989, J. A. Powell; South Ridge Rd, 220 m, 5 Jun 1990, r.f. Hemihyalea

edwardsii larvae, emgd. 12 Jul, 20, 31 Aug 1990 (JAP 90F12) Y.-F Hsu, J. A. Powell; South Highlands,

625-750 m, 30 Oct 1989; vie. French Camp, 750-800 m, manzanita-pine-oak woods, 6-8 Nov 1989,

J. A. Powell.

Acknowledgment. — We thank John Smiley for cooperation with facilities and

field work at the Big Creek Reserve and Paul H. Amaud, Jr. for comments on

the manuscript.

Yu-Feng Hsu and J. A. Powell, Department of Entomological Sciences, Uni¬

versity of California, Berkeley, California 94720.

Received 22 March 1991; accepted 8 May 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 66-68, (1992)

Scientific Note

LOW SUSCEPTIBILITY OF OVERWINTERING MONARCH

BUTTERFLIES TO BACILLUS THURINGIENSIS BERLINER

The large winter aggregations of the adult monarch butterfly, Danaus plexippus

L. (Danaidae: Lepidoptera), are spectacular phenomena that occur among selected

groves along the California coastline and in the high mountains of Mexico. The

overwintering areas in Mexico are periodically infested by larvae of Evita hyali-

naria (Grossbeck) (Geometridae: Lepidoptera), which defoliate the oyamel fir

trees, Abies religiosa Lindley, used by the butterflies. Foliage protection can be

attained with the use of Bacillus thuringiensis Berliner, but the application of this

biotic agent may inadvertently affect the monarch butterflies overwintering in the

sprayed region. Brower expressed concern over the widespread application of B.

thuringiensis because of its potential negative effects on the monarch butterflies

(Brower, L. 1986. Atala, 14: 17-19). He felt that large scale spraying of this biotic

agent should be avoided near an overwintering site unless the procedure could

be determined as safe or of minimal risk to the butterflies. Because the suscep¬

tibility of adult monarch butterflies to B. thuringiensis is unknown, we tested this

bacterium against overwintering butterflies under laboratory conditions.

Butterflies were collected from a central coast wintering site located in Oceano,

California (Leong, K. L. H. 1990. Ann. Entomol. Soc. Am., 83:906-910) and held

in cages without water at room temperature and humidity (21° C ± 0.25 SE, 42%

RH ±1.1 SE) for two days before treatment. The butterflies (10 insects per cage

and three cages per treatment) were initially subjected to twice the recommended

concentration of commercial B. thuringiensis [kurstaki] (Javelin®, 1.9 liters/3758

liters of H 2 0 [16,000 Spodoptera units/mg]) or to twice the recommended con¬

centration of dead transgenic Pseudomonas fluorescens (Trevisan) Migula con¬

taining endotoxin crystals (MVP®, 1.9 liters/3758 liters of H 2 0 [10,000 Dia-

mondback units/mg]). Control groups were provided water only. Adult monarchs

were exposed to B. thuringiensis by two ways: (1) spraying an aqueous suspension

onto the leaves of Eucalyptus sp. (tree species commonly used by overwintering

butterflies in California) within the cages until runoff (approximately 16 ml), and

(2) placing an aqueous suspension in petri dishes (50 ml) for 24 h.

The adults imbibed the preparation almost immediately after exposure. Five

days after exposure, the adults showed a low mortality to both Javelin® and

MVP®. The butterflies exposed to Javelin® spray, however, had a significantly

higher mortality (P < 0.05) than those sprayed with MVP® (20% ± 7 SE for

Javelin® vs 3% ± 3 SE for MVP®). The same relationship, but not significantly

different, was exhibited between the two groups of butterflies exposed to the

inocula in petri dishes (7% ± 5 SE for Javelin® vs 3% ± 3 SE for MVP®). None

of the control insects died during the study period. The results suggest that the

butterflies were slightly more susceptible to B. thuringiensis preparation than to

the transgenic P. fluorescens.

1992

SCIENTIFIC NOTE

To confirm the results of the B. thuringiensis preparation, another test was

conducted with Javelin® at the recommended (0.9 liters/3758 liters of H 2 0) and

at twice the recommended concentrations by spraying or by placing the aqueous

suspension in petri dishes (10 insects per cage, three cages per treatment). The

control adults were exposed to water only. The butterflies again exhibited low

total mortality rates at both the recommended and twice the recommended con¬

centration rates (7% vs 7% spray; 3% vs 7% petri dishes). The total mortality

among the control insects was zero for the spray and 3% for petri dish treatment.

Bacillus thuringiensis was isolated from the hemocoel of the dead butterflies

treated with Javelin, but not from the controls.

The fecal droppings of butterflies treated with twice the recommended concen¬

trations were collected daily and placed in 5 ml of sterile water and agitated. A

loopful of the suspension was then streaked onto nutrient agar and incubated at

25° C. Bacillus thuringiensis isolates were determined by colony growth charac¬

teristics and by microscopic examination (400 x). The bacterium was recovered

daily throughout the five day holding period. Fewer colonies (based on qualitative

observations) of B. thuringiensis were isolated after five days than after the first

two days. The bacterium was also recovered from the gut contents of the surviving

adults after the five-day holding period.

The overwintering butterflies, under laboratory conditions, showed low sensi¬

tivity to the B. thuringiensis used, both at the recommended and at twice the

recommended concentrations for control of lepidopterous larvae. The results

suggest that the application of this biotic agent within or near the butterfly’s winter

habitat presents a minimal threat to their survival. Because a small percentage of

the butterflies did succumb to this bacterium or its product (endotoxin), however,

the use of B. thuringiensis near the butterfly’s overwintering site should be min¬

imized.

Assuming that E. hyalinaria, the defoliator of fir trees, is equally susceptible

to B. thuringiensis and transgenic P. fluorescens, the latter may be a better choice

for controlling this insect. Our data suggest that the butterflies were less susceptible

to the endoxotin alone than to a bacterial spore endotoxin mixture. To ensure

another level of safety for the roosting butterflies from possible drift of the bioin¬

secticide, a spray-free buffer zone could be established around the overwintering

site. This buffer zone, however, may be subjected to defoliation by E. hyalinaria,

and could affect the roosting site. The possible threat of introducing the bacterium

or its endotoxin into the protected aggregation sites by affected adults (via fecal

droppings) exposed to the treated areas is minor due to the dilution effect, the

poor survivability of the spores on the leaf surfaces (Leong, K. et al. 1980. Environ.

Entomol., 9: 593-599) and the low adult susceptibility to B. thuringiensis. Another

possible threat is the contamination of the monarch butterfly eggs with B. thu¬

ringiensis deposited by affected adults (Ali, A. A. & T. F. Watson. 1982. J. Econ.

Entomol., 75: 596-598). Presumably, the overwintering butterflies will have elim¬

inated most of the bacterium before egg deposition occurs. The monarch larvae

sensitivity to B. thuringiensis from adults remains unknown and requires further

study.

Although our study suggests that B. thuringiensis presents a minimal threat to

the adult monarch butterflies, the susceptibility of the populations while in Mexico

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(1)

still needs to be investigated before wide scale spraying is employed near their

winter aggregation site.

Acknowledgment. — Authors thank the Mycogen Corporation, San Diego, Cal¬

ifornia for providing the Bacillus thuringiensis materials (Mycogen MVP®) used

in this study.

Kingston L. H. Leong, 1 Michael A. Yoshimura, 1 & Harry K. Kaya, 21 Biological

Sciences, California Polytechnic State University, San Luis Obispo, California

93407; department of Nematology, University of California, Davis, California

95616.

Received 5 April 1991; accepted 15 June 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 68-69, (1992)

Scientific Note

NEW DISTRIBUTIONAL RECORDS FOR SOME

CANDIDATE SPECIES OF LYTTA IN CALIFORNIA

(COLEOPTERA: MELOIDAE)

Five Californian species of Lytta blister beetles (Coleoptera: Meloidae) are

candidates for listing as endangered (i.e., a species that is likely to become extinct)

by the U.S. Fish and Wildlife Service. These species [L. hoppingi Wellman, L.

insperata (Horn), L. moesta (Horn), L. molesta (Horn), and L. morrisoni (Horn)]

have not been listed due to a lack of biological information. Developments in

California’s central valley and surrounding foothills continue to impact them and

their habitat because of three factors: our knowledge of their biology is limited,

a lack of adequate survey methods, and the beetles not being fully protected by

the Endangered Species Act (1973).

Recently, specimens of four of these candidate species were found in the R. S.

Wagner Collection at the Tulare County Agricultural Commissioner’s/Sealer’s

Office, Visalia, California. The specimens represent new distributional records for

three species, having been collected in the 1930s but overlooked by researchers.

For two of these species, an additional, more recent, distributional record is

reported. We present this information (see Records section) to improve and update

the distributional data on Lytta; to aid federal, state, and county regulatory agen¬

cies in their development, management, and protection efforts; and to encourage

researchers to utilize the Wagner Collecton and to study the endangered inver¬

tebrate fauna. Distributional information on Lytta has been summarized (Selan-

der, R. B. 1960. Illinois Biol. Monographs, 28) and recently updated (California

Department of Fish and Game. 1991. Natural Diversity Data Base, computer

data base of sensitive species. Sacramento, California).

Biological information for the five candidate species of Lytta is almost non¬

existent. Lytta molesta has been collected on Lupinus (Leguminosae) feeding on

1992

SCIENTIFIC NOTE

its flowers and seed pods (personal observations) and on Erodium cicutarium (L.)

(Geraniaceae) (Selander 1960). Like most meloid beetles, Lytta parasitize the nests

of wild bees, feeding on provisions or immature stages of the bees (Selander 1960).

Bee genera that are parasitized by various other Lytta species include: Anthophora

Latreille, Diadasia Patton, Emphoropsis Ashmead, Ptilothrix Smith (all Hyme-

noptera: Anthophoridae), Andrena subgenus Onagrandrena Linsley & MacSwain

(Hymenoptera: Andrenidae) and Colletes Latreille (Hymenoptera: Colletidae)

(Hurd, P. D. Jr. 1979. In Krombein et al. (eds.). Cat. of Hymenoptera in America

north of Mexico. Vol. II, Smith. Instit. Press, Wash., D.C., pp. 1199-2209; Se¬

lander 1960).

The fourth candidate species in the Wagner Collection, L. molesta, is repre¬

sented by three females and five males from a location previously reported by

Selander (1960) as Lanes Bridge, Fresno Co., California. These specimens, though

not providing new distributional information, have taxonomic value and dem¬

onstrate the merit of the Wagner Collection.

Because the candidate Lytta are very similar to each other, similar to other

Lytta species, and more than one species may occur at a site, we urge researchers

to collect voucher specimens and have them identified by a taxonomist to validate

records. We hope that this information encourages researchers or agencies (state,

federal, or private) to study or provide the funding necessary to determine the

biological status of these species.

Records.—Lytta hoppingi Wellman. New records: CALIFORNIA. KERN Co.: Taft, 29 Apr 1978,

W. H. Tyson (California Department of Fish and Game. 1991) [species determined by J. D. Pinto].

TULARE Co.: Ducor, 25 Mar 1934, 8 males and 8 females. Historical distribution: CALIFORNIA.

FRESNO Co.: Coalinga; TULARE Co.: Visalia; Delano.

Lytta moesta (Horn).—New records: CALIFORNIA. KERN Co.: Arvin, 29 Mar 1931, 2 females.

TULARE Co.: Springville, 28 May 1933, 1 female; 23 Jun 1936, 1 pair (in copulation). Historical

distribution: CALIFORNIA. FRESNO Co.: Friant. KERN Co.: Edison. MADERA Co.: Kismit. SANTA

CRUZ Co.: Santa Cruz. STANISLAUS Co.: Manteca; Modesto; Ripon; Westly. TULARE Co.: exact

location unknown (county label only); Kaweah; Potwisha, Sequoia National Park.

Lytta morrisoni (Horn).—New records: CALIFORNIA. FRESNO Co.: Panoche Road 13.1 km (8.2

miles) W of Interstate 5, 21 May 1978, F. G. Andrews (California Department of Fish and Game.

1991) [species determined by J. D. Pinto]. TULARE Co.: Plano, 1 May 1939, 1 male and 1 female.

Historical distribution: CALIFORNIA. FRESNO Co.: Coalinga. KERN Co.: exact location unknown

(county label only); Edison. TULARE Co.: Kaweah; Tipton; White River (south of). One data locale

lists only “southern California.”

Acknowledgment. — We thank the Kings River Conservation District for funding

this publication, J. D. Pinto (University of California, Riverside) for confirming

the identification of L. moesta; the former, F. G. Andrews, and W. H. Tyson

(both California Department of Food and Agriculture) for providing additional

information on the California Department of Fish and Game (1991) records; and

D. J. Burdick (California State University, Fresno) and two anonymous reviewers

for helpful editorial comments.

Jeffrey A. Halstead and Robert D. Haines, Kings River Conservation District,

4886 E. Jensen Avenue, Fresno, California 93725 and Tulare County Agricultural

Commissioner’s/Sealer’s Office, County Civic Center, Main and Woodland Drives,

Visalia, California 93291, respectively.

Received 20 February 1991, accepted 3 June 1991.

PAN-PACIFIC ENTOMOLOGIST

68(1): 70-71, (1992)

Scientific Note

RHOPALOSIPHUM RUFIABDOMINALIS (SASAKI) AND

APHIS ARMORACIAE CO WEN

(HOMOPTERA: APHIDIDAE) CONFIRMED ON

WHEAT IN IDAHO

Two species of aphids were found on wheat in Idaho for the first time during

October, 1990. Both were found on winter wheat ( Triticum aestivum L.) cv.

‘Stephens’ at the SW Idaho Research & Extension Center at Parma (Canyon

County). Neither of these species is likely to become a significant pest as both

have been in Idaho for at least five years and are only now being discovered on

wheat despite intensive surveys of this crop since 1986. However, these additions

to the wheat aphid fauna will further complicate species identifications of aphids

found on cultivated cereals by increasing the number of taxa that must be rec¬

ognized. Additionally, the genus Rhopalosiphum includes two common cereal

pests, R. maidis (Fitch) and R. padi (L.), plus R. insertum (Walker). Separation

of the latter three species requires detailed microscopic examination.

Rhopalosiphum rufiabdominalis (Sasaki) was collected between 16 Oct-19 Nov

1990 on winter wheat roots. Rhopalosiphum rufiabdominalis is permanently par-

thenogenetic on roots of rice and many other species of Gramineae in most parts

of the world, though holocyclic populations overwintering on Prunus spp. have

also been reported. It colonizes the roots of some dicots, especially Solanaceae

and can be a serious pest on plants in hydroponic systems (Blackman, R. & V.

Eastop. 1984. Aphids on the world’s crops: an identification and information

guide. John Wiley & Sons, New York; Blackman, R. et al. 1987. World perspec¬

tives on barley yellow dwarf. Proc. of intem’l. workshop, 6-11 July 1987, Udine,

Italy. Dipartimento Cooperatizione Alio Sviluppol International Maize and Wheat

Improvement Center (CIMMYT), pp. 197-214). It is known to vector barley

yellow dwarf virus (Jedlinski, H. 1981. Plant disease, 65: 975-978; Johnstone,

G. et al. 1987. World perspectives on barley yellow dwarf. Proc. of intem’l.

workshop, 6-11 July 1987, Udine, Italy. Dipartimento Cooperatizione Alio Svi¬

luppol International Maize and Wheat Improvement Center (CIMMYT), pp. 228-

239; Paliwal, Y. 1980. Can. J. PI. Path., 2: 90-92).

Rhopalosiphum rufiabdominalis was collected prior to these field collections in

suction traps at Parma, Burley, Caldwell and Craigmont. There are no previously

published records of collections from host plants in Idaho for R. rufiabdominalis.

A field survey of cereal aphids between 16 Oct and 5 Dec 1990, on winter wheat

showed decreasing numbers of R. rufiabdominalis as the season progressed (Table

1). This, along with late season suction trap collections and the presence of alatoid

nymphs late in the season indicate the possibility of holocyclic overwintering in

Idaho.

Aphis armoraciae Co wen was collected on 12 Oct 1990 from winter wheat using

a Berlese funnel, and on 3 Dec 1990 on bluebunch wheatgrass T-2950 ( Agropyron

spicatum (Pursh) Scribner & Smith) at Parma, Idaho. Aphis armoraciae infests

1992

SCIENTIFIC NOTE

Table 1. Rhopalosiphum rufiabdominalis (Sasaki) population on winter wheat ( Triticum aestivum

L.) cv. ‘Stephens’ at Parma, Idaho, 16 Oct-19 Nov 1990.

Aphid Stage

Total

Sample date*

Nymph 1

Nymph 2-4

Alatoid

Apterae

Alatae

16 Oct

22 Oct

8 Nov

19 Nov

5 Dec

a 20 tillers per sample and 10 samples.

roots and sometimes the aerial parts of plants in several families including Com-

positae, Cruciferae, Umbelliferae, and Gramineae (maize, wheat) (Blackman &

Eastop 1984). Aphis armoraciae was previously collected from Bear Lake Co. on

raspberry, Bingham Co. on potato, Canyon Co. on maize and roots of Sisymbrium

altissimum L., Caribou Co. on potato, cabbage and radish, Franklin Co. on maize

and Aster sp., Oneida Co. on Chrysothamnus nauseosus (Pallas) Britton and Teton

Co. on potato (Gittins, A. et al. 1976. An annotated list of the aphids of Idaho

(Homoptera: Aphididae). Rec. Bull., 95. College of Agric. Univ. of Idaho). Aphis

armoraciae was also collected in suction traps at various areas in Idaho for several

years from 1985-1990.

Voucher specimens from our collections from wheat are on deposit at the

University of Idaho, SW Idaho Research and Extension Center, Parma, Idaho.

Material Examined.—Rhopalosiphum rufiabdominalis : IDAHO. CANYON Co.: Parma, 22 Sep

1985, and 1,7, 15, 29 Sep 1987, suction trap, 1 specimen on each date; Caldwell, 21 Jul 1990, suction

trap, 1 alate. CASSIA Co.: Burley, 18 Aug 1986, suction trap, 1 alate. LEWIS Co.: Craigmont, 25

Aug 1990, suction trap, 1 alate.

Aphis amoraciae: IDAHO. CANYON Co.: Parma, 12 Oct 1990, winterwheat, Berlese funnel, 7

specimens; Parma, 3 Dec 1990, bluebunch wheatgrass T-2950, 2 oviparae; Parma, 16 and 22 Oct, 8

and 19 Nov, 5 Dec 1990, Triticum aestivum ‘Stephens,’ total 81 specimens of various stages (see

table).

Acknowledgment. — We thank D. Allison, Washington State University, for pro¬

viding suction traps data, and J. P. McCaffrey, F. W. Merickel and T. M. Mowry

for reviewing the manuscript. Published with the approval of the Director of the

Idaho Agricultural Experiment Station as Research Paper 91728.

Ali A. Al-Raeesi, Susan E. Halbert and James B. Johnson, Department of Plant

Soil & Entomological Sciences, University of Idaho, Moscow, Idaho 83843.

Received 12 April 1991; accepted 29 July 1991.

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Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons. New York.

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Volume 68

THE PAN-PACIFIC ENTOMOLOGIST

January 1992

Number 1

Contents

YOSHIDA, N. & H. KATAKURA—Evolution of oviposition habits in Aphodius dung beetles

(Coleoptera: Scarabaeidae). 1

HALBERT, S. E. & T. M. MOWRY—Survey of Myzus persicae (Sulzer) (Homoptera: Aphid-

idae) infestations on bedding plants for sale in eastern Idaho . 8

WELLS, J. D. & B. GREENBERG—Rates of predation by Chrysomya rufifacies (Macquart)

on Cochliomyia macellaria (Fabr.) (Diptera: Calliphoridae) in the laboratory: effect of

predator and prey development... 12

NEFF, J. L. & B. B. SIMPSON—Nest biology of Osmia {Diceratosmia) subfasciata Cresson in

central Texas (Hymenoptera: Megachilidae). 15

ARNAUD, P. H. Jr.-Obituary: James Wilson Tilden (1904-1988). 27

KAYA, H. K„ M. G. KLEIN, T. M. BURLANDO, R. E. HARRISON & L. A. LACEY-

Prevalence of two Bacillus popilliae Dutky morphotypes and blue disease in Cyclocephala

hirta LeConte (Coleoptera: Scarabaeidae) populations in California. 38

SCHOLL, A., R. W. THORP & E. OBRECHT—The genetic relationship between Bom-

bus franklini (Frison) and other taxa of the subgenus Bombus s.str. (Hymenoptera:

Apidae)... 46

HALBERT, S. E., B. J. CONNELLY & M.-G. FENG-Occurrence of Diuraphis {Holcaphis)

frequens (Walker) (Homoptera: Aphididae) on wheat, new to Idaho, and a key to North

American Diuraphis . 52

SCIENTIFIC NOTES

NILSSON, J. A. & C. D. JOHNSON—New host, Bauhinia variegata L., and new locality

records for Caryedon serratus (Olivier) in the New World (Coleoptera: Bruchidae: Pa-

chymerinae)... 62

JOHNSON, J. B. & L. M. WILSON— Heimbra opaca (Ashmead) (Hymenoptera: Eurytomidae)

discovered in Washington. 63

HSU, Y.-F. & J. A. POWELL— Hemihyalea edwardsii (Packard) (Lepidoptera: Arctiidae) is

the host of Paradejeania rutilioides (Jaennicke) (Diptera: Tachinidae) in central coastal

California. 64

LEONG, K. L. H., M. A. YOSHIMURA & H. K. KAYA—Low susceptibility of overwintering

monarch butterflies to Bacillus thuringiensis Berliner. 66

HALSTEAD, J. A. & R. D. HAINES—New distributional records for some candidate species

of Lytta in California (Coleoptera: Meloidae). 68

AL-RAEESI, A. A., S. E. HALBERT & J. B. JOHNSON— Rhopalosiphum rufiabdominalis

(Sasaki) and Aphis armoraciae Cowen (Homoptera: Aphididae) confirmed on wheat in

Idaho. 70

Announcement: Publications of the Pacific Coast Entomological Society. 72

Volume 68 April 1992 Number 2

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in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)

The Pan-Pacific Entomologist

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PAN-PACIFIC ENTOMOLOGIST

68(2): 73-96, (1992)

REVISION OF THE SPIDER BEETLE GENUS NIPTUS IN

NORTH AMERICA, INCLUDING NEW CAVE AND

PHOLEOPHILE SPECIES (COLEOPTERA: PTINIDAE)

Rolf L. Aalbu and Fred G. Andrews

Insect Taxonomy Laboratory,

California Department of Food and Agriculture,

Sacramento, California 95814

Abstract. — The genus Niptus is revised for North America. Four species of Niptus Boeildieu ( N .

giulianii NEW SPECIES, N. neotomae NEW SPECIES, N. sleeperi NEW SPECIES, and N.

arcanus NEW SPECIES) are described from the Great Basin area, southwestern Arizona, the

cape mountain region of Baja California, Mexico, and a California cave, respectively. Notes on

the biology of Niptus species, as well as Ptinus clavipes, are presented. A key is provided to the

species of Niptus found in North America. Phylogenetic considerations among Niptus, Pseudeu-

rostus, and Eurostus are discussed. Habitat conservation is stressed for species restricted to single

cave localities.

Key Words. — Insecta, Coleoptera, Ptinidae, Niptus, southwest United States, Mexico, biology,

caves

As a result of improved collecting techniques, such as overnight pitfall traps or

longer duration ethylene glycol (antifreeze) traps, and greater accessibility to pre¬

viously difficult to reach places, numerous specimens of small apterous beetles

are now available in collections. Most larval and adult Ptinidae feed on dried ''

plant and animal substances. Others have been recorded from dung. Many are

associated with mammals or birds and are often found in caves. Their biology,

including rearing methods of economically important species, is adequately cov¬

ered by Howe (1959).

One species, Niptus hololeucus (Faldermann), a stored product pest, is widely

distributed in the northern United States. Niptus kelleri (Brown) and N. hilleri

Reitter have previously been placed in the genus Pseudeurostus. One of these, N.

hilleri , is distributed widely, also in stored products (see Brown 1959: 629). Niptus

kelleri, known only from the type locality, was not examined.

Because all genera of ptinids are flightless (except for certain Ptinus ), the method

found most effective in capturing pholeophilic Ptinidae is the use of numerous

dry plastic “punch cup” containers as pit traps, especially near, or at, the entrance

to rodent burrows. These traps are set in the late afternoon and collected early

the next morning. This permits collection of live adult specimens for rearing and

provides additional biological information (substrate type, etc.). Adults are also

collected at night with the use of headlamps or lanterns to illuminate surface areas.

In 1978-1979,a year-long trapping survey of the Coleoptera of Mitchell Caverns

was conducted using ethylene glycol pitfall traps (Aalbu 1990). Mitchell Caverns

are located on the eastern slopes of the Providence Mountains (San Bernardino

County), California. A new species of Niptus was found to be endemic to one

cave.

Abbreviations. — The following abbreviations are used to denote the institutions

that loaned material: CASC, California Academy of Sciences, San Francisco,

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

California; CISC, University of California, Berkeley, California; CNCI, Canadian

National Collection, Ottawa, Ontario, Canada; CSLB, California State University,

Long Beach; FMNH, Field Museum of Natural History, Chicago, Illinois; KWBC,

Kirby W. Brown Collection, Stockton, California; MCZC, Harvard University

Museum of Comparative Zoology, Cambridge, Massachusetts; OSUC, The Ohio

State University, Columbus, Ohio; SDMC, San Diego Museum of Natural His¬

tory, California; USNM, United States National Museum, Washington D.C.;

UAIC, University of Arizona, Tucson.

Niptus arcanus Aalbu & Andrews, NEW SPECIES

(Figs. 1, 17, 19, 24 and 25)

Types. — HOLOTYPE (female) and ALLOTYPE (male): CALIFORNIA. SAN

BERNARDINO Co.: Providence Mountains State Recreation Area, Mitchell Cav¬

erns, el. 1340 m, El Pakiva Cave, 26 Aug-31 Dec 1978, Ethylene glycol pitfall

trap near Neotoma nest, #6. Type deposited in California Academy of Sciences

Collection. PARATYPES: CALIFORNIA. SAN BERNARDINO Co.: Providence

Mountains State Recreation Area, Mitchell Caverns, el. 1340 m, El Pakiva Cave,

26 Aug 1978 to 31 Dec 1978 trap #6; 17 Mar 1979 to 16 Jun 1979, trap #3 (4);

17 Mar 1979 to 16 Jun 1979, trap #4 (19); 17 Mar 1979 to 16 Jun 1979, trap #6

(34); 17 Mar 1979 to 16 Jun 1979 (1); 27 May 1978 to 26 Jul 1979, trap #5 (28);

31 Dec 1978 to 17 Mar 1979, trap #5 (1); 31 Dec 1978 to 17 Mar 1979, trap #6

(21); 8 May 1981 to 10 Aug 1981 (11), R. L. Aalbu, Ethylene glycol pitfall trap

near Neotoma nest. Paratypes deposited in USNM, CDFA, CISC, CASC, RLAC,

OSUC.

Description.— Female (holotype). Integument red-brown, elytra shiny; length 3.3 mm. HEAD with

surface vestiture of closely appressed, spatulate, scale-like setae with few longer fine setae on apical

margin of clypeus; antennal fossae with dorsal border not carinate, not laterally elevated; eyes minute,

three facets at minimum width, narrowly oval; antenna relatively long, slender, ratio of segment

lengths 14:11:10:10:9:9:9:9:10:10:18. PRONOTUM with surface sculpture of rugose, deep punctures

posteriorly forming moderately dense, fine tubercles; surface vestiture of one type, stout, arched,

recumbent setae; setae dense at anterior margin, at transverse row of four large tufts; tufts equal in

size, positioned near midlength. ELYTRA with surface smooth, shiny, strial punctures fine, nearly

obsolete; vestiture of two types, nearly equal in length; first consisting of short moderately slender,

erect, spatulate setae positioned in rows at regular distances along first to seventh intervals; second

arched, recumbent, moderately slender setae positioned in rows at elytral striae and elytral intervals;

setae short, dense at elytral margins. VENTRAL SURFACE: Sterna: ratio of segment lengths 17:19:

15:5:19; sternal surface vestiture short, golden, closely appressed, spatulate, scale-like setae intermixed

with sparse, slightly longer, less spatulate setae; fifth visible abdominal stemite with medial apical

area with closely packed postero-directed, semi-erect setae forming a rounded tubercle-like structure.

LEGS slender, femora moderately long, capitate, metafemora bent near apex; tibia slender; femoral

vestiture of dense, golden, short, appressed, scale-like setae only varying slightly in length; tibiae with

similar vestiture except protibiae with dense, slightly longer, slender, golden setae on lower margins,

mesotibiae with dense, slightly longer, slender golden setae on lower margins, on apical one-half of

outer margins; metatibiae with few sparse, slightly longer, golden setae on lower margins. Ratios of

segment lengths: prothoracic legs, 50:49; mesothoracic legs, 54:52; metathoracic legs, 60:65; protarsi,

10:6:6:6:9; mesotarsi, 12:7:6:6:9; metatarsi, 15:8:7:7:10.

Male (allotype).—Similar to holotype but smaller, approximate length 2.9 mm. Fifth visible ab¬

dominal stemite with medial apical area with setae only slightly less appressed, slightly longer than

rest of sternal setae; without tubercle-like structure.

Diagnosis. — The following combination of characters will serve to separate N.

arcanus : Head with eyes minute (Fig. 24) and antennal fossa with dorsal border

1992

AALBU & ANDREWS: REVISION OF NIPTUS

Figures 1-2. Figure 1. Niptus arcanus, habitus (stereo pair). Figure 2. Niptus neotomae, habitus

(stereo pair).

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

Figures 3-4. Figure 3. Niptus giulianii, habitus (stereo pair). Figure 4. Niptus abditus, habitus

(stereo pair).

1992

AALBU & ANDREWS: REVISION OF NIPTUS

Figures 5-6. Figure 5. Niptus ventriculus, habitus (stereo pair). Figure 6. Niptus abstrusus, habitus

(stereo pair).

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

Figures 7-8. Figure 7. Niptus absconditus, habitus (stereo pair). Figure 8. Niptus sleeperi, habitus

(stereo pair).

1992

AALBU & ANDREWS: REVISION OF NIPTUS

Figures 9-10. Figure 9. Niptus hilleri, habitus (stereo pair). Figure 10. Niptus hololeucus, habitus

(stereo pair).

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Figures 11-12. Figure 11. Niptus giulianii, anterior aspect of head (stereo pair). Figure 12. Niptus

hilleri, anterior aspect of head (stereo pair).

1992

AALBU & ANDREWS: REVISION OF NIPTUS

Figures 13-14. Figure 13. Niptus ventriculus, anterior aspect of head (stereo pair). Figure 14. Fecal

pellets of Neotoma lepida Thomas showing feeding damage by Niptus arcanus.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

1 3 K U X 3 0

0U CDF A

r • • ' • . .1

13 KU XI 00

10 0. 0U C D F A

1 3 K U X 7 2

Figures 15-20. Figure 15. Niptus hololeucus, sterna, ventral view. Figure 16. Niptus ventriculus,

dorsal-apical aspect of head. Figure 17. Niptus arcanus, apical aspect of sterna, ventral view. Figure

18. Niptus ventriculus, dorsal-apical aspect of pronotum. Figure 19. Niptus arcanus, lateral aspect of

metafemur. Figure 20. Niptus giulianii, lateral aspect of metafemur.

not carinate or laterally raised. Pronotum with medial and lateral transverse

pronotal tufts equal in size; anterior margin of pronotum without long erect setae.

Elytra with erect setae on intervals one to five short, spatulate. Legs long, with

metafemur capitate, metatibia slender, slightly curved. Sexually dimorphic: female

with fifth visible abdominal stemite with medial apical area bearing closely packed

patch of postero-directed, semi-erect setae forming a rounded tubercle-like struc¬

ture.

1992

AALBU & ANDREWS: REVISION OF NIPTUS

.011 CDF A

1 3 K U X 7 2

1 3KU X4 00

13KU X4@0

Figures 21-24. Figure 21. Niptus hilleri, sterna, ventral view. Figure 22. Niptus ventriculus, lateral

aspect of eye. Figure 23. Niptus absconditus, lateral aspect of eye. Figure 24. Niptus arcanus, lateral

aspect of eye.

Niptus arcanus is most closely related to N. neotomae, sharing short spatulate

elytral setae. These species also lack long erect setae on pronotal margins as well

as being sexually dimorphic, characters also shared by N. abscondidus Spilman.

Niptus neotomae differs from N. arcanus in having shorter setae both on the

pronotum and elytra and in the configuration of the legs. In N. neotomae, the legs

are short and stout, the metafemora clavate; in N. arcanus, the legs are long and

slender, the metafemora capitate. N. arcanus and N. abscondidus Spilman also

share strongly reduced eyes.

Distribution. —(Fig. 25) This species is only known from the type locality, El

Pakiva Cave, Mitchell Caverns, Providence Mountains, San Bernardino County,

California.

Label Biological Notations. — Ethylene glycol pitfall trap near Neotoma nest,

dry pit traps.

Biological Notes. — There are a number of caves in the Providence Mountains

State Recreation Area. Mitchell Caverns, located at about 1340 m, actually refers

to two separate limestone caves, believed to be Miocene in origin. Both caves are

at about the same level, although one cave, El Pakiva, contains a large secondary,

lower chamber at the far south end, which is approximately 18 meters lower.

These caves were exploited as a tourist attraction in the 1930s. In 1970, to facilitate

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

Figure 25. Distribution of Niptus arcanus (solid diamond), Niptus giulianii (solid circles), Niptus

abditus (solid squares) and Niptus neotomae (solid triangle).

visitor tours, a tunnel was completed connecting the two caves. During the 1979

survey, trapping periods (intended to sample seasonal differences during an entire

year) were segregated into four series, averaging approximately three months (see

Aalbu 1990).

Although nine years had elapsed since the construction of the tunnel connecting

the two caves during the faunal survey, some species of troglophilic Coleoptera

were found to remain concentrated or even completely restricted to one cave.

Niptus arcanus was the best example. Close to 100% (292) of the specimens were

found in both the main section and the lower caverns of El Pakiva (one specimen

found near an entrance) but was entirely absent from Tecopa, the other connected

cave. This is also one of the few species to be found in numbers deep in the lower

caverns of El Pakiva. Specimens of Niptus were trapped in greater numbers in

the fall but were present in large numbers throughout the year. Since this survey,

1992

AALBU & ANDREWS: REVISION OF NIPTUS

Figure 26. Known geographical distribution of Niptus abstrusus (solid triangles), Niptus absconditus

(solid diamond), Niptus sleeperi (star in circle) and Niptus ventriculus (solid circles).

other caves (Medicine Cave, Cave of the Winding Stairs) and mines in the area

have been surveyed or partially surveyed for insects. N. arcanus was not found

in any of these.

Most of the food energy in Mitchell Caverns comes in with packrats (Neotoma

lepida Thomas). The rats bring organic materials, such as twigs, cacti, grass, leaves,

etc., collected outside into their nests. The packrats and other rodents, such as

mice, also leave fecal pellets, which are found sometimes in great numbers in the

caverns. Rodents nesting in the caverns are in most instances not found in the

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

deeper areas. From data gathered from an analysis of substrate composition near

each trap (Aalbu 1990), Niptus arcanus was found most abundantly in substrate

consisting of mostly fine cave dust, with few calcite and limestone pebbles and

rocks, and a small amount of organic matter or in the lower caverns area of very

fine, highly organic dust and conglomerate (dust-clay-rocks). Niptus arcanus was

not abundant in packrat nesting areas. However, it appears there is an association

with packrats.

A number of species of ptinids are known to breed in rat dung (Howe 1959),

but no larvae were trapped or found in the cave substrate. Close examination of

Neotoma droppings in the areas of Niptus abundance proved interesting. Most of

these droppings, although relatively few in numbers compared with packrat nest¬

ing areas in other parts of the cave, contained numerous cavities with diameters

approximately equal to Niptus specimens in size. No other insect in the area is

known to create similar cavities. It appears that the larvae, and possibly also the

adults, of this species feed on Neotoma pellets (Fig. 14). Unfortunately, attempts

to rear live adults on the dung were unsuccessful as adults died within a short

period of time.

An additional ptinid, Ptinus feminalis Fall, was also trapped in the caves. This

species has a wide geographical range. It is known to feed on dried vegetable

matter and animal substances. P. feminalis was found in both caves during the

survey. Most were found near the entrances.

Material Examined.. — 313 specimens (see types), from the type locality distributed as follows: 292

trapped during cave survey (see Aalbu 1990: table 6); 12 from substrate samples and nine collected

alive in pitfall traps 6 Jun-20 Jun 1988.

Niptus giulianii Aalbu & Andrews, NEW SPECIES

(Figs. 3, 11, 20 and 25)

Types. — HOLOTYPE (female): ARIZONA. COCONINO Co.: 6.3 km SE of

Moenkopi, sand dunes/ dry canyon, 31 Jul-1 Aug 1983, Rolf L. Aalbu coll., dry

overnight “punch cup” pitfall trap. ALLOTYPE (male): UTAH. UINTAH Co.:

16.1 km SSW of Vernal, 28-29 Jul 1983, Rolf L. Aalbu coll., dry overnight “punch

cup” pitfall trap. Holotype and allotype deposited in the collection of the California

Academy of Sciences. PARATYPES: ARIZONA. COCONINO Co.: 3.2 km S of

Moenkopi, 3 Jul 1972, F. Andrews & E. A. Kane [CDFA] (49); 1.6 km E of

Moenkopi, 31 Jul 1983 to 1 Aug 1983, R. L. Aalbu pitfall near Neotoma nest,

[RLAC] (9); 6.3 km SE of Moenkopi, sand dunes/dry canyon, 31 Jul 1983 to 1

Aug 1983, R. L. Aalbu, pit traps [RLAC] (161); 3.9 km S of Moenkopi, Moenkopi

Dunes, 17 Jul 1975, F. Andrews & A. Hardy, cereal bowl trap [CDFA] (11);

Moenkopi sand dunes, Mar 1983 to Sep 1983, D. Giuliani, antifreeze pit trap

[CDFA] (21); Waheap, 4.8 km NW of Lake Powell, 1 to 2 Aug 1983, R. L. Aalbu

[RLAC] (10); 19.3 km ENE of Tuba City, 31 Jul 1983 to 1 Aug 1983, R. L. Aalbu,

sand dunes/dry canyon, [RLAC] (25). CALIFORNIA. INYO Co.: Deep Springs

Valley Sand Dunes, 17 July 1975, [CDFA] (1); 17 Jun 1978 to 28 Sep 1978,

[CDFA] (1); 28 Sep 1979 to 15 Dec 1979, [CDFA] (1); 13 May 1980 to 29 Sep

1980; Deep Springs Valley Sand Dunes, (5000 ft), D. Giuliani, antifreeze pit trap,

[CDFA] (2). MONO Co.: Mono Lake, sand dunes, 17 Aug 1979 to 21 Nov 1979

[CDFA] (6); same locality, 21 Nov 1979 to 7 Jun 1980 [CDFA] (2); same locality,

1992

AALBU & ANDREWS: REVISION OF NIPTUS

7 Jun 1980 to 1 Sep 1980 [CDFA] (17), D. Giuliani, antifreeze pit trap; Mono

Lake, NE end, sand dunes, 15 Jun 1979 to 17 Aug 1979, D. Giuliani, antifreeze

pit trap, [CDFA] (56); Mono Lake, 1 km E of Sulfur Pond, #17, no date, J. H.

Harris, ethylene glycol pit trap [CDFA] (4). SANTA CRUZ Co.: Watsonville, May

1936, E. L. Kellogg [CDFA] (1). NEVADA. CHURCHILL Co.: Sand Mountain,

19 Jul 1977, D. Giuliani, UY light [CDFA] (1); Sand Mountain dunes, 16 Sep

1974, F. Andrews & A. Hardy [CDFA] (18). ESMERALDA Co.: Clayton Valley

Sand Dunes, 17 Sep 1974, F. Andrews & A. Hardy [CDFA] (3); Clayton Valley

Sand Dunes, 11.3 km S of Silver Peak, 1280 m, 2 May 1974, T. Eichlin & A.

Hardy [CDFA] (3); 9.7 km N of Oasis, 3.2 km W of Fish Lake Valley, 1376 m,

24 Feb 1982 to 24 Jun 1982, D. Giuliani, antifreeze pit trap, [CDFA] (3); Fish

Lake Valley Sand Dunes, 17 Aug 1976, D. Giuliani, cereal bowl pit trap [CDFA]

(1); 11.3 km N of Dyer, sand dunes, 1494 m, 24 Jun 1982 to 30 Sep 1982, D.

Giuliani, antifreeze pit trap [CDFA] (2); 16.1 km SE of Dyer, sand dunes, 1494

m, 24 June 1982 to 30 Sep 1982, D. Giuliani, antifreeze pit trap [CDFA] (4); 21

km N, 12.9 km E of Dyer, Fish Lake Valley, 1433 m, Sep 1986 to Sep 1987, D.

Giuliani, antifreeze pit trap [CDFA] (11); 4.8 km N of Goldfield, sand dunes,

1676 m, 28 Mar 1982 to 1 Oct 1982, [CDFA] (6); same locality, Mar 1983 to

Sep 1983, [CDFA] (4); same locality, Oct 1982 to Mar 1983, D. Giuliani, antifreeze

pit trap [CDFA] (1). HUMBOLDT CO.: 19.3 km NW of Winnemucca, 18 Jul

1977, D. Giuliani, cereal bowl trap [CDFA] (23). MINERAL Co.: Teels Marsh,

17 Feb 1979 to 16 Jul 1979, D. Giuliani, antifreeze pit trap, sand dune association

[CDFA] (6); Teels Marsh, sand dunes, 22 May 1976 [CDFA] (25); same locality,

16 Aug 1979 to 22 Nov 1979, [CDFA] (13); 22 Sep 1979 to 30 Jan 1980, [CDFA]

(7); 7 Jun 1980 to 31 Aug 1980, D. Giuliani, antifreeze pit trap, [CDFA] (11); 8

km W of Marietta, sand dunes, 1920 m, 6 Jun 1980, D. Giuliani [CDFA] (1);

12.9 km S of Mina, sand dunes, 22 May 1976, D. Giuliani [CDFA] (14); 14.5

km S of Mina, 30 Jun 1965, M. E. Irwin, sand dune association [CDFA] (2);

Huntoon Valley Sand Dunes, 16 Aug 1979 to 22 Sep 1979 [CDFA] (3); same

locality, 7 Jun 1980 to 31 Aug 1980 [CDFA] (4); same locality, 31 Aug 1980 to

25 May 1981 [CDFA] (1); D. Giuliani, antifreeze pit trap. NYE Co.: Current,

14.5 km S, 3.2 km W of Railroad Valley, 1494 m, Sep 1986 to Sep 1987, D.

Giuliani, antifreeze pit trap [CDFA] (3). PERSHING Co.: Woolsey, 27 Jun 1972,

T. R. Haig [CDFA] (1); Woolsey RR Stn., 6 June 1973, T. R. Haig, blacklight

[CDFA] (1). WASHOE Co.: Pyramid Lake Dunes, 7 Sep 1941, LaRivers [CASC]

(3). UTAH. EMERY Co.: 21A km N of Hanks ville, sand dunes nr. Glison Butte

Well, 26 Jul 1978, F. Andrews & A. Hardy, cereal bowl trap [CDFA] (22); 20.8

km N, 11.2 km E of Hanks ville, sand dunes, Sep 1983 to Mar 1984, D. Giuliani,

antifreeze pit trap [CDFA] (9). GRAND Co.: Arches Nat. Mon., Devils Garden

Campgr., 14 Sep 1983, R. L. Aalbu [RLAC] (19). KANE Co.: 8 km SE of Glen

Canyon City, sand dunes, 15-17 Jun 1988, R. L. Aalbu, pit traps, sand dune/

rodent burrows, [RLAC] (4); Lake Powell, Lone Rock Campgr., 15/17 Jun 1988,

R. L. Aalbu, pit traps, sand dune/rodent burrows, [RLAC] (44); Kanab, 16.1 km

N of Kanab cyn., 18-19 Jun 1988, R. L. Aalbu, pit trap sandstone overhang

Neotoma nest, [RLAC] (3). SAN JUAN Co.: Bluff Sand Dunes, 8 km W of Bluff,

24 Jul 1978, F. Andrews & A. Hardy, cereal bowl trap [CDFA] (13). UINTAH

Co.: 16.1 km SSW of Vernal, 13 Sep 1983, R. L. Aalbu [RLAC] (14). Paratypes

deposited in USNM, CASC, CISC, CASC, RLAC, OSUC.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

Description. —Female (holotype). Integument red-brown, elytra shiny; length approximately 2.9 mm.

HEAD with surface vestiture consisting of closely appressed, spatulate, scale-like setae with few longer

fine setae on apical margin of clypeus; antennal fossae with dorsal border not carinate, not laterally

elevated; eyes large, at least seven facets wide at minimum width, oval; antenna of moderate length,

stout, ratio of segment lengths 11:9:8:8:8:8:8:8:8:9:15. PRONOTUM with surface sculpture consisting

of rugose, deep punctures posteriorly forming moderately dense, small tubercles; surface vestiture of

two types, first of sparse, long setae in a row at anterior margin; second of short, stout, arched, recumbent

setae, dense, often subspatulate, forming dense ring at anterior margin; midlength transverse row of

four tufts unequal in size, medial tufts prominent, lateral tufts small to nearly obsolete. ELYTRA with

surface smooth, shiny, strial punctures obsolete; vestiture of two types: first of long, fine, erect setae

sparsely positioned at regular intervals on first, third, fifth, seventh intervals; second of moderately

long, stout, arched, recumbent setae, positioned in rows on elytral intervals, more abundant and

shorter, around suture, on lateral margins. VENTRAL SURFACE: Sterna: ratio of segment lengths

15:17:15:5:19; sternal surface vestiture of short, dense golden closely appressed, spatulate setae in¬

termixed with less dense, slightly longer, fine setae; fifth visible abdominal stemite with medial apical

area with closely packed patch of short appressed, scale-like setae. LEGS stout, with femora short,

clavate; tibiae short; metatibia curved proximal-posteriorly; femoral vestiture consisting of dense,

golden, short, appressed, scale-like setae only slightly varying in length; tibiae with similar vestiture

except protibiae with dense, longer, slender, golden setae on lower margins; mesotibiae with dense,

longer, slender, golden setae on lower margins, on apical one-half of outer margins; metatibiae with

few sparse, longer, golden setae on lower margins. Ratios of segment lengths: prothoracic legs, 41:40;

mesothoracic legs, 46:45; metathoraciclegs, 60:59; protarsi, 8:4:4:4:7; mesotarsi, 10:4:4:4:9; metatarsi,

15:7:6:6:11.

Male (allotype). —Similar to holotype but smaller, approximate length 2.6 mm. Fifth visible ab¬

dominal stemite with medial apical area with setae similar to rest of sternal area.

Diagnosis. — The following combination of characters will serve to separate N.

giulianii : Head with eyes large and antennal fossa with dorsal border not carinate

or laterally raised. Pronotum with medial transverse pronotal tufts, larger than

lateral tufts; anterior margin of pronotum with long erect setae. Elytra with erect

setae on intervals three and five long and slender, short on one and absent on

two and four. Legs short, with metafemur clavate, metatibia stout, curved prox¬

imal-posteriorly. Sexually dimorphic: female with fifth visible abdominal stemite

with medial apical area bearing closely packed patch of minute, scale-like setae.

Label Biological Notations. — Dry overnight “punch cup” pitfall trap, ethylene

glycol pitfall trap near Neotoma nest, pit traps sand dune/rodent burrows, pit

traps sandstone overhang Neotoma nest.

Distribution. —(Fig. 25) The peculiar east-west distribution of this species prob¬

ably reflects lack of adequate collections from this middle area instead of a real

distributional gap. There is, however, a curious absence of this species from the

Eureka Valley sand dunes region, an area that has undergone intensive trapping;

whereas, the species is present in Deep Springs Valley sand dunes, only eight

miles away.

Biological Notes. — This species is often associated with rodent burrows near or

on sand dunes, although it is also found off of the dunes.

Material examined.— See types.

Niptus neotomae Aalbu & Andrews, NEW SPECIES

(Figs. 2 and 25)

Types. - HOLOTYPE (female) and ALLOTYPE (male): ARIZONA. GRAHAM

Co.: Pinaleno Mountains, Heliograph Peak, 3055 m elevation, 9 Sep 1987, G. E.

Haas col. Holotype and allotype deposited in the collection of the California

1992

AALBU & ANDREWS: REVISION OF NIPTUS

Academy of Sciences. PARATYPES: 5, same data. Deposited in USNM, CDFA

and RLAC.

Description. — Female (holotype). Integument red-brown, elytra shiny; length approximately 2.6 mm.

HEAD with surface vestiture consisting of closely appressed, short, spatulate, scale-like setae; antennal

fossae with dorsal border not carinate, not laterally elevated; eyes small, five facets at minimum width,

oval in shape; antenna of moderate length, ratio of segment lengths 10:8:7:6:6:6:6:6:6:7:13. PRO-

NOTUM with surface sculpture consisting of rugose punctures; surface vestiture of short, stout,

recumbent setae; setae dense at anterior margin, at transverse row of four weakly developed tufts;

tufts equal in size, positioned near midlength. ELYTRA with surface shiny, sculpture small deep

regular punctures; vestiture of two types, nearly equal in length; first of short, moderately slender,

erect, strongly spatulate setae positioned in rows at regular distances along first to seventh intervals;

second arched, recumbent, moderately slender setae positioned in approximate rows on both elytral

striae and elytral intervals; setae short, dense at elytral margins. VENTRAL SURFACE: Sterna: ratio

of segment lengths 12:15:14:4:17; sternal surface vestiture consisting of short, closely appressed,

spatulate, scale-like setae; fifth visible abdominal stemite with medial apical area with dense patch of

apically directed, semi-erect setae. LEGS stout, femora short, clavate, metafemora slightly bent near

apex; tibia stout; femoral vestiture consisting of dense, golden, short, appressed, scale-like setae only

slightly varying in length; tibiae with similar vestiture except protibiae with dense, slightly longer,

slender, golden setae on lower margins, mesotibiae with dense, slightly longer, slender, golden setae

on lower margins, on apical one-half of outer margins; metatibiae with few sparse, slightly longer,

golden setae on lower margins. Ratio of segment lengths: prothoracic legs, 32:31; mesothoracic legs,

35:33; metathoracic legs, 39:40; protarsi, 5:4:3:4:7; mesotarsi, 5:4:4:4:6; metatarsi, 8:5:4:4:7.

Male (allotype).—Similar to holotype but slightly smaller, approximate length 2.3 mm; eyes slightly

smaller than female, four facets in width; fifth visible abdominal stemite with setal pattern unmodified.

Diagnosis. — The following combination of characters will serve to separate N.

neotomae : Head with eyes small and antennal fossa with dorsal border not carinate

or laterally raised. Pronotum with medial, lateral transverse pronotal tufts equal

in size, only slightly developed; anterior margin of pronotum without long erect

setae. Elytra with erect setae on intervals one to five short, spatulate. Legs short,

stout, with metafemur clavate. Sexually dimorphic: female with fifth visible ab¬

dominal stemite with medial apical area bearing closely packed apically directed,

semi-erect setae forming a rounded patch.

Niptus neotomae is most closely related to N. arcanus, sharing short spatulate

elytral setae. These species also lack long erect setae on pronotal margins and

have sexual dimorphism, characters also shared by N. abscondidus Spilman. Nip¬

tus neotomae differs from N. arcanus in having shorter setae both on the pronotum

and elytra and in the configuration of the legs: short and stout, metafemora clavate

in N. neotomae ; long and slender, metafemora capitate in N. arcanus.

Distribution.—{ Fig. 25) This species is only known from the type locality.

Label Biological Notations. —Nest of Neotoma mexicana in U.S.F.S. shed.

Biological Notes. — Haas (T. J. Spilman, personal communication) mentions

finding the beetles while searching for fleas in a rather dry and dusty nest composed

of shredded cloth, newspapers, wrappers, cardboard, and packing material sur¬

rounded by cones, bark, sticks and various dried green plant material on the floor

of the shed between some storage boxes.

Material Examined.—See, types.

Niptus abstrusus Spilman

(Figs. 6 and 26)

Niptus abstrusus Spilman, 1968: 195.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Diagnosis. — The following combination of characters will serve to separate N.

abstrusus : Head with eyes small and antennal fossa with dorsal border carinate

and laterally raised. Pronotum with medial and lateral transverse pronotal tufts

equal in length; anterior margin of pronotum with long erect setae. Elytra with

erect setae on intervals three and five only slightly longer than those on intervals

one, two and four; legs with metafemur clavate, metatibia stout, curved. Not

sexually dimorphic.

Distribution.—{ Fig. 26) Southwestern Texas and north-central Mexico. Known

from caves in Texas (Fern Cave [Val Verde Co.], Bat Cave [Brewster Co.) and

Mexico (Pedrigosa Circle Cave, Pedrigosa Pipe Cave, and Cueva de San Vincente

[Coahuila]).

Label Biological Notations. — On pineapple, on dry beans, with Ariocarpus lloydi.

Biological Notes. — Ashworth (1973) reports finding fragments of individuals of

this species in a 12,000 year old fossil Neotoma nest in western Texas. Individuals

have been reported on raccoon droppings (Reddell 1966) and on bat guano (Red-

dell 1970).

Material Examined. -Twenty-one from the following seven localities: TEXAS. VAL VERDE Co.:

Fern Cave, 27.4 km N of Comstock (7); bat room (3). MEXICO. (2) (state unknown) DURANGO:

Tepehuanes (8). COAHUILA: (1).

Niptus absconditus Spilman

(Figs. 7, 23 and 26)

Niptus absconditus Spilman, 1968: 197.

Diagnosis. — The following combination of characters will serve to separate N.

absconditus: Head with eyes small and antennal fossa with dorsal border not

carinate or laterally raised. Pronotum with medial and lateral transverse pronotal

tufts equal in size; anterior margin of pronotum without long erect setae. Elytra

with erect setae on intervals one to five short; legs long, with metafemur capitate,

metatibia stout, almost straight. Sexually dimorphic: female with fifth visible

abdominal stemite with medial apical area bearing closely packed patch of dense,

short scale-like setae. Niptus abscondidus is most closely related to N. arcanus.

See discussion under N. arcanus.

Distribution. — This species is only known from the type locality.

Label Biological Notations. —None.

Material Examined.— Four specimens (PARATYPES) from: MEXICO. HIDALGO: Grutas de

Xoxafi, VIII-19-65, J. Reddell, J. Fish & W. Bell cols.

Niptus abditus Brown

(Figs. 4 and 25)

Niptus abditus Brown, 1959: 631.

Diagnosis. — The following combination of characters will serve to separate N.

abditus : Head with eyes minute and antennal fossa with dorsal border not carinate

or laterally raised. Pronotum with lateral transverse pronotal tufts more developed

than medial pronotal setal tufts; anterior margin of pronotum without long erect

setae. Elytra with erect setae on intervals three and five longer than those on

intervals one, two and four; legs with metafemur capitate, metatibia slender,

1992

AALBU & ANDREWS: REVISION OF NIPTUS

straight. Sexually dimorphic: female with fifth visible abdominal stemite with

medial apical area bearing closely packed patch of postero-directed, semi-erect

setae forming a rounded tuberclelike structure.

Distribution. — This species is only known from the three localities mentioned.

Label Biological Notations. — Ex. nest of Neotoma sp., ethylene glycol pit trap.

Material Examined.— UTAH. SAN JUAN Co.: 12.9 km E of Bluff, 1402 m, September 1984 to

March 1985, ethylene glycol pit trap, D. Giuliani col. (2). TOOELE Co.: Great Salt Lake, Stansbury

Island, floor of Spider Cave, 9.1 m from the entrance, 29 Nov 1952, J. R. Keller col., PARATYPE

#6916, [CNCI]. UTAH Co.: Rock Canyon near Provo, 15 Jun 1964, V. J. Tipton col., (8).

Niptus sleeperi Aalbu & Andrews, NEW SPECIES

(Figs. 8 and 16)

Type. — Holotype (male). MEXICO. BAJA CALIFORNIA SUR: 21A air km

ENE of Todos Santos, Sierra Laguna, La Laguna, 4-7 Jun 1973, E. L. Sleeper col.

Type deposited in California Academy of Sciences Collection.

Description .—Male (holotype). Integument dark red-brown, vestiture golden to yellow; length ap¬

proximately 2.4 mm. HEAD with surface vestiture consisting of closely appressed, spatulate, scale¬

like setae with few longer fine setae on apical margin of clypeus; antennal fossae with dorsal border

carinate, laterally elevated; eyes small, four facets at minimum width, narrowly oval in shape; antenna

short, stout; ratio of segment lengths 10:9:7:6:6:6:6:6:6:7:13. PRONOTUM with surface sculpture

consisting of rugose, deep punctures posteriorly forming moderately dense, small tubercles; surface

vestiture of two types, first of few, sparse, moderately long, line setae (with apical ends occasionally

finely spatulate) positioned near anterior margin; second of short, stout, dense, arched, recumbent

setae; setae denser, shorter, stouter at anterior margin; denser at midlength transverse row of four

tufts; tufts equal in size. ELYTRA with surface sculpture of deeply impressed, large, contiguous strial

punctures, equal in size, impression throughout; surface vestiture of two types: first of moderately

long, fine, sparse, erect setae (equal in length to erect setae on pronotal margin) positioned at regular

intervals along first to seventh elytral intervals; second of shorter, stout, arched, recumbent setae

positioned throughout elytral surface, more abundant on intervals. VENTRAL SURFACE: Sterna:

ratio of segment lengths 11:12:6:3:15; sternal surface vestiture of short, golden, closely appressed, fine

setae; fifth visible abdominal stemite with medial apical area unmodified. LEGS short, stout, with

femora clavate; tibiae short; metatibia curved proximoposteriorly; femoral vestiture consisting of

dense, golden, short, appressed, scale-like setae slightly varying in length; tibiae with similar vestiture

except: protibiae with dense, longer, slender, golden setae on lower margins, mesotibiae with dense,

longer, slender golden setae on lower margins, on apical one-half of outer margins; metatibiae with

few sparse, longer, golden setae on lower margins. Ratio of segment lengths: prothoracic legs, 28:31;

mesothoracic legs, 40:33; metathoracic legs, 37:42; protarsi, 5:3:3:3:6; mesotarsi, 7:3:3:3:7; metatarsi,

9:4:4:4:7.

Female. — Unknown.

Diagnosis.— The following combination of characters will separate N. sleeperi :

Head with eyes small and antennal fossa with dorsal border carinate, laterally

raised. Pronotum with medial and lateral transverse pronotal tufts equally de¬

veloped; anterior margin of pronotum with long erect setae. Elytra with deeply

impressed, large, contiguous strial punctures, equal in size and impression

throughout; erect setae on intervals one to five short. Legs with metafemur clavate,

stout, metatibia stout, curved. Sexual dimorphism unknown.

Distribution.— (Fig. 26) This species is only known from the type locality.

Label Biological Notations. — Berlesed from oak duff.

Material Examined. — Holotype; only it is known.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Niptus ventriculus LeConte

(Figs. 5, 13, 16, 18, 22, and 26)

Niptus ventriculus LeConte, 1859: 13.

Diagnosis. — The following combination of characters will separate N. ventric¬

ulus: Head with eyes large and antennal fossa with dorsal border carinate, laterally

raised. Pronotum with medial and lateral transverse pronotal tufts equally de¬

veloped; anterior margin of pronotum with long erect setae. Elytra with erect setae

on intervals three and five long; short on intervals one, two and four. Legs with

metafemur clavate, metatibia broad, curved. Not sexually dimorphic.

Elytral strial setal length and punctures vary greatly in populations from sub¬

equal setal length and completely smooth punctures, except for the ninth interval

in specimens from Coahuila, to slight variation in strial setal length and few rows

of punctures in specimens from Glamis, California, to strongly punctate with long

setae in specimens from near Bakersfield, California.

Label Biological Notations. — UV light; rodent nest; ex mouse nest Peromyscus

eremicus; kangaroo rats: in burrows of, excavating and sifting burrow of; sifting

beach dunes under ambrosia; base of Palo Verde; at night: walking dunes, pitfall;

pitfalls: cereal bowl trap, under Larrea and Petelonyx thurberi, ethylene glycol

trap, antifreeze trap on sand dune with creosote and sand verbenas, rye bread

trap, dry overnight “punch cup” trap, interdune traps. Spilman (1968) mentions

records from nests of Neotoma, and the kangaroo rats Dipdomys deserti Stephens

and Dipdomys spectabilis Merriam, pitfall traps in sand dunes, under seaweed

and rocks at high tide line, sifting sand on dunes, pit traps sand dune/rodent

burrows, antifreeze pit trap on sand dune.

Distribution.—{ Fig. 26) Widespread throughout southwest U.S. and Mexico.

Material Examined. —(945 from the following 103 localities). ARIZONA (76 specimens/12 local¬

ities). no locality (3). COCHISE Co.: 6.4 km E of Portal (1); A.M.S.W.R.S. (7). COCONINO Co.:

Moenkopi, Moenkopi sand dunes (12); 6.3 km SE of Moenkopi, sand dunes/dry canyon (11); 3.2 km

S of Moenkopi (3); 16.1 km S and 8 km W of Page (29). GREENLEE Co.: Guthrie (3). LA PAZ Co.:

4.8 km SE of Parker (1). MOHAVE Co.: Littlefield, 580 m (7). NAVAJO Co.: (1). PIMA Co.: Santa

Rita Mts. (5). YUMA Co.: Yuma (4); CALIFORNIA (302 specimens/50 localities). FRESNO Co.:

Monocline Ridge Sand Dunes (1); 12.9 km NNW of Coalinga, Los Gatos Cyn. (2); 29 km SW of

Mendota, Cievo Hills (5). IMPERIAL Co.: Holtville (1); 12.9 km ESE of Holtville, East Mesa Geo¬

thermal Site (11); Seely (17); Glamis (31); 1.6 km S of Glamis (6); 4.8 km NW of Glamis (1); 22.5

km NW of Glamis (5); 1.6 km N of Glamis (25); 3.2 km N of Glamis (1); 3.2 km NW of Glamis

(13); 5.6 km WNW of Glamis (2); 5.6 km NW of Glamis (9); 11.3 km SE of Glamis, Algodones

Dunes, 32 o 55'20" N, 114°59T4' / W, Site 4 (12); 6.5 km W of Ogilby, 32°48'48" N, 104°53'51" W (1);

6.4kmSSWofOgilby, 32°45'33"N, 104°51'32"W, Site 7 (3); Algodones Dunes, 4 km NE of Coachella

Bridge No. 1, 32°51'41" N, 115°4'6" W, Site 24, (1); Algodones Dunes, 20 km ESE of Holtville,

32°44'34" N, 115°1153" W, Site 30, (1). INYO Co.: Chicago Valley Sand Dunes (2). KERN Co.: 12.9

km N and 4.8 km W of Ridgecrest (1); 1.6 km E of Bakersfield Hart Peak (1). KINGS Co.: no locality

(2); 7.7 km W of Kettleman City, 7.7 km W (8) of, and 3.2 km S of Leemoore. RIVERSIDE Co.:

Hopkins Well (2); Palm Springs (1); Rice Dunes (13); Palen Dunes (16); 11.3 km SE of Freda (2); 4.8

km W of Blythe (7); 1.6 km W of Blythe (10); Indio (1); 8 km E of Indio (2); La Quinta (1); Mule

Mts. (3); 3.2 km NW of Gilman Hot Springs, Lamb Canyon (9). SAN BERNARDINO Co.: Cadiz

Dunes (33); Kelso Dunes (4); 28.2 km SE of Baker; Cronese Valley (2); 14.5 air km S of sand dunes

S ofZzyzx (3); 14.5 km N and 16.1 km E of Ridgecrest (3); 16.1 km N and 16.1 km E of Ridgecrest

(1); 9.7 km N and 3.2 km W of Ridgecrest (1); 30.6 km N of Ridgecrest, Baby Mt. (7); Amargosa

River at st. hwy. 127 (1). SAN DIEGO Co.: Borrego (1). SAN LUIS OBISPO Co.: 12.1 km W of

Simmler (18); 24.9 km NW of Reyes Station (1). SANTA CRUZ Co.: (1); Watsonville (1). NEW

1992

AALBU & ANDREWS: REVISION OF NIPTUS

MEXICO. (32 specimens/5 localities). Hot Springs (4). LUNA Co.: Deming (2); E of Deming at base

of Red Mt. on Humocky Rd. (14). SAN JUAN Co.: Ship Rock (11). SOCORRO Co.: Sevilletta Sand

Dunes (1). TEXAS. (4 specimens/2 localities). EL PASO Co.: El Paso (3). PRESIDIO Co.: Marfa (1).

UTAH (55 specimens/5 localities). JUAB Co.: Fish Springs Range, 40.2 km SE of Callao, Sand Pass

(4). KANE Co.: 8 km SE of Glen Canyon City, sand dunes (3); Lake Powell, Lone Rock Campgr. (27).

SAN JUAN Co.: 3.2 km S and 32.2 km W of Bluff (10). WASHINGTON Co.: 17.9 km N ofSt. George,

red sand dunes (1). MEXICO. BAJA CALIFORNIA (327 specimens/13 localities): Miller’s Landing

(84); 16.1 km S of Punta Prieta (2); 12. 4 km NW of Catavina (2); El Crusero (22); 41.4 km SE of

Laguna Chapala (15); 19.3 km NW of San Bartolo (1); 9.7 km N of Guerrero Negro (154); 5 km N

of Guerrero Negro (8); 11.3 km N of Guerrero Negro (6); 25.7 km E of Rosarito, Rancho San Ignacito

(22); 10 km NE of Rosarito (9); 5.0 km SW of Colonet (1); Bahia San Quintin, Santa Maria Beach

(1). BAJA CALIFORNIA SUR (96 specimens/8 localities): 22.5 km E of Guerrero Negro (1); 55.4 km

SE of Guerrero Negro (4); 13.7 km ESE and 8.6 km S of Guerrero Negro (2); 11.3 km SE of Guerrero

Negro (27); 20.9 km SW of Guillermo Prieto (43); 19.3 km S of Guillermo Prieto (10); 20.9 km S of

Rancho Tablon (8); Tortugus (1). COAHUILA (42 specimens/1 locality): 12.9 km N of Viesca, sand

dunes at Bilbao (42). DURANGO (26 specimens/1 locality): 43.5 km S of Ceballos (26). SONORA

(12 specimens/7 localities): Puerto Penasco, 0.5 km from coast (1); Desemboque (1); El Golfo (4); 80.

5 km SW of Sonoyta (1); 16.1 km N of C. Sotelo nr. Bahia Adair (1); San Carlos Bay (1); 9.7 km W

of San Carlos Bay, Los Algodones (3).

Phylogenetic Considerations

Pseudorostus and Eurostus have historically either been separated from Niptus

(Brown 1940: 119, 1944: 19, 1959: 627; Hinton 1941: 343; Spilman 1968: 193)

or included as synonyms of Niptus (Papp 1959: 258, 1962: 385; Spilman [North

American Beetle Fauna Project] 1975: R62-1). Of these, Eurostus has generally

been accepted as being congeneric with Pseudeurostus. Pseudeurostus has been

separated from Niptus based on the carinate frons between the antennal fossae in

Pseudeurostus (Fig. 12), which is not narrowly flat as in Niptus (Fig. 11). Clearly,

this character is unique and synapomorphic in species of Pseudeurostus. However,

P. hilleri and P. kelleri and all species of Niptus except N. hololeucus (Fig. 15)

share a strongly reduced fourth visible abdominal stemite (Figs. 17, 21), another

clearly synapomorphic character. Thus, if Pseudeurostus is to be generically sep¬

arated from Niptus, then N. hololeucus, the type species of Niptus, needs also to

be separated from both groups, making it necessary for a new generic name for

the eight “wild” species of Niptus. It is clearly preferable to lump all these under

the genus Niptus as indicated by Spilman (1975: R62-1).

Key to North American Species of Niptus

1. Body large (usually above 3.8 mm in length), golden throughout, color

result of scale-like setae that completely conceal integument of entire

insect; fourth visible abdominal stemite only slightly shorter than

third (Fig. 15); pronotum lacking distinct tufts of setae (Fig. 10) ..

. hololeucus

1'. Body smaller (usually under 3.4 mm in length), red-brown above, elytra

not completely covered with scale-like setae; fourth visible abdom¬

inal stemite less than one-half length of third. 2

2(T). Frons carinate between antennal fossae (Fig. 12); pronotum lacking

distinct transverse row of four tufts of setae ( Pseudeurostus group) 3

2'. Frons narrow but flat between antennal fossae (Figs. 11, 13); pronotum

with distinct transverse row of four tufts of setae (Fig. 18) . 4

Vol. 68(2)

94 THE PAN-PACIFIC ENTOMOLOGIST

3(2). Elytral intervals with closely placed, recumbent setae as well as single

row of semi-erect setae. kelleri

3'. Elytral intervals with single row of semi-erect setae, lacking closely

placed, recumbent setae (Fig. 12). hilleri

4(2'). Antennal fossa with dorsal border distinctly carinate and laterally

strongly elevated (Figs. 13, 16). 5

4'. Antennal fossa with dorsal border not distinctly carinate and laterally

not strongly elevated (Fig. 11). 7

5(4). Elytra with strial punctures deeply impressed, large, contiguous, equal

in size and impression throughout; erect setae on intervals one to

five short; legs with metafemur clavate, very stout; eyes small (Fig.

8) . sleeperi

5'. Elytra with strial punctures unevenly impressed, always fine to minute

at apex and lateral margins; erect setae on intervals three and five

long to moderately long; legs with metafemur clavate; eyes variable

. 6

6(5'). Elytra with erect setae on intervals three and five long; short on intervals

one, two and four; strial punctures often large and deeply impressed

on disc; eyes large (Figs. 5, 22). ventriculus

6'. Elytra with erect setae on intervals three and five only slightly longer

than those on intervals one, two and four, strial fine to minute; eyes

smaller (Fig. 6). abstrusus

7(4'). Elytra with length of erect setae on intervals three and five greater than

width of one interval; pronotal tufts not equal in size; eyes variable

. 8

7'. Elytra with length of erect setae on intervals three and five less than

width of one interval; pronotal tufts equal in size; eyes very small 9

8(7). Pronotum with medial transverse pronotal setal tufts more developed

than lateral pronotal setal tufts; anterior margin of pronotum with

long erect setae; elytra with erect setae absent on intervals on two

and four; legs with metafemur clavate, metatibia stout, strongly

curved; eyes large; stemites sexually dimorphic (Fig. 3). giulianii

8'. Pronotum with lateral transverse tufts more developed than medial

pronotal setal tufts; anterior margin of pronotum lacking long erect

setae; elytra with erect setae short but present on two and four; legs

with metafemur capitate, metatibia slender, straight; eyes small; ster-

nites not sexually dimorphic (Fig. 4). abditus

9(7'). Erect setae on elytra spatulate at tip (Figs. 1, 2); width of metatibiae

variable . 10

9'. Erect setae on elytra unmodified, pointed at tip; width of metatibiae

at apex equal to widths of eighth and ninth intervals combined (Fig.

7) . absconditus

10(9). Erect spatulate setae on elytra very short; legs short, stout, metafemora

clavate (as in Fig. 20), width of metatibia at apex equal to widths of

eighth and ninth intervals combined (Fig. 2) . neotomae

10'. Erect spatulate setae on elytra short (Fig. 1); legs long, slender, meta¬

femora capitate (Fig. 19), width of metatibia at apex subequal to

widths of eighth and ninth intervals combined . arcanus

1992

AALBU & ANDREWS: REVISION OF NIPTUS

Biology

North American species of Niptus not associated with stored products seem to

be distributed in two seemingly different habitats: caves and sand dune areas.

These two habitats do share one important aspect of the microhabitat in which

Niptus species are found. This is a fine to very fine substrate (in the form of fine

sand or cave dust) with a varying amount of organic debris due to rodent activity.

All species seem to be associated with various desert rodents especially species

of packrats ( Neotoma ), but also mice ( Peromyscus ), and kangaroo rats ( Dipdomys ).

Niptus arcanus, N. abstrusus, N. absconditus, and N. kelleri are found in caves.

These reveal to varying degrees, morphological characteristics typically associated

with cave coleoptera (see Aalbu 1990). Of these N. arcanus, N. Kelleri and N.

absconditus are restricted to single cave habitats. Niptus arcanus is considered to

be a true troglobite (Aalbu 1990) a relative rarity in Northwestern American

beetles. It is possible that upon further study, other species will also be classified

as troglobites rather than troglophiles. We can only stress the importance of

conserving these unique cave habitats, especially in caves where considerable

environmental impact is present due to high visitor traffic (such as in Mitchell

Caverns). This would entail assuring species survival by providing for long term

microhabitat protection in terms of the least amount of habitat disturbance pos¬

sible.

Acknowledgment

The following individuals and institutions are gratefully acknowledged for loan

of their material: David Kavanaugh, California Academy of Sciences, San Fran¬

cisco, California; John T. Doyen, University of California, Berkeley, California;

Ives Bousket, Canadian National Collection, Ottawa, Ontario, Canada; E. L.

Sleeper, California State University, Long Beach, California; A1 Newton, Field

Museum of Natural History, Chicago, Illinois; Kirby W. Brown, Stockton, Cal¬

ifornia; Scott Shaw, Harvard University Museum of Comparative Zoology, Cam¬

bridge, Massachusetts; Charles A. Triplehom, The Ohio State University, Co¬

lumbus, Ohio; Dave Faulkner, San Diego Museum of Natural History, San Diego,

California; Ted J. Spilman, United States National Museum, Washington D.C.;

Carl A. Olson, University of Arizona, Tucson, Arizona. The following individuals

deserve special mention: Ives Bousket of The Canadian National Collection for

loan of paratype of N. abditus; Phillips L. Claud, Jim Hart, Bill Wisehart and

John Kelso-Shelton of the California Department of Parks and Recreation, for

granting collection permits and aiding in collections of N. arcanus.

Literature Cited

Aalbu, R. L. 1990. An analysis of the Coleoptera of Mitchell Caverns, San Bernardino County,

California. N. Speol. Soc. Bull., 51: 1-10.

Ashworth, A. C. 1973. Fossil beetles from a fossil wood rat midden in western Texas. Coleop. Bull.,

27: 139-140.

Brown, W. J. 1940. A key to the species of Ptinidae occurring in dwellings and warehouses in Canada

(Coleoptera). Can. Entomol., 72: 115-122.

Brown, W. J. 1944. Some new and poorly known species of Coleoptera, II. Can. Entomol., 76: 4-

10.

Brown, W. J. 1959. Niptus Boield. and allied genera in North America (Coleoptera: Ptinidae). Can.

Entomol., 91: 627-633.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

Hinton, H. E. 1941. The Ptinidae of economic importance. Bull. Entomol. Res., 31: 331-381.

Howe, R. W. 1959. Studies on beetles of the family Ptinidae, XVII. Conclusion and additional

remarks. Bull. Entomol. Res., 50: 287-326.

Papp, C. S. 1959. Discussions on the taxonomic status of some Ptinidae of economic importance.

(Notes on North American Coleoptera, No. 5) Entomol. Berichten, 19: 256-259.

Papp, C. S. 1962. An illustrated and descriptive catalogue of the Ptinidae of North America. (Notes

on North American Coleoptera, No. 7) Dtsch. Entomol. Ztschr. (N.F.), 9: 367-423.

Reddell, J. R. 1966. A checklist of the cave fauna of Texas. II. Insecta. Texas J. Sci., 18: 25-56.

Reddell, J. R. 1970. A checklist of the cave fauna of Texas. V. Additional records of Insecta. Texas

J. Sci., 22: 47-65.

Spilman, T. J. 1968. Two new species of Niptus from North American caves (Coleoptera: Ptinidae).

Southwestern Natur., 13: 193-200.

Spilman, T. J. 1975. Ptinidae. Page R62.1. In Blackwelder, R. E. & R. H. Arnett, (eds.) Checklist

of the beetles of Canada, United States, Mexico, Central America and the West Indies, Volume

1, part 5, The darkling beetles, ladybird beetles and related groups (red version). Biol. Res. Inst.

Amer. Inc. Rensselaerville, New York.

Received 1 May 1990; accepted 15 July 1991.

PAN-PACIFIC ENTOMOLOGIST

68(2): 97-99, (1992)

THE SOLITARY BEE

MELISSODES THELYPODII THELYPODII COCKERELL

(HYMENOPTERA: ANTHOPHORIDAE) COLLECTS

POLLEN FROM WIND-POLLINATED

AMARANTHUS PALMERI WATSON

James H. Cane, 1 Stephen L. Buchmann, 2 and Wallace E. LaBerge 3

1 Department of Entomology and Alabama Agricultural Research Station,

Auburn University, Alabama 36849-5413

2 USDA, ARS, Carl Hayden Bee Research Center, 2000 East Allen Rd.,

Tucson, Arizona 85719

3 Illinois Natural History Survey, Champaign, Illinois 61820

Abstract.— The native solitary bee Melissodes thelypodii thelypodii Cockerell was observed to

harvest pollen from panicles of the anemophilous plant Amaranthus palmeri Watson in south¬

eastern Arizona. Pure Amaranthus pollen loads were removed by females foraging at this plant,

suggesting floral fidelity and this bee’s potential value for commercial pollination of the related

grain amaranths.

Key Words.— Insecta, pollination, anemophily, Melissodes, Amaranthus, pollen-foraging, bees

Foragers of social bees will sometimes collect pollen from flowering plants that

rely upon wind to transport pollen to receptive pistils (Faegri & van der Pijl 1978).

Honey bees {Apis mellifera L.) and sometimes stingless bees {Trigona s.l.) collect

pollen from diverse anemophilous (wind-pollinated) plants (Sharma 1970; O’Neal

& Waller 1984; C. D. Michener, unpublished data). Less commonly, bumble bees

{Bombus sp.) may collect pollen from anemophilous plants, such as bahia grass,

Paspalum notatum var. saurae Parodi (JHC, unpublished data).

In contrast, nonsocial or solitary bees have rarely been reported to gather pollen

from anemophilous plants. The pollen of oaks ( Quercus ), which are considered

anemophilous, may be gathered by solitary bees when their preferred pollen hosts

are not available {Andrena erythronii Robertson [Michener & Rettenmeyer 1956];

Osmia rufa [Raw 1974]; Habropoda laboriosa (Fabr.) [Cane & Payne 1988]).

Several British Andrena reportedly collect pollen periodically from several ane¬

mophilous trees, such as oak and chestnut (Chambers 1945). Nomiine bees of

the Old World genus Rhopalomelissa collect and may depend on grass pollen for

larval provisions (C. D. Michener, unpublished data). The sweat bee Dialictus

illinoiensis (Robertson) avidly harvests pollen from dallis grass, Paspalum dila-

tatum Poiret, augmenting the seed set of this grass (Adams et al. 1981).

Careless-weed {Amaranthus palmeri Watson) is a weedy, dioecious amaranth

occurring through much of the central and western United States and Mexico

(Munz 1959). The species exhibits several characteristics that typify anemophilous

plants. It produces copious pollen that bears little of the oily pollenkitt typical of

pollen usually collected by bees. Its small (24-26 pm diam) periporate “cheno-

am” (Chenopodiaceae-Amaranthaceae) type pollen grains are commonly impli¬

cated in human hayfever allergies (Wodehouse 1971). Bees have not been reported

to visit this anemophilous plant.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

We observed female Melissodes thelypodii thelypodii Cockerell working the

spike-like panicles of male plants of A. palmeri for pollen during mornings of

August, 1990, along the edge of a cotton field in the San Simon Valley of south¬

eastern Arizona (Cochise Co.). In this vicinity, we previously noted this bee species

sonicating flowers of Solanum elaeagnifolium Cavanilles and S. rostratum Dunal

(Solanaceae) for pollen. Bees of other species that worked nearby Solanum flowers

were never seen at Amaranthus (Protoxaea gloriosa (Fox), Protandrena mexica-

norum (Cockerell), Bombus sonorus Say, and Caupolicana yarrowi (Cresson)).

The first female M. thelypodii thelypodii to visit a given inflorescence frequently

released a visible cloud of pollen upon alighting. Females walked along the spikes

gathering pollen, proceeding distally from mid-base along those spikes that were

upright. They would then fly to a neighboring spike to continue collecting pollen.

Some females accumulated a full load of Amaranthus pollen from five to seven

spikes in as few as six minutes. A microscopic survey of the taxonomic constitution

of their pollen loads revealed good floral fidelity. Pure loads of Amaranthus pollen

were carried by five of six females collected at Amaranthus. The remaining female

bore 11% cotton pollen and 89% Amaranthus pollen. We have found only one

other reference to species of Melissodes gathering pollen from anemophilous plants.

Adams et al. (1981) reported M. bimaculata (Lepeletier) to occasionally collect

pollen from dallis grass.

The role, if any, of bees in the pollination of dioecious anemophilous plants

has been debated in the pollination literature. Usually, solitary or social bees that

harvest anemophilous pollen are considered mere pollen thieves, removing pollen

from male plants but never subsequently visiting the nectarless female plants.

Their contribution to pollination can not be discounted, however. Foragers may

mistakenly visit female flowers upon occasion. Even if they only visit male flowers,

wing and leg movements during pollen collection may dislodge prodigious quan¬

tities of pollen which, once airborne, can travel to female flowers.

The relative contributions of wind and insects, specifically bees, to the polli¬

nation of wild and cultivated amaranths also remains equivocal. Several weedy

species have been implicated in human respiratory allergies, including A. palmeri

(Wodehouse 1971). Unlike many anemophilous pollens, such as conifer pollen,

the pollen of A. palmeri seems to be moderately nutritious for bees, containing

3.5% nitrogen, or 18.38% crude protein by micro-Kjeldahl analysis (SLB, un¬

published data). Bee activity varies greatly at amaranths. Kaufman (1979) reported

an absence of bees at cultivated grain amaranth. In contrast, Singh (1961) some¬

times observed abundant bees at grain amaranth in India, and O’Neal & Waller

(1984) found Amaranthus pollen to constitute 6% of the average annual pollen

intake of honey bee colonies in the Sonoran desert, near Tucson, Arizona. Using

pollen traps on honey bee colonies, one of us (SLB) found that cheno-am pollen

constituted from 2-8% of the annual colony pollen harvest near Tucson during

the years 1981-1989. The solitary bee Hylaeus bisinuatus Forster has been re¬

corded visiting members of the Amaranthaceae (Krombein et al. 1979). We con¬

clude that amaranth pollen is not a mere scopal contaminant in these cases. Bees

will actively collect amaranth pollen, perhaps reflecting the ease with which quan¬

tities of this pollen can be harvested relative to other competing floral species.

Pox Amaranthus retroflexus L., Murphy (1978) demonstrated that insects alone

could provide cross-pollination. Further, Hauptli & Jain (1985) found wide dis-

1992

CANE ET AL.: POLLEN COLLECTION BY MELISSODES

parities in outcrossing rates for their field experiments with cultivated A maranthus

cruentus. They attributed this result to variable densities of pollen-foraging honey

bees in their plots.

Our observations demonstrate that females of the solitary bee M. t. thelypodii

avidly visit staminate flowers of amaranth for pollen, exhibiting good species

fidelity on a given foraging trip. For at least hermaphroditic species of commercial

amaranth, the visitations of pollen-foraging bees have promise to improve out-

crossing rates for the enhancement of genetic variation and perhaps even seed set

of desirable cultivars.

Acknowledgment

Voucher bees are deposited with the Illinois Natural History Survey. This article

is contribution number 17-91299IP of the Alabama Agricultural Experiment

Station, which partly supported this research (H-803).

Literature Cited

Adams, D. E., W. E. Perkins & J. R. Estes. 1981. Pollination systems in Paspalum dilatatum Poir.

(Poaceae): an example of insect pollination in a temperate grass. Am. J. Bot., 68: 389-394.

Cane, J. H. & J. A. Payne. 1988. Foraging ecology of the bee Habropoda laboriosa (Hymenoptera:

Anthophoridae), an oligolege of blueberries (Ericaceae: Vaccinium) in the southeastern United

States. Ann. Entomol. Soc. Am., 81: 419-427.

Chambers, V. H. 1945. British bees and wind-borne pollen. Nature, 155: 145.

Faegri, K. & L. van der Pijl. 1978. The principles of pollination ecology (3rd ed). Pergamon Press,

New York.

Hauptli, H. & S. Jain. 1985. Genetic variation in outcrossing rate and correlated floral traits in a

population of grain amaranth (Amaranthus cruentus L.). Genetica, 66:21-27.

Kaufman, C. S. 1979. Grain amaranth research: an approach to the development of a new crop. pp.

88-91. In Proc. second amaranth conf. Rodale Press, Emmaus, Pennsylvania.

Krombein, K. V., P. D. Hurd, Jr., D. R. Smith & B. D. Burks. 1979. Catalog of Hymenoptera in

America north of Mexico. Vol. 2. Smithsonian Institution Press, Washington, D.C.

Michener, C. D. & C. W. Rettenmeyer. 1956. The ethology of Andrena erythronii with comparative

data on other species (Hymenoptera, Andrenidae). Univ. Kansas Sci. Bull., 37: 645-684.

Munz, P. A. 1959. A California flora. University of California Press, Los Angeles.

Murphy, J. C. 1978. Pollination in the weedy amaranths. Abstr. Amer. Soc. Bot. Nat’l. Meet., 1978.

O’Neal, R. J. & G. D. Waller. 1984. On the pollen harvest by the honey bee (Apis mellifera L.) near

Tucson, Arizona (1976-1981). Desert Plants, 6: 81-109.

Raw, A. 1974. Pollen preferences of three Osmia species. Oikos, 25: 54-60.

Sharma, M. 1970. An analysis of pollen loads of honeybees from Kangra, India. Grana, 10: 35-42.

Singh, H. B. 1961. Grain amaranthus, buckwheat and chenopods. Indian Counc. Agric. Res. Cereal

Crop. Ser. I.

Wodehouse, R. P. 1971. Hayfever plants (2nd ed). Hafner Publishing Co., New York.

Received 17 May 1991; accepted 13 August 1991.

PAN-PACIFIC ENTOMOLOGIST

68(2): 100-104, (1992)

SPHERICAL HYPHAL BODIES OF PANDORA NEOAPHIDIS

(REMAUDIERE & HENNEBERT) HUMBER

(ZYGOMYCETES: ENTOMOPHTHORALES) ON

ACYRTHOSIPHON PISUM (HARRIS)

(HOMOPTERA: APHIDIDAE): A POTENTIAL

OVERWINTERING FORM

Ming-Guang Feng , 1 Robert M. Nowierski , 1 Robert E. Klein , 2

Albert L. Scharen , 3 and David C. Sands 3

1 Entomology Research Laboratory, Montana State University,

Bozeman, Montana 59717

irrigated Agriculture Research and Extension Center,

Washington State University, Prosser, Washington 99350

department of Plant Pathology, Montana State University,

Bozeman, Montana 59717

Abstract.— Cadavers of the pea aphid, Acyrthosiphon pisum (Harris), occurred abundantly on

commercial alfalfa in Kennewick, Washington, during late autumn of 1990. An aphid-specific

fungal pathogen, Pandora neoaphidis (Remaudiere & Hennebert) Humber, was responsible for

the death. Numerous hyphal bodies of the fungus inside the cadavers were spherical and averaged

11.5 (9.3-15.0) ftm in diameter (n = 100). Such spherical hyphal bodies apparently developed

from regular hyphal bodies forming septa, which has never been recorded for P. neoaphidis.

Over 300 cadavers collected in the field on 15 Oct were randomly sorted into three batches and

then maintained under different environmental conditions for studying the overwintering po¬

tential of the fungus. Cadavers maintained in a dark refrigerator at approximately 4° C or placed

within nylon-chiffon mesh bags (ca. 5x5 mm) and secured to the branches of shrubs (approx¬

imately 0.5 m above the ground in Bozeman, Montana) were capable of producing conidia and

infecting aphids in monthly observations from November to April with consistently visible snow

cover. In contrast, cadavers placed in polypropylene microcentrifuge tubes (38 x 13 mm), corked

with sterile cotton and then buried in the field soil (approximately 6 cm deep), were found to

have exhausted all their sporulation potential and infective capability in the first observation of

late November. The results indicate that P. neoaphidis may survive winter months in the form

of hyphal bodies on plant substrates above the ground rather than in the soil.

Key Words.— Insecta, Acyrthosiphon pisum, Entomophthorales, Pandora neoaphidis, aphid-spe¬

cific fungal pathogen, overwintering

A mycosis of the pea aphid, Acyrthosiphon pisum (Harris) (Homoptera: Aphid-

idae), was observed in a commercial alfalfa field in Kennewick, Washington,

during late September through mid-October, 1990. Alfalfa stems were heavily

infested with aphids (100% of the plants infested and more than 100 aphids per

alfalfa stem). Aphid cadavers, resulting from fungal infection, were observed in

abundance. Approximately 10% of the axillary shoots on alfalfa stems contained

at least one cadaver, and some of the shoots contained 10 or more.

Aphid cadavers collected on 27 Sep, 1 Oct and 9 Oct (the last date coinciding

with alfalfa harvest) were shipped via overnight mail to MGF in Bozeman, Mon¬

tana, for identification of pathogens involved. The aphid-specific fungus, Pandora

neoaphidis (Remaudiere & Hennebert) Humber (Zygomycetes, Entomophthora-

1992

FENG ET AL.: OVERWINTERING APHID PATHOGEN

101

les), was found to be the only pathogen responsible for the mycosis observed.

This was based on microscopic examination of nearly 200 cadavers individually

mounted on slides with aceto-orcein following maintenance in a moist chamber

at approximately 25° C for 20 h. No secondary infection by other entomophtho-

ralean fungi was detected.

Morphological features including conidiophores, conidia (Fig. la) and hyphal

bodies (Fig. lb) coincided well with those previously documented for P. neoaphidis

(e.g., Feng et al. 1990). Measurements of 100 primary conidia randomly taken

from 20 slides (cadavers) averaged 22.0 (17.5-27.5) x 11.3 (9.25-14.3) im i, falling

within the previously defined range of P. neoaphidis (Waterhouse & Brady 1982).

Spherical hyphal bodies (SHB) (Figs, lc, Id, lh), not previously documented

for P. neoaphidis, appeared with primary conidia and regular hyphal bodies (RHB)

in all the cadavers examined. The relative abundance of these unusual hyphal

bodies seemed to be negatively correlated with the abundance of primary conidia

and RHBs. Some of the cadavers were nearly filled with SHBs. The frequency of

cadavers containing SHBs tended to increase with each successive collection date.

The SHBs measured 11.5 (9.3-15.0) jum in diameter (n = 100), and were nearly

equal to the diameter (width) of primary conidia (Fig. la) and RHBs (Fig. lb).

Like uninucleate primary conidia of P. neoaphidis, most SHBs contained only a

single large nucleus (Fig. lc). Some SHBs were found to have two or more nuclei

(Fig. Id). SHBs with multiple nuclei were usually larger in size than those with

only one nucleus.

The SHBs apparently developed from RHBs, as shown in Figs, le-lh. A septum

sometimes appeared in the hyphal body, preceding the formation of a SHB (Figs,

le-lg). Septa are usually absent from the vegetative cells in the Entomophthorace-

ae (Humber 1989) and have never been recorded for P. neoaphidis. Subsequently,

the single cell separated by a septum became spherical, often at the end of the

hyphal body (Fig. lg). Eventually, the remainder of the hyphal body gradually

disappeared as its contents (protoplasts) entered the new SHBs (Fig. lh).

The appearance of SHBs late in the season suggests that SHBs may function as

an overwintering form in the life cycle of P. neoaphidis. This hypothesis was tested

by tracing the infectivity of cadavers collected from the field, then exposed to

different environments during winter months. Over 300 cadavers were collected

from uncut alfalfa plants on the border strips of the field in Kennewick on 15 Oct

and carried back to the laboratory in Bozeman. These cadavers were then separated

into 3 batches and about 15 from each batch were placed into polypropylene

microcentrifuge tubes (38 x 13 mm) corked with sterile cotton for two batches

or nylon-chiffon mesh bags (approximately 5x5 cm; four threads per mm) for

the third batch. The two batches of tubes were maintained in a dark refrigerator

of about 4° C and buried in the field soil (approximately 6 cm deep) on the Montana

State University campus, respectively. Mesh bags of the third batch were secured

to the branches of outdoor shrubs approximately 0.5 m above the ground on the

university campus. Thereafter, one tube or bag of cadavers was randomly taken

from each batch every month and used to inoculate aphids reared in the laboratory

by suspending the cadavers over the aphids for a spore shower. It was found that

cadavers hung in bushes or maintained in the refrigerator were capable of pro¬

ducing conidia and infecting aphids throughout the cold winter with consistently

visible snow cover from November to April. However, cadavers in the tubes

102

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Figure 1. Common morphological characteristics (a, b) and unusual spherical hyphal bodies [SHB]

(c-h) of P. neoaphidis associated with cadavers of A. pisum on alfalfa in Kennewick, Washington, in

autumn, (a) Primary conidia [PC], germinating PC [GPC] and secondary conidium [SC] formed from

PC. (b) Regular hyphal bodies [RHB]. (c) SHB uninucleate, (d) SHB binucleate. (e, f) Well-defined

septum [SP], at arrow, seen in RHB. (g) SHB forming at the end of RHB. (h) Newly-formed SHB and

remains [RM] of RHB. Scale bars: 20 /im; the bar for (a) also applies to (c-e) and (g, h).

buried in the soil were found to have exhausted all their sporulation potential in

the first observation on 30 Nov. A layer of conidia were then visible on the inside

wall of the tube and the cadavers became indistinguishable from one to another.

As a result, the cadavers in the soil could not infect aphids at that time. This

appeared to be attributable to the high humidity in the soil under snow cover

during the relatively mild November.

Therefore, our observations indicate that the P. neoaphidis hyphal bodies in

aphid cadavers can survive winter months only in relatively dry environments

1992

FENG ET AL.: OVERWINTERING APHID PATHOGEN

103

(e.g., on plant substrates above the ground) rather than in the moist soil, as

postulated by some authors (e.g., Latteur & Godefroid 1983). This is similar to

a report that hyphal bodies of P. neoaphidis may maintain infectivity in cadavers

for up to 32 weeks at regimes of 0° C and < 50% relative humidity (Wilding 1973).

In contrast, other entomophthoralean fungi generally overwinter as resting spores,

as seen in Conidiobolus obscurus (Hall & Dunn) Remaudiere & Keller (Latge et

al. 1978) and Zoophthora radicans (Brefeld) Batko (Perry & Regniere 1986).

Although it was claimed that resting spores of P. neoaphidis had been obtained

in vitro (Uziel & Kenneth 1986), resting spores have never been observed from

aphids infected by P. neoaphidis in the field. The SHBs observed in this study

are unlikely to be resting spores (zygospores or azygospores) because they are thin-

walled and too small for resting spores typically reported for the Entomophthorales

(R. A. Humber, personal communication). Resting spores have been observed in

the field for another entomophthoralean species, Entomophthora planchoniana

Cornu, but the primary overwintering form of this latter fungus is hyphal bodies

that are distinct from those usually found for the same species (Keller 1987).

SHBs in the pea aphids infected by P. neoaphidis late in the season seem to be

analogous to the hyphal bodies of E. planchoniana.

It remains unknown what environmental stimuli may induce the information

of septa in the hyphal bodies, thus forming the SHBs. During the period from 16

Sep to 15 Oct, 1990, local day length decreased from about 12.5 h to 11 h, while

the daily minimum temperature was 9.3 (range: 1.1-16.1)° C, daily maximum

22.4 (10.6-30.6)° C, and daily mean 15.7 (7.8-22.8)° C. Whether these environ¬

mental conditions (short day and low temperature) may be conducive to physi¬

ological changes in the aphid hosts, which in turn may influence fungal devel¬

opment, is unclear.

Finally, P. neoaphidis may require a variety of host species from different crops

or non-crop plants to complete the life cycle. Plant hosts that remain in the field

through late autumn or are perennial (e.g., alfalfa) may provide a source of in¬

oculum to initiate infections in aphid populations that infest spring and summer

crops the following year (e.g., small grains). This hypothesis warrants further

studies.

Acknowledgment

We thank R. A. Humber, S. A. Woods and D. A. Streett for their critical review

of this manuscript. This article is published as Journal Series No. J-2587 from

the Agricultural Experiment Station, Montana State University, Bozeman, Mon¬

tana.

Literature Cited

Feng, M. G., J. B. Johnson & L. P. Kish. 1990. Survey of entomopathogenic fungi naturally infecting

cereal aphids (Homoptera: Aphididae) of irrigated grain crops in southwestern Idaho. Environ.

Entomol., 19: 1534-1542.

Humber, R. A. 1989. Synopsis of a revised classification for the Entomophthorales (Zygomycotina).

Mycotaxon, 34: 441^-60.

Keller, S. 1987. Observations on the overwintering of Entomophthora planchoniana. J. Invertebr.

Pathol., 50: 333-335.

Latge, J. P., D. Perry, B. Papierok, J. Coremans-Pelseneer, G. Remaudiere & O. Reisinger. 1978.

104

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

Germination d’azygospore d' Entomophthora obscura Hall & Dunn, role du sol. C.R. Acad. Sci.

Paris Ser. D, 287: 946-946.

Latteur, G. & J. Godefroid. 1983. Trial of field treatments against cereal aphids with mycelium of

Erynia neoaphidis (Entomophthorales) produced in vitro, pp. 2-10. In Cavalloro, R. (ed.).

Aphid antagonists. A.A. Balkema, Rotterdam.

Perry, D. F. & J. Regniere. 1986. The role of fungal pathogens in spruce budworm population

dynamics: frequency and temporal relationships, pp. 167-170. In Samson, R. A., J. M. Vlak

& D. Peters (eds.). Fundamental and applied aspects of invertebrate pathology. Wageningen,

Netherlands.

Uziel, A. & R. G. Kenneth. 1986. In vitro resting-spore formation in Erynia neoaphidis. p. 230.

In Samson, R. A., J. M. Vlak & D. Peters (eds.).Fundamental and applied aspects of invertebrate

pathology. Wageningen, Netherlands.

Waterhouse, M. & B. L. Brady. 1982. Key to the species of Entomophthora sensu lato. Bull. Br.

Mycol. Soc., 16: 113-143.

Wilding, N. 1973. The survival of Entomophthora spp. in mummified aphids at different temper¬

atures and humidities. J. Invertebr. Pathol., 21: 309-311.

Received 25 June 1991; accepted 13 August 1991.

PAN-PACIFIC ENTOMOLOGIST

68(2): 105-121, (1992)

RECENT COLONIZATION OF THE

SAN FRANCISCO BAY AREA, CALIFORNIA,

BY EXOTIC MOTHS (LEPIDOPTERA: TINEOIDEA,

GELECHIOIDEA, TORTRICOIDEA, PYRALOIDEA)

Jerry A. Powell

Department of Entomological Sciences, University of Califomia-Berkeley,

Berkeley, California 94720

Abstract. — Records are given documenting the establishment of seven species of moths in the

San Francisco Bay area, California, during 1955-1988: Opogona omoscopa (Meyrick) (Tineidae),

Oegoconia quadripuncta (Haworth) (Blastobasidae), Mirificarma eburnella (Denis & Schiffer-

miiller) (Gelechiidae), Crocidosema plebiana Zeller (Tortricidae), and three pyraloids, Uresiphita

reversalis (Guenee), Parapediasia teterrella (Zincken), and Achroia grisella (Fabr.). Ten additional

Microlepidoptera that have colonized this region in the past 50 years are tabulated with literature

sources. Most of these species spread to the San Francisco area after establishment in southern

California, often following long periods (17-60 years) of naturalization there.

Key Words. — Insecta, Lepidoptera, Tineoidea, Gelechioidea, Tortricoidea, Pyraloidea, coloni¬

zation

Insects make up an important part of the alien fauna that has been transported

by humans to colonize different parts of the world. By 1982, 1700 such immigrants

had become established in the 48 contiguous U.S. states, including 134 Lepidop¬

tera (Sailer 1983). Although Sailer calculated that Lepidoptera are poorly repre¬

sented, relative to their species numbers, as contrasted to Coleoptera, Hymenop-

tera and particularly Homoptera, some Microlepidoptera and Pyraloidea have

become frequent travelers via their association with human activities.

California is an adopted home to more than 60 species of these smaller moths,

including many of our most notorious insects, in households (clothes moths, stored

foods moths), gardens (e.g., azalea leafminer, buddleia budworm, cotoneaster

webworm), and agricultural situations (codling moth, Oriental fruit moth, pink

bollworm, etc.). Other species are detritivores, scavengers, or fungus feeders and

seldom attract attention. For example, Opogona omoscopa (Meyrick), Nemapogon

granellus (L.), Oinophila v-flavum (Haworth), Batia lunaris (Haworth), Endrosis

sarcitrella (L.), and Oegoconia quadripuncta (Haworth) are all common members

of the urban insect community in California but are seldom noticed except by

lepidopterists. A few adventive colonists are even encouraged for possible weed

suppression, such as Agonopteryx alstroemeriana (Clerck), a specialist on poison

hemlock, and A. nervosa (Haworth) and Uresiphita reversalis (Guenee), which

feed on genista and other brooms, although the last species sometimes eats other

ornamental legumes or native lupines and causes mixed emotions, varying with

circumstances.

Many of these lepidopterous colonists became established in California so early

in the immigration of European and Oriental humans that a history of their

introduction and spread cannot be reconstructed. There is essentially no record

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

of the Microlepidoptera fauna of the Pacific coast of North America prior to the

remarkable expedition in 1871-1872 in northern California and Oregon by Lord

Walsingham, during which he collected and later described many of our native

species (see Essig 1941, Powell 1964a: 5). More extensive collections in urban

and agricultural situations were made by Koebele and Coquillett during the 1880s

and 1890s, primarily in Alameda and Los Angeles counties. Otherwise, there are

few records of Microlepidoptera in California prior to the turn of the century,

and any other record that may have existed of the fauna in the San Francisco Bay

area during the 19th century was lost in the 1906 fire that destroyed the collections

of the California Academy of Sciences.

Despite federal and state efforts at quarantine against imported insects, as the

human population has increased and ease of transportation improved, the parade

of incoming insects has continued. California’s population increased an appalling

25%, and that of the S. F. Bay area 16%, during the 1980s alone. Hence, it is not

surprising that at least 17 species of small moths have taken up residence in this

area during the past half century (Table 1). Included are six species that appear

to have colonized during the 1980s. The occurrence of two of these is documented

elsewhere: Athrips rancidella (Powell 1985) (Fig. 4) and Agonopteryx alstroemer-

iana (Powell & Passoa in press). Here, I give data for the remaining five, and for

three species that have been established for longer periods, but apparently not

documented in detail in the literature.

Methods. — I recovered data from specimens in the major California collections

through 1990, by searching the unidentified accessions and confirming identifi¬

cations in the determined material. Voucher specimens in collections are indicated

in the text by the following abbreviations:

CAS, California Academy of Sciences, San Francisco; CDFA, California De¬

partment of Food & Agriculture, Sacramento; EME, Essig Museum of Entomol¬

ogy, University of California, Berkeley; FAC, Fresno County Agricultural Com¬

missioner’s Office, Fresno; LACM, Los Angeles County Museum of Natural

History; SDNH, San Diego Natural History Museum; SJAC, San Joaquin County

Agricultural Commissioner’s Office, Stockton; SJS, San Jose State University,

Department of Entomology; UCD, University of California, Davis, Bohart En¬

tomological Museum; USNM, U.S. National Museum of Natural History, Wash¬

ington, D.C. In addition, card- and computer-file records at CDFA were made

available. Most of these are not represented by voucher specimens. Data from

the identified material in the USNM were recorded, but not from unidentified

accessions, through 1977 ( Opogona, Oegoconia, Achroia ) and 1988 ( Crocidose -

ma).

I made blacklight trap collections in suburban sites at Walnut Creek, Contra

Costa Co., from 1961 to 1973 (EME). In the first six years, samples were made

most nights I was in residence, near the foot of Shell Ridge, while those during

August, 1966, to 1973 were sporadic, at a site near San Ramon Creek. The two

localities are respectively about 2.75 airline km NW and 2.5 km SW of the

Highway 24-680 interchange. I recorded moths in urban Berkeley on nearly all

dates I was residence from May 1978 through 1990. During 1978-June, 1984, I

sampled at a site 3.0 airline km NNW of the University of California west gate

and during July 1984 through 1990, at a second locality 0.33 airline km north of

the 1978-1984 site. This area has been residential since 1915-1920.

1992

POWELL: MOTH COLONIZATION

107

Table 1. Exotic Microlepidoptera and Pyraloidea that became established in the San Francisco

Bay area during 1939-1988. (1 = local colonization; w = widespread occurrence in S. F. Bay area;

u = uncertain status.)

Taxa

Earliest record

Present

status

Source

Tineidae:

Oiophila v-flavum (Haworth)

1947, Stanford

Tilden 1951, Powell

1964b

Opogona omoscopa (Meyrick)

1972, Berkeley

CDFA, Davis 1978

Oecophoridae:

Agonopteryx alstroemeriana

1983, Berkeley

Powell & Passoa, in press

(Clerck)

Batia lunaris (Haworth)

1956, Marin

Powell 1964c

Esperia sulphurella (Fabr.)

1966, El Cerrito

Powell 1968

Pyramidobela angelarum Keifer

1942, San Jose,

San Mateo

Keifer 1942

Blastobasidae:

Oegoconia quadripuncta (Haworth)

1959, Redwood

City, 1976,

Berkeley

present data

Symmoca signatella

(Herrich-Schaeffer)

1959, Redwood

City

Powell 1960

Gelechiidae:

Athrips rancidella

1983, Berkeley

Powell 1985

(Herrich-Schaeffer)

Mirificarma eburnella

1985, Morgan Hill

present data

(Denis & Schiffermiiller)

Tortricidae:

Crocidosema plebiana Zeller

1988, Berkeley

present data

Spilonota ocellana (Denis & Schiffer-

1939, San Jose

Keifer 1939

muller)

Cnephasia longana (Haworth)

1947, San Mateo

Keifer 1948, Powell 1964a

Platynota stultana (Walsingham)

1967, Antioch,

Albany

Powell 1983

Pyralidae:

Uresiphita reversalis (Guenee)

(1966, Stevens

Cr.), 1980, San

Jose

present data

Parapediasia teterrella (Zincken)

1988, Berkeley

present data

Achroia grisella (Fabr.)

1955, San Jose

present data

Tineidae

Opogona omoscopa (Meyrick)

(Fig. 1)

Opogona omoscopa was originally described from Australia in 1893 and since

has been found widely distributed in pan-global warm regions, probably in large

part the result of man’s activities. The larvae feed in decaying, often damp plant

108

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

i i i * i i i i i i i i l i l i i i i i i l i i l < i i i i i i i i i i m i i i i i i i i i i i i i

i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i i a i i i i i i i i i i i i i i

Figures 1-8. Figure 1. Opogona omoscopa-, Berkeley, October 1978. Figure 2. Oegoconia quadri-

puncta; Berkeley, June 1988. Figure 3. Mirificarma eburnella; Nevada Co., May 1980. Figure 4. Athrips

rancidella; Berkeley, May 1983. Figure 5. Crocidosema plebiana Zeller; Berkeley, November 1990.

Figure 6. Uresiphita reversalis\ Berkeley, December 1983. Figure 7. Achroia grisella; Berkeley, August

1983. Figure 8. Parapediasia teterrella; Berkeley, May 1989.

material, including wood, bark and dead leaves (Davis 1978) and evidently are

easily transported with roots and other plant material. This species has been known

in Hawaii since 1905, where it is widespread and abundant (Zimmerman 1978),

a likely source for introduction into California.

1992

POWELL: MOTH COLONIZATION

109

The earliest Pacific coast record is at Goleta, Santa Barbara Co., California, in

May, 1969 (CDFA and Davis 1978), which probably was soon after establishment,

because the adults come to lights readily, and I collected at Goleta for five weeks

during June and July, 1965, without finding O. omoscopa. The moths were taken

at nearby Santa Barbara and Summerland in June and July, 1969 (USNM). In

fall, 1970, C. Nagano collected a series at Santa Monica, Los Angeles Co. (LACM),

and by summer, 1971,0. omoscopa was widespread in southern California, having

been detected in Gardena (LACM), Los Angeles (USNM) and Rancho Santa Fe,

San Diego Co. (EME).

The first records in the S. F. Bay area were two larval collections on ginger

roots in a market in Berkeley, in May, 1972, and May, 1973 (CDFA). Ginger

roots sold in this area are normally imported from Hawaii. Hence, it is possible

that a separate introduction from overseas, rather than from southern California,

initiated the S. F. Bay area population. There also were larval collections from

Corte Madera, Marin Co. in January, 1974, and Fremont and Livermore, Alameda

Co. in 1976 (CDFA), but we do not have documented records of colonies outside

of buildings until adults began appearing at lights in the ‘east bay’ in 1978 (EME).

The species has been common in Berkeley since that time, having been recorded

on 15-30 dates each year. The moths are seen in every month but are most

prevalent in September-November (50% of all records during 1985-1990: JAP,

unpublished data).

The peculiar, widely divergent labial palpi, flattened, smooth front and elongate,

plicate maxillary palpi make O. omoscopa easily recognizable among all California

Lepidoptera.

Blastobasidae

Oegoconia quadripuncta (Haworth)

(Fig- 2)

This Palaearctic species is distinctive in the urban fauna of California, having

black forewings spotted with yellow. In Europe the larvae are reported to feed on

decaying vegetable matter, and we reared O. quadripuncta from leaf litter beneath

Quercus by P. Rude (EME). In England there is a single annual generation, with

adults active in July and August, occurring in habitats such as hedge-bottoms

(Emmet 1979).

Oegoconia quadripuncta was introduced into the Atlantic states long ago; it was

redescribed as novimundi Busck, a synonym, in 1915, and it was established in

Pennsylvania and New jersey by 1920 (USNM). The adventive range had reached

Washington, D.C. by 1927, Martha’s Vineyard, Massachusetts by 1941 and Il¬

linois by 1956 (USNM).

There does not seem to be a published report of this species’ occurrence in

California, such as during Keifer’s summaries of introductions during 1935-1955

(Powell 1991). However, O. quadripuncta evidently was introduced into southern

California, presumably from the eastern U.S., more than 50 years ago. There are

specimens from South Pasadena, L. A. Co., collected in August, 1938, and June,

1940 by C. Henne (USNM), and the range extended to Ventura Co. (Ojai) by

1961 (EME) and Orange Co. by 1962 (CDFA). The species had become common

in Los Angeles by the time Donahue began sampling there in the early 1970s

(LACM).

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

The first record I have seen in the S. F. Bay area is August, 1959, at Redwood

City, San Mateo Co. (EME), but O. quadripuncta was not known east of the bay

until adults appeared at lights in Berkeley in 1976. The species seems to be

becoming more prevalent at Berkeley; it was observed on two or three dates per

season until 1986, then five dates in 1987 and 1988, six in 1989, and nine in

1990 (despite 44 nights absence from sampling during summer), when the flight

period extended from early June to mid September.

Oegoconia evidently colonized the Central Valley about a decade later than the

S. F. Bay area. I did not find the species in Davis when I sampled there in 1956,

but there are more than a dozen collections records from Sacramento (1967—

1968), Davis (1969-1971) and Fresno (1970-1971) (CDFA, UCD).

Gelechiidae

Miriftcarma eburnella (Denis & Schiffermuller)

(Fig. 3)

This moth was reported in North America under the names M. formosella

(Hubner) (Anonymous 1969, Dowell & Gill 1989) and M. flamella (Hubner)

(Hodges 1983), which are considered to be synonyms of M. eburnella (Pitkin

1984). The species is widespread in Europe and the Mediterranean region, where

it feeds on Medic ago, including alfalfa, and other legumes (Pitkin 1984).

This gelechiid is distinctive in the California fauna, having rust-orange and

yellow patterned forewings. It was first recognized in North America when larvae

were found defoliating Ladino clover, Trifolium repens L., in the Sacramento

Valley in Sutter, Placer and Sacramento counties, in April, 1969. Identifications

at the time revealed that I had collected specimens near Georgetown, El Dorado

Co., in June, 1967 (Anonymous 1969; CDFA, unpublished report). The species

was already widely established, however, as evidenced by specimens determined

later that had been taken by A. Keuter and G. Keuter in May, 1965, and May-

June, 1967, at Citrus Heights, Sacramento Co. (CAS). There are also two speci¬

mens labelled 12 Oct 1967, in the Keuter material, suggesting a bivoltine life

cycle.

Berkeley students and I found M. eburnella at additional localities in El Dorado

Co. (Greenville, Somerset) during May and June, 1967 and 1978, and at several

sites around the Sierra Foothill Field Station (near Smartville), Yuba Co. and

Rough and Ready, Nevada Co., in May, 1980. The species appeared at La Grange,

Stanislaus Co. in 1971 (CDFA), at a site that has been sampled for many years

by R. P. Allen. The 1971-1980 localities are 55 km NW to 134 km SE of a line

between Citrus Heights and Georgetown, along the foothills of the Sierra Nevada.

During a census of Lepidoptera of serpentine grasslands in Santa Clara Co., D.

D. Murphy and I collected two specimens of M. eburnella at Kirby Canyon Ridge

(approx. 6 airline km NE of Morgan Hill), on 29 Apr 1985. This suggested that

populations of this gelechiid had spread across the Central Valley and inner Coast

Range into the Santa Clara Valley. However, more intensive survey on numerous

dates at this locality and serpentine grasslands at a dozen other sites in Santa

Clara, San Mateo and Marin counties during March through May, 1986-1987

and 1990 failed to recover M. eburnella. Possibly the four year drought following

1985 suppressed the clover or other hostplants severely, limiting or eradicating

1992

POWELL: MOTH COLONIZATION

111

the immigrant moth population from this habitat. Hence, the residency status of

M. eburnella in the S. F. Bay area is uncertain.

Tortricidae

Crocidosema plebiana Zeller

(Fig. 5)

This species was originally described from Sicily in 1847, but its native distri¬

bution is unknown. It occurs pan-globally in warmer regions, probably having

been transported by man since early times. Crocidosema plebiana was reported

from Hawaii by several early authors, but Zimmerman (1978) regards the Ha¬

waiian Crocidosema as three distinct, endemic species. Differentiation of Pacific

island populations also is discussed by Clarke (1971, 1986). The evidence suggests

that C. plebiana (sens, lat.) occupied a broad range, and that North American

populations probably originated from the Mediterranean. The larvae of C. ple¬

biana feed in flowers and fruit of various Malvaceae, including Hibiscus , and

have been taken on cotton several times in California.

Heinrich (1923) reported C. plebiana in the U.S. from California and Texas.

In addition, the species has been collected widely in the south, in Louisiana (1916),

Florida (1918 onwards) and South Carolina (1944) (USNM; Kimball 1965). The

species has long been established and abundant in southern California; the earliest

available record is June, 1911 at San Diego, collected by W. S. Wright (USNM).

In 1917-1918, it was collected at Chula Vista, San Diego Co. (CAS). By about

1920 it occurred in the San Bernardino area (Barnes collection: USNM), at Riv¬

erside by 1932, and on Santa Catalina Island by 1931 (CDFA, LACM). By that

time probably it was established throughout much of the Los Angeles basin and

Orange Co., where its colonization was documented in the 1940s during the

statewide Oriental fruit moth survey by dimalt bait traps (CDFA). Specimens

were reared from Hibiscus buds at Exposition Park, Los Angeles in 1942 (LACM).

The distribution also extended to the coast in the Ventura (1943) and Santa

Barbara (1936) areas (CDFA, LACM), and San Luis Obispo Co. (Pismo Beach)

by 1959 (EME). Crocidosema plebiana was found in the San Joaquin Valley in

Kern Co. in 1968 (CDFA).

I had not seen any subsequent records north of a line between Pismo and

Bakersfield until C. plebiana appeared in Berkeley recently. In late September

and October, 1988, two males came to a blacklight, but none was observed in

1989, suggesting that the moths captured in 1988 did not represent an established

population. In 1990, however, C. plebiana reappeared, with males attracted to

blacklight on 9, 12 Jul and on 10 dates between 11 Sep and 17 Oct, confirming

the colonization.

Pyralidae

Uresiphita reversalis (Guenee)

(Fig. 6)

This large pyraustine, which is known as “the genista caterpillar,” has bright

rust-brown forewings and ochreous-yellow hindwings. The moths are primarily

nocturnal and come to lights but are easily flushed into activity during the daytime.

The larvae are aposematic in color and behavior; they are orange and black

spotted, live exposed, without a shelter, and are rendered distasteful by sequestered

112

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

alkaloids (Bemays & Montllor 1989). They often occur in defoliating numbers.

Hence, populations are easily seen in the field, and this species is not likely to

colonize unnoticed for long.

Although its relatives are Old World species, Uresiphita reversalis is believed

to be a native Nearctic species, having been described originally in 1854 from

“North America” without a specified locality. The natural distribution is un¬

known, but it may have encompassed parts of the southeastern U.S. and Mexico.

The present range is reported to be “southern Canada to southern Florida and

west to California” (Munroe 1976), but it is likely sustained in northern areas by

migrations, not continuously resident populations. In California, moreover, pop¬

ulations are dependent upon introduced plants, particularly Genista, grown as

ornamentals or in weedy situations, and there are no known records prior to 1930,

so U. reversalis is assumed to be an introduced or adventive exotic.

In the west there are records as early as 1912 in the Davis Mountains, Texas

(LACM) and 1927 in the mountains of southern Arizona (CAS), and there are

scattered collections from the Mexican plateau and coastal Sinaloa (EME, UCD),

suggesting that native populations may have lived in these areas.

The earliest known record in California is a series collected in Los Angeles in

September, 1930, by J. A. Comstock (LACM). By late 1931, U. reversalis occurred

widely in urban Los Angeles, Orange, San Diego and Ventura counties and had

been reared from “Genista and other brooms” at several localities (Keifer 1931).

There are records at Riverside and San Bernardino by September, 1932, and Santa

Barbara in 1933 (CDFA, CAS, LACM). McKenzie (1933), who described the

early stages and recorded hostplants, stated that the initial appearance of this

insect in California had been noted only recently.

Populations seemed to stabilize in cismontane southern California during the

following 30 years, and there are collection records for nearly every year, indicating

that residency was continuous.

The earliest known collection of U. reversalis in the San Francisco Bay area is

August, 1966, at Stevens Creek [5 km SW of Cupertino], Santa Clara Co., by R.

Denno (UCD). This record is puzzling because one would expect this species to

have appeared first in an urban area, rather than a forested canyon in the foothills,

and because if the collection sampled an established population, it is surprising

that no other colonies were detected in the south bay area during the subsequent

13 years. Continuous residency is documented beginning in 1980. Larvae were

collected from Laburnum in Fremont in July, 1980 (CDFA) and on Genista at

San Jose at least three times between September, 1980, and September, 1981

(CDFA, EME, SJS), the first by F. litis (W. E. Ferguson, personal communication);

and the species rapidly colonized northward in the S. F. Bay area during the next

10 years. In 1983, larvae of U. reversalis were found in Oakland by P. Neyland

(EME), and adults began appearing at localities that R. L. Langston, W. W.

Middlekauff or I had been sampling regularly: Antioch, Contra Costa Co. (August),

Berkeley (November), San Bruno Mts., San Mateo Co. (December), in Contra

Costa Co. at El Cerrito the following year, and Fish Ranch Canyon and Kensington

by 1985 (CAS, EME).

Uresiphita reversalis was widely established north of San Francisco Bay by the

late 1980s, in Marin (1986) (including Angel Island and Marin Island, 1989),

Napa (1988) and Solano (1987) counties (CDFA, EME).

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POWELL: MOTH COLONIZATION

113

It is likely that U. reversalis also spread through the Central Valley and Sierra

Nevada foothills during or preceding the 1980s. By 1968, when records were

suspended in a card file system at CDFA 1 , there were no listings of this pyralid

from counties north of the Transverse Ranges; also, there are no voucher speci¬

mens for records during the following 11 years. Larvae were collected at Tracy,

San Joaquin Co. in late 1982 (SJAC), and at Bakersfield, Kern Co. and near

Fresno, Fresno Co. a year later (CDFA, FAC). By 1987, when computerization

came on line at CDFA, U. reversalis was found in Kern, Tulare, Merced, Placer,

and Sacramento counties; by 1988 in Yolo Co. (January) and in the northern

Sacramento Valley, in Sutter and Butte (October), and Shasta (1989) counties

(CDFA) (Fig. 9). Simultaneously, Uresiphita colonized the foothills of the Sierra

Nevada in Amador, Nevada, and Tuolumne counties, to elevations of 550-800

m at Sonora and Grass Valley (CDFA).

The genista caterpillar feeds on a variety of legumes, particularly those of the

tribe Genisteae (Fabaceae) including Genista, Cytisus, and Lupinus, as well as on

Baptisia and Sophora (Dyar 1901, McKenzie 1933, Kimball 1965, Munroe 1976,

Bemays & Montllor 1989). In California, the adventive populations evidently are

dependent primarily on Genista (= Cytisus ) monspessulana (L.) (French broom)

and horticultural hybrids. Cytisus scoparius (L.) (Scotch broom) has been recorded

as the larval host on several occasions, but at least some of these evidently originate

from plant misidentifications or equating the common names “genista,” “broom”

and “Scotch broom” as applied to various Genista species. For example, there

are records from “Scotch broom” in San Diego County in 1931 (CDFA) and 1967

(EME), but Cytisus scoparius was not established anyplace south of Monterey

County by 1978 (Mountjoy 1979).

There are numerous records of larvae having been collected on nonleguminous

plants, including Buddleia (Loganiaceae) (McKenzie 1933; CDFA [1964]), as¬

paragus fern (Liliaceae), Taxus (Taxaceae), Gardenia (Rubiaceae) (CDFA), and

“chamise” (SDNH). Such records, along with other evidence (“pupating in door¬

way,” “barbeque cover,” etc. [CDFA]), probably reflect a propensity of late instar

larvae to wander. Particularly when colony densities are high and Genista is

defoliated, larvae of U. reversalis are liable to be found on various other plants

in the vicinity.

In the S. F. Bay area, larvae of U. reversalis feed on native Lupinus, including

L. chamissonis Eschscholtz when growing in proximity to Genista monspessulana

(Pt. Molate, JAP 87G4, EME), and on L. arboreus Sims according to Bemays &

Montllor (1989). In no-choice feeding tests in the laboratory, Bemays and Montllor

found that larvae did not feed and soon died when offered certain legumes, in¬

cluding Medicago, Trifolium, Vicia, and Pickeringia. They fed successfully on

1 Uresiphita reversalis is rated “C” in pest status by CDFA (a native or established species, against

which no agricultural quarantine action may be needed). Most “C” and “D” (beneficial or non-

phytophagous non-economic) rated insects ceased to be routinely entered in the CDFA card system

in 1968, due to system size restrictions; those rated “Q” [old “X”] (unassessed exotic), “A” (quarantine

action mandated) or “B” (county level quarantine), however, continued to be automatically entered

in the post-1968 card database. Individual instances of “C” or “D” rated insects also could have been

entered, if requested by a CDFA taxonomic specialist for the group; their post-1968 absence on cards

does not necessarily mean that no CDFA identification was done. In 1987, the CDFA data system

was computerized, and all data from CDFA identifications was once again routinely entered.—(Ed.)

114

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Figure 9. Geographical distribution of Uresiphita reversalis in California: by 1931 (shaded area);

later dated localities refer to first records in peripheral areas of southern California and first records

in counties north of the Transverse Ranges.

Lupinus arboreus, Cytisus striatus (Hill), G. monspessulana, and Cytisus scoparius,

and late instar larvae significantly preferred Lupinus over G. monspessulana, when

given the choice.

Bemays & Montllor (1989) believed that the data indicate that the main hosts

of U. reversalis in California are species of Lupinus. However, I have not seen

any evidence that populations inhabit less disturbed plant communities where

they would be sustained solely by native plants. Moreover, there are no specimen

voucher records of larvae on native plants outside of urban situations after more

than half a century residency in southern California and none in more northern

areas (CAS, CDFA, EME, LACM, SDNH, SJS, UCD). The CDFA has records

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POWELL: MOTH COLONIZATION

115

of larval collections from “Lupinus sp.” from Castro Valley, Alameda Co. (1988),

Redding, Shasta Co. (1989) and Santa Maria, Santa Barbara Co. (1988) in garden,

park, and nursery settings. Some exotic ornamental legumes also serve as hosts,

including Laburnum at El Cerrito (JAP 84K1) (EME) and Piptanthus at the

Strybing Arboretum, San Francisco (CAS).

Parapediasia teterrella (Zincken)

(Fig. 7)

Described in 1821 from Georgia, this was one of the first pyralids known in

North America. It is widespread in the eastern U.S. and is often extremely abun¬

dant at lights in urban areas, such as around Washington, D.C. The original

geographical distribution no doubt was modified by human colonization of North

America; by the late 1800s it encompassed the Atlantic and midwestem states.

Murtfeldt (1893) reported that P. teterrella had become more abundant during

the past two or three years around Kirkwood, Missouri, than all other crambids

combined.

There are records in the southwest as early as half century ago: Tulsa, Oklahoma

(1940); Albuquerque, New Mexico (1944); Tucson (1935) and Madera Canyon

(1947), Arizona (LACM). Hence, P. teterrella may have spread into that region

with urbanization during the early 1900s.

The earliest known occurrence in California is August, 1954, at South Gate,

Los Angeles Co. (LACM). Records from other parts of southern California and

the Central Valley indicate that this lawn moth had colonized in the early 1950s,

then spread southward and northward within a few years (Fig. 10). I collected the

urban lawn moths, Crambus sperryellus Klots and Tehama bonifatella (Hulst)

and did not find Parapediasia teterrella in San Diego during 1953-1956; but in

1957-1959, light trapping by A. A. Lee and R. A. Mackie produced P. teterrella

at widely separated inland localities: Escondido and Otay, San Diego Co. and at

several coastal sites in 1958-1959 (SDNM, EME). The colonization reached

Bakersfield, Kern Co., by July, 1959, then quickly spread through the Central

Valley, recorded at Madera (1960), Sacramento (1967), and Davis, Yolo Co.

(1969) (CDFA). I did not find P. teterrella in urban Davis when I sampled there

during the summer of 1956.

The adventive populations appear to have extended through the delta region

to S. F. Bay, having been recorded at the Antioch National Wildlife Refuge in

August, 1981, and at Berkeley beginning in 1988 (EME), when two adults came

to blacklight in October. The following season I recorded P. teterrella on 16 dates

between 19 May and 17 Oct; in 1990 the species became more abundant and

seasonally extensive, flying from 9 Apr to 24 Oct (34 dates recorded), often with

5-10 individuals observed. At Berkeley this species has become the most prevalent

lawn moth, while Tehama bonifatella (Hulst) appears to have declined in numbers

(eight and 13 dates in 1989 and 1990, down from 19-29 dates in 1985-1988),

although its adult activity was more prolonged than that of P. teterrella in 1990

(25 Mar to 12 Nov). The data suggest that competitive displacement is occurring

at this site.

Parapediasia teterrella may be better adapted to inland than coastal areas in

California, as I have not seen records of its occurrence in urban areas of the coastal

counties, from Ventura to San Francisco.

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

Figure 10. Geographical distribution of Parapediasia teterrella in California: dated localities refer

to first record in each country.

Achroia grisella (Fabr.)

(Fig. 8)

The lesser wax moth is described in the stored products and general entomo¬

logical literature as a cosmopolitan insect, but evidently it has not been formally

reported in California. Achroia grisella is not mentioned by Essig (1926), nor by

Keifer during 1927-1954 (Powell 1991); and there were no records from the Pacific

states in the USNM in 1977. Larvae of this moth, which is uniformly mouse gray

with a contrasting pale yellow head, typically live in old honeycombs but also are

said to feed on dried fruit and “apparently” on dried insects (Forbes 1923). The

species was originally described from Europe but has been widely established in

the Atlantic states at least since the 1890s (USNM).

1992

POWELL: MOTH COLONIZATION

117

There are records of Achroia grisella in southern California dating back to the

early 1900s, but apparently the adults are not readily attracted to lights, and

populations likely have been more prevalent and widespread than records indicate.

A series was collected by W. S. Wright in San Diego on at least six dates between

1908 and 1915 (SDNH); there are two specimens from the E. Piazza collection,

probably from San Diego, taken in 1921, and A. grisella was taken at Del Mar,

San Diego Co. in 1934-1942 (CDFA, LACM).

Circumstantial evidence suggests that the lesser wax moth colonized central

parts of California at a much later date; there are at least 20 collection records

from the San Francisco Bay area and Sacramento Valley in the past 40 years but

none before that. The earliest vouchered record that I have seen is September,

1952, at Courtland, Sacramento Co. (CDFA); but probably A. grisella was wide¬

spread in central California by that time, as there are specimens from San Jose,

Santa Clara Co. taken in 1955 by J. W. Tilden (SJS) and from Prunedale and

Soledad, Monterey Co., in 1956 (CDFA). In the east bay, I took one specimen in

13 years sampling at Walnut Creek, Contra Costa Co. (June, 1964), and adults

have been collected sporadically in Berkeley since 1968 (EME). A long series of

A. grisella was reared by P. A. Rude from larvae in the honeycombs of an aban¬

doned beehive in Kensington in 1978 (EME), but only four individuals, taken on

four dates in 1983, 1987 and 1989, have been observed during the past 12 years

sampling in Berkeley.

Discussion

Collection records indicate that at least 17 species of exotic Microlepidoptera

and pyraloid moths have colonized the San Francisco Bay area during the past

half century, including six during the most recent 10 years (Table 1). Five of these

evidently were introduced independently from other populations in California,

either directly from the Old World ( Batia lunaris, Esperia sulphurella), or from

the Pacific northwest or eastern U.S. ( Agonopteryx alstroemeriana, Athrips ran-

cidella, Cnephasia longana ). The others colonized secondarily from southern

California, by local introduction or gradual spread by adventive populations.

Among species that have reached the S. F. Bay area via southern California,

several underwent a sequence of introduction-establishment then a long period

of naturalization, followed by rapid range extension northward (e.g., Fig. 9) (or

colonization via secondary introduction in the bay area). This pattern parallels

that shown by other introduced insects, for example the passion vine-feeding

butterfly, Agraulis vanillae (L.) (Powell 1961), the Old World earwig, Euborellia

annulipes (Lucas) (Langston & Powell 1975) in California, and the European

hesperiid, Thymelicus lineola (Ochsenheimer) in midwestem and northeastern

U.S. and adjacent Canada (Powell 1983). Such delayed ecogeographical expan¬

sions by introduced insects may involve genetic adaptation to environmental

situations to which the founder or even source populations were not adapted. The

delay cannot always be documented because of incomplete records of adventive

populations while they are at low levels, but gaps appear to have been as much

as 17 years for Symmoca signatella, at least 40 years for Platynota stultana, 50

for Uresiphita reversalis, and 60 for Crocidosema plebiana, following widespread

establishment in southern California.

By contrast, a few species have colonized in southern California and then

apparently began expanding their range without appreciable delay during natu-

118

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

ralization. Parapediasia teterrella (Fig. 10) colonized the Sacramento Valley within

6-13 years after detection in the Los Angeles basin (but 21 more years passed

before establishment in the east bay); Pyramidobela angelarum was established

in Santa Clara and San Mateo counties eight years after its discovery in Los

Angeles (Keifer 1942), and Opogona omoscopa reached the bay area within three

years of first notice at Santa Barbara, although this may have been via independent

introduction, and was widely established after two (San Diego Co.) to nine years

(S. F. Bay area).

The data are too fragmentary to document the history of Oinophila v-flavum

(Powell 1964b) and Achroia grisella in California. It would not be surprising to

discover that such species have been established in the S. F. Bay area for a half

century or more, as was the case for the urban tortricids, Acleris variegana (Schif-

fermuller) (Powell 1964a) in the bay area and Clepsis unifasciana (Hubner) in the

Pacific northwest (Powell 1988).

Dowell & Gill (1989) compiled a list of 208 invertebrates that they classified

as exotic and believed had been discovered in California between 1955 and 1988,

based on several USD A and CDF A publications. They include 24 species of

Lepidoptera, of which 16 are Microlepidoptera and Pyraloidea. The list is neither

complete nor restricted to exotic species. Included are at least four species that

likely are native insects:

Bucculatrix tridenticola Braun (erroneously given as Brown), which was origi¬

nally described in 1963 from southern and eastern Oregon, eastern Washington,

Colorado, Utah, and Nevada, occurs in association with Artemisia tridentata

Nuttall in natural communities in Modoc County, California (several records in

1960s: Hall 1965; EME) and probably throughout the Great Basin. The probable

origin inexplicably was given by Dowell and Gill as eastern U.S.

Periploca nigra Hodges was described originally from Sacramento in 1962 and

was found to be widely established in the S. F. Bay area on ornamental junipers

(Koehler & Tauber 1964). This may be an introduced species, but it is reported

to range from New York to Louisiana “then west to Sacramento and San Diego,

California” (Hodges 1978). The natural hostplants and geographical distribution

in the west are unknown.

Choristoneura conflictana (Walker) is widespread across boreal America in

association with Populus tremuloides Michaux and was reported from several sites

in native aspen forests of the Warner Mountains, Cascades and Sierra Nevada,

having been collected in California from 1922-1962 (Powell 1964a).

Eumysia mysiella (Dyar) is a widespread native insect of the Great Basin and

southwest. It was described from Stockton, Utah, in 1905 and by the 1960s was

recorded in Arizona, New Mexico, and Nevada (Heinrich 1956) (EME). Probably

its natural range included California, east of the Sierra Nevada. The larval host

is unrecorded, but the closely related E. idahoensis Mackie feeds on several species

of Atriplex (Chenopodiaceae) (Mackie 1958).

Dowell and Gill’s list omits several species that were first detected in California

between 1955-1988, including Opogona omoscopa, discussed above; Batia lu-

naris, which was established on both sides of S. F. Bay by 1962 (Powell 1964c);

Esperia sulphurella, an early spring, diurnal moth that was discovered at El Cerrito

and Berkeley in 1966 and 1967 (Powell 1968) and has been recorded many times

during the subsequent two decades (CAS, EME); and Agonopteryx alstroemeriana,

1992

POWELL: MOTH COLONIZATION

119

which was widely established in the bay area by 1984 (CAS, EME; Powell &

Passoa in press) (Table 1).

Combining species validly listed by Dowell and Gill and those for which data

are given here, yields a total of at least 27 species of exotic Microlepidoptera and

Pyraloidea that have been discovered in California during the half century after

1940. The residency status of several of those included by Dowell and Gill is

unknown; populations have been subject to eradication procedures, and/or we

lack subsequent collections to confirm colonization and spread (e.g., Homadaula

anisocentra (Meyrick) and Endothenia albolineana (Kearfott)).

Acknowledgment

I thank all the collectors who have made the special efforts needed to acquire

study specimens of Microlepidoptera that have enabled us to document the fauna

in California during the past 35 years. Particular acknowledgment is due those

who sampled in urban areas, including J. P. Donahue (Los Angeles), R. L. Langston

(Kensington), R. H. Leuschner (Gardena, Manhattan Beach), C. D. MacNeill

(Richmond, El Cerrito), R. A. Mackie (San Diego Co.), W. W. Middlekauff (El

Cerrito), D. C. Rentz (San Francisco, Albany), and P. A. Rude (Oakland). Rose¬

mary Leen provided records of Uresiphita from her search of 1980-1987 archive

files at CDFA, gave me information on the hostplant nomenclature of U. reversalis,

and offered critical comments on the ms. Crocidosema data at San Diego Natural

History Museum and the U.S. National Museum of Natural History were com¬

piled by J. W. Brown. Records from computer and card files at the California

Dept, of Food & Agriculture were provided by T. D. Eichlin and R. Somerby;

and K. W. Brown and N. J. Smith provided records from the San Joaquin and

Fresno County Agricultural Commissioner’s Offices, respectively. Cooperation by

the following enabled me to use the collections in their care: V. Lee and N. D.

Penny (California Academy of Sciences), J. P. Donahue (Los Angeles County

Museum of Natural History), D. K. Faulkner (San Diego Natural History Mu¬

seum), R. Stecker (San Jose State University, Department of Entomology), the

late R. O. Schuster (University of California, Davis, Bohart Entomological Mu¬

seum), and D. R. Davis and R. W. Hodges (U.S. National Museum of Natural

History, Washington, D.C.).

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Bemays, E. A. & C. B. Montllor. 1989. Aposematism of Uresiphita reversalis larvae (Lepidoptera).

J. Lepid. Soc., 43: 261-273.

Clarke, J. F. G. 1971. The Lepidoptera of Rapa Island. Smithson. Contr. Zool., 56.

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Davis, D. R. 1978. The North American moths of the genera Phaeoses, Opogona, and Oinophila,

with a discussion of their suprageneric affinities (Lepidoptera: Tineidae). Smithson. Contr. Zool.,

282.

Dowell, R. V. & R. Gill. 1989. Exotic invertebrates and their effects on California. Pan-Pacif.

Entomol., 65: 132-145.

Dyar, H. G. 1901. Notes on the winter Lepidoptera of Lake Worth, Florida. Proc. Entomol. Soc.

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Emmet, A. M. 1979. A field guide to the smaller British Lepidoptera. Brit. Entomol. & Nat. Hist.

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Essig, E. O. 1941. Itinerary of Lord Walsingham in California and Oregon, 1871-72. Pan-Pacif.

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Hall, R. C. 1965. Sagebrush defoliator outbreak in northern California. U.S.D.A. Forest Serv.

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Heinrich, C. 1923. Revision of the North American moths of the subfamily Eucosminae of the

family Olethreutidae. U.S. Natl. Mus. Bull., 123.

Heinrich, C. 1956. American moths of the subfamily Phycitinae. U.S. Natl. Mus. Bull., 207.

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Hodges, R. W. 1983. Checklist of the Lepidoptera of America north of Mexico. Classey Ltd. and

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Koehler, C. & M. J. Tauber. 1964. Periploca nigra, a major cause of dieback of ornamental juniper

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Received 1 July 1991; accepted 18 August 1991.

PAN-PACIFIC ENTOMOLOGIST

68(2): 122-132, (1992)

REVISION OF THE GENUS TACHYCOLPURA BREDDIN

(HEMIPTERA: HETEROPTERA: COREIDAE: COLPURINI)

Harry Brailovsky, Ernesto Barrera, and William Lopez-Forment

Departamento de Zoologia, Instituto de Biologia,

Universidad Nacional Autonoma de Mexico,

Apdo. Postal 70153, Mexico 04510 D.F., Mexico

Abstract. —The genus Tachycolpura Breddin (Coreidae: Colpurini) is revised to include T. luteola

NEW SPECIES, from Borneo, and T. sumatrana NEW SPECIES, from Sumatra. Xenocolpura

Blote NEW SYNONYM, is synonymized within Tachycolpura with the binomial T. elongata

(Blote) NEW COMBINATION. The dorsal habitus, pronotum, and female genital plate of each

species, and the male genital capsule and parameres of the new species, are illustrated. A key to

species is provided.

Key Words.— Insecta, Heteroptera, Coreidae, Colpurini, Tachycolpura, NEW SPECIES, Sumatra,

Borneo.

The tribe Colpurini contains about 16 genera ( Hygia with nine subgenera) and

134 species, with several genera and many species awaiting description. Members

of the tribe are distributed from Fiji and Australia to India and the eastern

Palaearctic region, reaching their greatest diversity in Malaysia, Indonesia and

Papua New Guinea (Dolling 1987). The species are usually black or dark colored,

with a striking diversity of structure in the male genital capsule and in the female

genital plate (Brailovsky 1990).

Breddin (1900) described the genus Tachycolpura to include Lybas penicillatus

Walker, 1871 as the type. Distant (1901) and Bergroth (1913) cited this species

only superficially, without adding new morphological or distributional data. Blote

(1936) described and illustrated the new genus and species Xenocolpura elongata

Blote, from Sumatra. Within his generic treatment, Blote does not allude to the

affinities that this genus might have with other Colpurini, but only emphasizes,

as diagnostic characters, the reduced wings and the conical projections of the

humeral angles of the pronotum.

During this revision, we had no doubt in recognizing the close relationship

between both genera. In this paper we synonymize Xenocolpura with Tachycol¬

pura, and create a new binomial, Tachycolpura elongata. Two new species, col¬

lected in Sumatra and Borneo, are also described.

Tachycolpura is the only genus of Colpurini in which the humeral angles of the

pronotum are projected as a conical tooth of variable length, width and trajectory.

The tylus, jugae, and the antenniferous tubercles are unarmed and the femora are

armed with a double row of spines and granules that decorate their ventral side.

The shape of the posterior edge of the genital capsule, the length and width

of the gonocoxae I and of paratergite IX, the development of the wings, and the

color of the hemelytral membrane, the corium and the tibiae, all characterize the

genus.

The following abbreviations identify the institutions where types are deposited,

and specimens were loaned: Bernice P. Bishop Museum, Honolulu, Hawaii (BPBM);

The Natural History Museum, London (BMNH); Coleccion Entomologica del

1992

BRAILOVSKY ET AL.: REVISION OF TACHYCOLPURA

123

Institute) de Biologia, Universidad Nacional Autonoma de Mexico (IBUNAM);

Museum d’Histoire Naturelle, Geneva, Switzerland (MGHN); Rijksmuseum van

Naturlijke Histoire, Leiden, Netherlands (RNHL); Zoologisches Musem, Univer-

siteit Van Amsterdam, Netherlands (ZMUA).

Tachycolpura Breddin

Tachycolpura Breddin, 1900. Rev. d’Entomol. 19: 215.

Tachycolpura : Bergroth, 1913. Mem. Soc. Entomol. Belg. 22: 142.

Xenocolpura Blote, 1936. Zool. Meded. 19: 44, NEW SYNONYM.

Type Species. —Lybas penicillatus Walker.

Redescription. —Narrow body, moderately elongated, with an average length from 16.48 mm to

20.15 mm. Head. Longer than wide, elongate, cylindrical and slightly narrowed basally; tylus unarmed,

apically truncate, extending anterior to jugae, and seen laterally extending above them; antenniferous

tubercles unarmed with truncate apex; jugae unarmed; antennal segment I robust, cylindrical, slightly

curved outwards and longer than head; segment II longest, segment IV shortest and fusiform; segments

II and III cylindrical; ocelli not elevated; preocellar pit deep, diagonally excavated; eyes spherical;

tubercles postocular protuberant; side of head in front of eyes straight, slightly convergent; bucculae

rounded, short, not projecting beyond antenniferous tubercle, with sharp mesial projection and anterior

edges thickened; rostrum long, reaching the medial one-third of abdominal stemite V, or almost to

apex of VII; rostral segment IV longest. III longer than II and II longer than I, which is shortest.

Thorax. Pronotum. Wider than long, moderately sloped; anterior collar wide; anterolateral edges

ranging from oblique and gently rounded to almost straight; humeral angles projected into conical

tooth, directed upwards and slightly backwards, with variable length (Figs. 1-5); posterior edge straight.

Anterior lobe of metathoracic scent gland globose and reniform, posterior lobe sharp, small. Legs.

Femora with two rows of granules and small spines along ventral surface, less abundant on metafemur;

tibiae with shallow sulcus, sometimes difficult to see; metatibiae longer than metafemur. Scutellum.

Triangular, longer than wide, with sharp apex. Hemelytra. Macropterous, reaching median one-third

of abdominal segment VII of male or median one-third of VIII, or anterior one-third of IX in female,

or coleopteroid and extending to anterior third of abdominal segment V in both sexes (see Slater

1975); claval suture evident or barely so (coleopteroid individuals); claval commissure shorter than

total length of scutellum; apical border obliquely straight, with short apical angle not reaching middle

one-third of hemelytral membrane; hemelytral membrane with few bifurcate veins. Abdomen. Con-

nexival segments higher than body, forming a case where hemelytra rest; posterior angle of connexival

complete, or extended into a very short, wide projection; abdominal stemites with medial sternal

furrow projecting to posterior border of stemites V or VI. Integument. Body surface rather dull. Head,

pronotum, scutellum, clavus, corium, thorax, abdominal sterna and exposed parts of genital segments

of both sexes strongly punctate. Antennae and legs minutely granulate. Head, pronotum, scutellum,

clavus, corium, thorax and abdominal sterna with long, decumbent to suberect conspicuous golden

or silvery bristle-like hairs. Pronotum, thorax and abdominal sterna with circular gray-white farinose

punctures.

Male Genitalia. —Genital Capsule. Posteroventral edge bidentate (Figs. 14-16). Parameres. Simple

and straight body; apical projection widened, with the anterior lobe convex or continuous with body

and the posterior lobe ending in a sharp and short projection (Figs. 20-24).

Female Genitalia. — Abdominal stemite VII with plica and fissure evident; plica narrow or elevated

and transversely evolved; gonocoxae I nearly square, large; paratergite VIII short, square, with spiracle

visible; paratergite IX nearly square, larger than former paratergite VIII (Figs. 6-13). Spermatheca.

Bulb long and dilated, duct coiled, with short membranous duct (Fig. 25).

Diagnosis. — Tachycolpura is the only genus within the Colpurini that has the

humeral angles of the pronotum projected into a sharp and robust conical pro¬

jection, of variable length and trajectory. Other typical characters are an unarmed

tylus, jugae and antenniferous tubercles, an armed femora of all three pairs of

124

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Figures 1-5. Pronotum view of Tachycolpura spp. Figures 1, 2. T. penicillata (Walker). Figure 1.

Male. Figure 2. Female. Figure 3. T. elongata (Blote) NEW COMBINATION. Figure 4. T. luteola,

NEW SPECIES. Figure 5. T. sumatrana NEW SPECIES.

1992

BRAILOVSKY ET AL.: REVISION OF TACHYCOLPURA

125

Figures 6-9. Frontal view of the female genital plates of Tachycolpura spp. Figure 6. T. penicillata

(Walker). Figure 7. T. elongata (Blote) NEW COMBINATION. Figure 8. T. luteola NEW SPECIES.

Figure 9. T. sumatrana NEW SPECIES. Figures 10-13. Lateral view of female genital plates of

Tachycolpura spp. Figure 10. T. penicillata (Walker). Figure 11. T. elongata Blote NEW COMBI¬

NATION. Figure 12. T. luteola NEW SPECIES. Figure 13. T. sumatrana NEW SPECIES.

legs, and a notoriously elongated head. The presence of a fissure and a plica in

the female, together with the spiny projection of the buccula, confirm the generic

diagnosis of the genus.

Discussion .—Wing development in the Colpurini is notoriously variable, in-

126

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

eluding apterous, coleopteroid, micropterous, submacropterous and macropterous

species, even within a genus ( Sciophyrus ) and a species ( Brachylybas spp.). There¬

fore, wing character is not a reliable tool for a generic definition.

Blote (1936) in describing and illustrating Xenocolpura, noted that its charac¬

teristic features are especially a brachypterous condition, the presence of a sub-

conical tooth in the humeral angles of the pronotum and a thorny projection in

the bucula. In examining the type material of X. elongata Blote and Tachycolpura

penicillata (Walker), both monotypic genera, we could not find any definitive

characters to be used. Both species have the same degree of development of the

humeral angles, the bucula and of the genital plates of the female. Therefore, we

synonymized Xenocolpura within Tachycolpura, and included X. elongata as the

second known species of Tachycolpura.

Distribution. = Four species are known from Malaya, Sumatra, Singapore and

Borneo.

Biology. —Apparently a very scarce genus restricted to forested areas.

Key to Tachycolpura Species

1. Coleopteroid individuals, with the hemelytral membrane not extending

beyond abdominal segment V; claval suture not evident; gonocoxae

I long, with a maximum length of 3.00 mm (Figs. 7, 11); posterior

border of genital capsule with two short projections, with robust and

truncated apices (Figs. 16, 19) (Sumatra) .

. T. elongata (Blote) NEW COMBINATION

1Macropterous individuals, with the hemelytral membrane reaching ab¬

dominal segment VII of male, or IX in female; claval suture evident;

gonocoxae I shorter than 2.90 mm. 2

2(1'). Apical angle and apical margin of corium yellow; genital capsule elon¬

gate, with posterior margin oblique and convergent, with two short

projections with rounded apices (Figs. 15, 18) (Borneo) .

. T. luteola NEW SPECIES

2'. Apical angle and apical border of corium black or brown-red; genital

capsule globose, with the posterior edge widened, with two short

rounded lobes (Figs. 14, 17). 3

3(2'). Humeral angle of pronotum with long, thin, slender conical projections

(Fig. 5); clavus and corium pallid red-orange; tibiae dark orange, with

two yellow rings, one subbasal and the other almost apical (Sumatra)

. T. sumatrana NEW SPECIES

3'. Humeral angles of pronotum with short and robust projections (Figs.

1-2); clavus and corium black; tibiae dark red-brown, without yellow

rings (Singapore, Borneo) . T. penicillata (Walker)

Tachycolpura penicillata (Dallas)

(Figs. 1, 2, 6, 10, 14, 17, 20, 21, 26)

Lybas penicillatus Walker, 1871. Cat. Hem. IV: 150-151.

Lybas penicillatus'. Lethierry & Severin, 1894. Cat. Gen. 2: 42.

Tachycolpura penicillata'. Breddin, 1900. Rev. d’Entomol. 19: 216.

Colpura penicillatus : Distant, 1901. Ann. Mag. Nat. Hist. Ser. 7(7): 20.

Tachycolpura penicillata'. Bergroth, 1913. Mem. Soc. Entomol. Belg. 22: 142.

Figures 14-16. Frontal view of the male genital capsule of Tachycolpura spp. Figure 14. T. pen-

icillata (Walker). Figure 15. T. luteola NEW SPECIES. Figure 16. T. elongata (Blote) NEW COM¬

BINATION. Figures 17-19. Lateral view of the male genital capsule of Tachycolpura spp. Figure 17.

T. penicillata (Walker). Figure 18. T. luteola NEW SPECIES. Figure 19. T. elongata (Blote) NEW

COMBINATION. Figures 20-24. Parameres of Tachycolpura spp. Figures 20, 21. T. penicillata

(Walker) Figures 22, 23. T. luteola NEW SPECIES. Figure 24. T. elongata (Blote) NEW COMBI¬

NATION. Figure 25. Spermatheca of Tachycolpura luteola NEW SPECIES.

128

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Types. —Lybas penicillatus Walker. We designate a female, collected in Sin¬

gapore and deposited in the Natural History Museum, London, as a Lectotype.

Redescription. —Female. Color. Black with the following areas pale ochre or pale orange: upper side

of the postocular tubercles, apex of scutellum, a very small discoidal dot on middle one-third of apical

margin of corium, posterior one-third of connexivum, anterior and posterior lobes of metathoracic

scent gland, and posterior angle or pleural margin of abdominal stemites III to VII; antennal segments

II and III, rostral segments I to IV and tibiae and tarsi dark red-brown; antennal segment I black, and

IV dark ochre, with basal one-third red; hemelytral membrane dirty yellow with veins red-brown,

basal angle and anterior margin pale yellow. Structures. Rostrum reaching posterior border of sternal

segment V; humeral angles of pronotum projecting into a conical, short, robust tooth, pointed backward

(Fig. 2); hemelytra macropterous, with claval suture evident and membrane reaching middle one-

third of abdominal segment VIII; posterior angle of connexival segments V and VI not projecting out

from surface; gonocoxae I conspicuously long, with the maximum width large; paratergite IX nearly

square, short and barely reaching beyond the external border of gonocoxae I (Figs. 6, 10). Measure¬

ments: Head length: 2.85 mm; interocellar space: 0.64 mm; interocular space: 1.44 mm; width across

eyes: 2.15 mm; preocular distance: 1.85 mm; length antennal segments: I, 4.00 mm; II, 5.20 mm; III,

3.70 mm; IV, 2.25 mm. Pronotal length: 3.70 mm; width across frontal angles: 1.70 mm; width across

humeral angles: 4.90 mm. Scutellar length: 2.35 mm; width: 2.00 mm. Maximum length of gonocoxae

I seen frontally: 2.85 mm; maximum length of gonocoxae I seen laterally: 1.35 mm. Total body length:

17.65 mm.

Male. — Color. Similar to female, but hemelytral membrane dirty yellow with veins and anterior

margin brown and only basal angle yellow. Structures. Humeral angles produced into a short conical

tooth, barely projecting beyond posterolateral edge of pronotum (Fig. 1). Macropterous hemelytra and

membrane reaching middle one-third of abdominal segment VII. Genital capsule globose with posterior

margin widened and with two short rounded mounds (Figs. 14, 17). Parameres. Figs. 20-21. Mea¬

surements: Head length: 2.84 mm; interocellar space: 0.64 mm; interocular space: 1.25 mm; width

across eyes: 2.13 mm; preocular distance: 1.68 mm; length antennal segments: I, 4.00 mm; II, 5.16

mm; III, 3.70 mm; IV, 2.23 mm. Pronotal length: 3.30 mm; width across frontal angles: 1.70 mm;

width across humeral angles: 3.92 mm. Scutellar length: 2.20 mm; width: 1.65 mm. Total body length:

16.48 mm.

Diagnosis. — Macropterous species, characterized by having the humeral angles

of the pronotum projected into a short, conical robust tooth (Fig. 1), or very small

(Fig. 2), and in each condition pointed backward, with the middle one-third of

the apical margin of corium with a discoidal small yellow patch. The male genital

capsule is globose, with the posterior margin widened and apices produced into

two rounded mounds (Figs. 14, 17). Paratergite IX of female square, short, and

barely surpasses the external border of gonocoxae I (Figs. 6, 10). The basal angle

of the hemelytral membrane yellow.

Distribution. —Originally described from Singapore and northern Borneo (Sa¬

rawak).

Material Examined.— One male and three females, among them the female lectotype. Data: MA¬

LAYA. Ulu Gombok. INDONESIA. BORNEO: Without localities.

Tachycolpura elongata (Blote) NEW COMBINATION

(Figs. 3, 7, 11, 16, 19, 24, 29)

Xenocolpura elongata Blote, 1936. Zool. Meded. 19: 44-45.

Types.— Female holotype deposited in the Rijksmuseum van Naturlijke His-

toire, Leiden, Netherlands.

Redescription.—Female. Color. Black, with following areas orange ochre: dorsum of postocular

tubercles, apex of scutellum, posterior margin of connexivum, and anterior and posterior lobes of

1992

BRAILOVSKY ET AL.: REVISION OF TACHYCOLPURA

129

Figures 26-29. Dorsal view Tachycolpura spp. Figure 26. T. penicillata (Walker). Figure 27. T.

luteola NEW SPECIES. Figure 28. T. sumatrana NEW SPECIES. Figure 29. T. elongata (Blote) NEW

COMBINATION.

130

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

metathoracic scent glands; rostral segments I to IY, trochanters, most of tibiae and tarsi red-brown;

hemelytral membrane dirty yellow, with veins and basal angle red-brown. Structures. Rostrum reaching

posterior border of sternal segment V; humeral angles of pronotum produced into a robust, short,

conical tooth, projected backward (Fig. 3); hemelytra coleopteroid, with claval suture not evident, and

membrane reaching anterior one-third of abdominal segment Y; posterior angle of connexival segments

V-VI well marked against surface; gonocoxae I conspicuously elongated, with well developed maxi¬

mum width; paratergite IX square, conspicuously surpassing external border of gonocoxae I (Figs. 7,

11). Measurements: Head length: 3.06 mm; interocellar space: 0.76 mm; interocular space: 1.40 mm;

width across eyes: 2.35 mm; preocular distance: 2.12 mm; length antennal segments: I, 3.90 mm; II

to IV absent. Pronotal length: 3.48 mm; width across frontal angles: 1.74 mm; width across humeral

angles: 4.55 mm. Scutellar length: 1.95 mm; width: 1.85 mm. Maximum length of gonocoxae I seen

frontally: 3.00 mm; maximum length of gonocoxae I seen laterally: 1.80 mm. Total body length:

18.25 mm.

Male. — Color. Similar to female. Structures. Rostrum reaching anterior margin of sternal segment

VII; coleopteroid, with hemelytral membrane reaching anterior one-third of abdominal segment V.

Genital capsule globose, posterior margin widened, with two short robust lateral projections with

truncate apices (Figs. 16, 19). Parameres. Fig. 24. Measurements: Head length: 3.00 mm; interocellar

space: 0.67 mm; interocular space: 1.38 mm; width across eyes: 2.30 mm; preocular distance: 2.00

mm; length antennal segments: I, 3.83 mm; II to IV absent. Pronotal length: 3.09 mm; width across

frontal angles: 1.69 mm; width across humeral angles: 3.90 mm. Scutellar length: 1.80 mm; width:

1.55 mm. Total body length: 17.18 mm.

Diagnosis. — This is the only species in the genus with coleopteroid hemelytra;

the claval suture is not evident and the membrane is very short, not extending

beyond the anterior one-third of abdominal segment V. The aspect of the humeral

angles of the pronotum, as well as the length of the gonocoxae I, place it near T.

penicilliata (Walker), but in T. elongata (Blote) the gonocoxae I is clearly wider

and paratergite IX extends well beyond the external border of gonocoxae I (Figs.

6,7,10,11).

The genital capsule of T. elongata is wide and posseses two robust projections

with truncated apices (Figs. 16, 19), whereas the other species have two very short

mounds with rounded apices (Figs. 14, 17).

Distribution. — Restricted to Sumatra, from Lubu Raja and Tapanuli.

Material Examined.—One, male and three females, among which was the holotype. INDONESIA.

( WEST) SUMATRA: PADANG: Pandjang.

Tachycolpura luteola Brailovsky, Barrera & Lopez-Forment NEW SPECIES

(Figs. 4, 8, 12, 15, 18, 22, 23, 25, 27)

Types. — Holotype: male; data: INDONESIA. ( CENTRAL) BORNEO: Sg. Pa-

jau, 1925, Mjoberg. Deposited in the Zoologisches Museum, Universiteit Van

Amsterdam, Netherlands. Paratypes: 3 males, 5 females; same data as holotype.

(2 males and 4 females deposited in the Zoologisches Museum, Universiteit Van

Amsterdam, Netherlands and 1 male and 1 female in the “Coleccion Entomologica

del Instituto de Biologia, UNAM, Mexico”); INDONESIA. ( NORTHWEST)

BORNEO: Kuching, Jan 1900, Dyak, 4 females (3 deposited in the Rijksmuseum

van Naturlijke Histoire, Leiden, Netherlands and 1 in the “Coleccion Entomo¬

logica del Instituto de Biologia, UNAM, Mexico”).

Description. —Male (holotype). Color. Black, with the following areas ochre or yellow ochre, some¬

times with orange reflections: apex of scutellum, apical angle and apical margin of corium, posterior

margin of connexivum, internal side of trochanters, anterior and posterior lobes of metathoracic scent

glands, and angle or posterior margin of pleural margin of abdominal stemites IV to VII; antennal

1992

BRAILOVSKY ET AL.: REVISION OF TACHYCOLPURA

131

segments II, III and tibiae dark red-brown; rostral segments I to IV and tarsi lighter red-brown;

antennal segment I black, IV yellow with basal one-third brown; external side of trochanters shiny

red-brown; hemelytral membrane dirty yellow with veins and subbasal large brown blotch and pallid

yellow basal angle. Structures. Rostrum reaching anterior border of sternal segment VI; pronotal

humeral angles projecting into a short, robust, conical tooth pointed outwards and slightly downwards

(Fig. 4); macropterous hemelytra, claval suture evident, membrane reaching middle one-third of

abdominal segment VII; posterior angle of connexival segments V and VI not marked on surface.

Genital capsule long, posterior margin becoming narrower with conspicuous oblique border, and two

short lateral projections with rounded apices (Figs. 15, 18). Parameres. Figs. 22, 23. Measurements:

Head length: 3.00 mm; interocellar space: 0.72 mm; interocular space: 1.26 mm; width across eyes:

2.15 mm; preocular distance: 1.95 mm; length antennal segments: I, 4.75 mm; II, 6.70 mm; III, 4.65

mm; IV, 2.70 mm. Pronotal length: 3.45 mm; width across frontal angles: 1.62 mm; width across

humeral angles: 4.10 mm. Scutellar length: 2.25 mm; width: 1.90 mm. Total body length: 17.80 mm.

Female. — Color. Similar to male. Structures. Macropterous, with hemelytral membrane reaching

posterior margin of abdominal segment IX. Gonocoxae I short lengthwise with well developed width;

paratergite IX square, reaching beyond external margin of gonocoxae I (Figs. 8, 12). Spermatheca.

Fig. 25. Measurements: Head length: 2.85 mm; interocellar space: 0.70 mm; interocular space: 1.17

mm; width across eyes: 2.10 mm; preocular distance: 1.90 mm; length antennal segments: I, 4.15

mm; II, 5.70 mm; III, 4.05 mm; IV, 2.40 mm. Pronotal length: 3.60 mm; width across frontal angles:

1.65 mm; width across humeral angles: 4.85 mm. Scutellar length: 2.20 mm; width: 1.95 mm. Max¬

imum length of gonocoxae I, seen frontally: 2.25 mm; maximum length of gonocoxae I, seen laterally:

1.55 mm. Total body length: 17.90 mm.

Diagnosis. —This distinctive species is recognized by the light yellow color of

the apical angle and apical margin of the corium. In T. penicillata and T. elongata,

the corium are entirely black. The length of the gonocoxae I is very short (2.25

mm) and the posterior margin of the genital capsule is narrowed apically, with

conspicuously oblique margins and two short rounded apical projections (Figs.

15, 18). In the other species, the gonocoxae I is longer (2.80-3.00 mm), and the

genital capsule is wider and globose, with both projections truncated apically

(Figs. 16, 19), or rounded (Figs. 14, 17).

Etymology. — The taxon name is based on the yellow color of the apical angle

and the apical margin of the corium.

Material Examined.—See types.

Tachycolpura sumatrana, Brailovsky, Barrera & Lopez-Forment

NEW SPECIES

Type.— Holotype: female; data: INDONESIA. SUMATRA: Deli (Bed Piet),

without date. Deposited in the Museum d’Histoire Naturelle, Geneva, Switzer¬

land. The left wing of the holotype is destroyed.

Description. —Female (holotype). Color. Head, pronotum, scutellum, thorax and abdominal stemites

black with pale red reflections at apex of tylus, scutellar disc, acetabula of three pairs of legs, abdominal

stemites, genital plates and pleural margin of abdominal stemites III to VII; ochre yellow on: upper

side of postocular tubercles, apex of scutellum, semi-discoidal spot on middle one-third of apical

margin of corium, posterior margin of connexival, anterior and posterior lobes of metathoracic scent

gland, and posterior margin of pleural margin of abdominal stemites III to VII; antennal segment I

dark red-brown, segments II and III pale red-orange, IV yellow with basal one-third pale orange;

clavus, corium, connexivum and dorsal segments of abdomen red-orange; hemelytral membrane dirty

yellow with veins and large subbasal brown spot and basal angle dark ochre; coxae and femora red-

brown; trochanters bicolored, with external side red-brown, and internal side yellow; tibiae dark orange

with two yellow rings, one subbasal, other almost apical; orange tarsi with ochre reflections; rostral

segments I to IV brown ochre. Structures. Rostrum reaching posterior border of sternal segment V;

humeral angles of pronotum produced into long, thin, slightly backwards inflected conical prominence

132

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

(Fig. 5); macropterous hemelytra with claval suture evident and membrane reaching middle one-third

of abdominal segment IX; posterior angle of connexival segments V and VI slightly remarked on

surface; gonocoxae I well developed longitudinally and transversely widened; paratergite IX square,

length exceeding posterior margin of gonocoxae I (Figs. 9, 13). Measurements: Head length: 3.05 mm;

interocellar space: 0.65 mm; interocular space: 1.30 mm; width across eyes: 2.30 mm; preocular

distance: 1.90 mm; length antennal segments: I, 4.10 mm; II, 5.65 mm; III, 3.80 mm; IV, 2.25 mm.

Pronotal length: 3.60 mm; width across frontal angles: 1.80 mm; width across humeral angles: 5.85

mm. Scutellar length: 2.45 mm; width: 2.15 mm. Maximum length of gonocoxae I, seen frontally:

3.00 mm; maximum width of gonocoxae I, seen laterally: 1.60 mm. Total body length: 17.30 mm.

Male. — Unknown.

Diagnosis. — The peculiar long and slender (Fig. 5) projections of the humeral

angles of the pronotum, the pale red-orange coloration of the clavus, corium,

connexivum and the abdominal segments, and the two yellow rings on the tibiae,

are diagnostic characters of T. sumatrana. All the other species have shorter and

more robust conical projections of the humeral angles; their clavus, corium, con¬

nexivum and abdominal segments are black, and their tibiae lack two yellow rings.

Etymology. —Named for its occurrence on the Island of Sumatra.

Material Examined.—Set types.

Acknowledgment

We are indebted to the following individuals and institutions for the loan of

specimens and other assistance: Gordon M. Nishida (Bernice P. Bishop Museum,

Honolulu); J. Palmer and Janet Margerison Knight (British Museum [Natural

History], London); Jan Van Tol (Rijksmuseum van Naturlijke Histoire, Leiden);

W. Hogenes (Zoologisches Museum, Universiteit Van Amsterdam); B. Hauser

(Museum d’Histoire Naturelle, Geneva).

Special thanks are extended to the Direction General de Asuntos del Personal

Academico of the Universidad Nacional Autonoma de Mexico (DGAPA) and to

the Consejo Nacional de Ciencia y Tecnologia (CONACyT) for financial assistance

to the senior author.

We thank the reviewers for reading and comments.

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Received 1 July 1991; accepted 18 October 1991.

PAN-PACIFIC ENTOMOLOGIST

68(2): 133-139, (1992)

DESCRIPTIONS OF IMMATURES OF EOEURYSA

FLAVOCAPITATA MUIR FROM TAIWAN

(HOMOPTERA: DELPHACIDAE)

Stephen W. Wilson , 1 James H. Tsai , 2 and C. C. Chen 3

department of Biology, Central Missouri State University,

Warrensburg, Missouri 64093

2 Fort Lauderdale Research and Education Center, University of Florida,

IFAS, Fort Lauderdale, Florida 33314

3 Taichung District Agricultural Improvement Station, Changhua, Taiwan

Abstract. — Adult male and female genitalia, the egg, and first through fifth instar nymphs of the

delphacid planthopper Eoeurysa flavocapitata Muir, collected from sugarcane ( Saccharum offi-

cinarum L.) from Taiwan are described and illustrated and a key to instars is provided. Features

useful in separating nymphal instars include differences in body size and proportions; spination

of metatibiae, metatibial spurs, and metatarsomeres; and number of metatarsomeres.

Key Words.— Insecta, Homoptera, Delphacidae, Eoeurysa flavocapitata, immature stages, Tai¬

wan, sugarcane

The delphacid planthopper Eoeurysa flavocapitata Muir has been recorded from

northeastern India, Bangladesh, Malaysia, China, Indonesia, and Taiwan (Chat-

terjee 1971, Chatterjee & Choudhuri 1979, Chu & Chiang 1975, Metcalf 1943,

Mirza & Qadri 1964, Qadri 1963). Adult females insert eggs in, and adults and

nymphs feed on, leaves of sugarcane {Saccharum ojflcinarum L.). Feeding causes

leaf desiccation, development of red streaks on damaged tissue, and growth of

sooty mold on the honey dew produced by the planthopper (Chatterjee & Choud¬

huri 1979, Fennah 1969). Sugarcane is the only recorded host, and E. flavocapitata

may be monophagous on this grass as are over 20 species of sugarcane delphacids

in the genus Perkinsiella (Wilson 1988). However, the Neotropical sugarcane pest

Saccharosydne saccharivora (Westwood) was found to have two species of An-

dropogon as its natural hosts (Metcalfe 1969) and apparently included the related

grass sugarcane (all in the tribe Andropogoneae [Clayton and Renvoize 1986]) in

its range of food plants after introduction of sugarcane to the New World. The

only other Eoeurysa, E. arundina Kuoh and Ding, has been found only on Arundo

donax L. (Yang 1989), another economically important grass, which is not closely

related to sugarcane (Clayton & Renvoize 1986).

The biology of E. flavocapitata on sugarcane was studied by Chatterjee & Choud¬

huri (1979) and Jiang (1976) who provided information on oviposition, feeding

sites, and duration of stadia. Adults were described and illustrated by Chu &

Chiang (1975), Jiang (1976) and Yang (1989). Brief descriptions of immatures

were provided by Chatterjee & Choudhuri (1979), who also included somewhat

diagrammatic illustrations, and Jiang (1976 [in Chinese]); the fifth instar was

described, and partial illustrations provided, by Wu & Yang (1985). The present

paper includes detailed descriptions and illustrations of adult male and female

genitalia, and the immature stages; it also gives a key for the separation of nymphal

instars.

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THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Figures 1-4. Eoeurysa flavocapitata adult genitalia. Figure 1. Male, left lateral view of complete

genitalia. Figure 2. Male, right lateral view of aedeagus. Figure 3. Male, caudal view of pygofer and

styles. Figure 4. Female, lateral view of complete genitalia. Scale bar = 0.5 mm.

Methods

Terminology used in the description of the female genitalia follows Asche (1985)

and Heady & Wilson (1990). The fifth instar is described in detail but only major

differences are described for fourth through first instars. Arrangement and number

of pits is provided for the fifth and fourth instars; this information is not given

for earlier instars because the pits are extremely difficult to discern (those that

could be observed relatively easily are illustrated). Measurements are given as

mean ± SD. Length was measured from apex of vertex to apex of abdomen,

width across the widest part of the body, and thoracic length along the midline

from the anterior margin of the pronotum to the posterior margin of the metano-

tum. Eggs were obtained by excising them with a fine needle from sections of field

collected sugarcane leaves.

1992

WILSON ET AL.: IMMATURE EOEURYSA FLAVOCAPITATA

135

Figure 5. Eoeurysa flavocapitata female, ventral view of complete genitalia. Scale bar = 0.5 mm.

Eoeurysa flavocapitata Muir

Descriptions.—Adults (Figs. 1-5). Adult E. flavocapitata were briefly described by Muir (1913);

detailed descriptions and illustrations provided by Jiang (1976) and Yang (1989) should be referred

to for non-genitalic adult morphology.

Male genitalia (Figs. 1-3). —Pygofer, in lateral view, subquadrate, with broadly produced diaphragm

armature. Anal tube, in lateral view, with a single spinose process originating at the dorsocaudal aspect

of the tube, and a pair of bifid spinose processes each originating at the ventrocaudal aspect of the

tube. Styles, in caudal view, broadest across basal one-third, narrowing and strongly divergent in

apical one-third. Aedeagus subcylindrical, with a dorsally directed terminal bifid tooth and a dorso-

caudally directed process in apical one-third on right side.

Female genitalia (Figs. 4, 5).—Tergite nine oriented anteroventrally (see Asche 1985), elongate,

longitudinally concave in ventral midline. Anal tube subcylindrical, style somewhat bulbous. Genital

scale (or atrium plate) subtriangular. Valvifers of segment eight each covering approximately one-

third of tergite nine anterolaterally; medial margin deeply notched in anterior one-third. Lateral

gonapophyses of segment nine elongate, broadly rounded posteriorly. In lateral view, median gon-

apophyses of segment nine saber-shaped, with approximately 15 strong teeth on dorsal margin in

distal one-half (not all teeth apparent in ventral view). Gonapophyses of segment eight slender, subacute

apically.

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

Figures 6-8. Eoeurysa flavocapitata fifth instar. Figure 6. Habitus, dorsal view. Figure 7. Ventral

view of male. Figure 8. Apical part of venter of female abdomen. Scale bar = 0.5 mm.

Fifth instar nymph (Figs. 6-8).—Length 3.6 ±0.17 mm; thoracic length 1.1 ± 0.06 mm; width 1.2

± 0.08 mm (n = 10). Body white with gray to fuscous markings on frons, clypeus, and apex of

abdomen. Form elongate, subcylindrical, flattened dorsoventrally, widest across mesothoracic wing-

pads. Vertex subtriangular; posterior margin nearly straight, narrowing anteriorly. Frons border with

clypeus concave; lateral margins strongly convex and carinate (outer carinae) and paralleled by second

pair of very weak carinae (inner carinae) continuous with lateral margins of vertex; area between inner

and outer carinae with nine pits on each side (six visible in ventral aspect, three in dorsal aspect);

three pits between each outer carina and eye. Clypeus subconical, narrowing distally. Beak three-

segmented, cylindrical, segment one hidden by anteclypeus, segment two subequal in length to segment

three, segment three with black apex. Antennae three-segmented; scape short, cylindrical; pedicel

subcylindrical, 2.0 x length of scape; flagellum bulbous basally, with elongate bristle-like extension

distally, bulbous base approximately 0.3 x length of pedicel. Thoracic nota divided by middorsal line

into three pairs of plates. Pronotal plates subtriangular (in dorsal view); anterior margin convex;

posterior border sinuate; each plate with a weak posterolaterally directed carina and seven pits ex¬

tending anteriorly from near middorsal line posterolaterally to lateral margin (lateralmost pits often

not visible in dorsal view). Mesonotum with median length 2.0 x that of pronotum; elongate lobate

wingpads almost extending to tips of metanotal wingpads; each plate with very weak posterolaterally

directed carina (not illustrated); two pits near middle of non-lobate portion of plate and two pits near

lateral margin. Metanotum with median length approximately 0.7 x that of mesonotum; lobate wing¬

pads extending to fourth tergite; each plate with one very weak pit near middle of plate (not illustrated).

Pro- and mesocoxae elongated and directed posteromedially; metacoxae fused to sternum. Metatro¬

chanter short and subcylindrical. Metatibia with two spines on lateral aspect of shaft, an apical

transverse row of five black-tipped spines on plantar surface and a subtriangular flattened movable

spur with one apical tooth and 13-15 other teeth on posterior margin. Pro- and mesotarsi with two

1992

WILSON ET AL.: IMMATURE EOEURYSA FLAVOCAPITATA

137

Figures 9-13. Eoeurysa jlavocapitata immature stages. Figure 9. Egg. Figure 10. First instar. Figure

11. Second instar. Figure 12. Third instar. Figure 13. Fourth instar. Scale bars = 0.5 mm (top = 9-

11, bottom =12, 13).

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

tarsomeres, tarsomere one wedge-shaped; tarsomere two subconical, with pair of apical claws and

median membranous pulvillus. Metatarsi with three tarsomeres; tarsomere one with apical transverse

row of eight black-tipped spines; tarsomere two cylindrical, approximately 3.5 x length of tarsomere

one, with apical transverse row of four black-tipped spines on plantar surface; tarsomere three sub-

conical, slightly longer than tarsomere two, with pair of apical claws and median pulvillus. Abdomen

nine segmented; flattened dorsoventrally; widest across fourth abdominal segment. Tergite one small,

sub triangular, hidden by juncture of thorax and abdomen (not visible in illustration); two subrectan-

gular, not extending to lateral aspect of segment; tergites five to eight each with three pits on each side

(lateralmost pits not always visible in dorsal view). Segment nine surrounding anus, with three pits

on each side; female with one pair of acute processes extending from juncture of stemites eight and

nine; males lacking processes.

Fourth instar nymph (Fig. 13).—Length 2.8 ± 0.18 mm; thoracic length 0.8 ± 0.04 mm; width 0.08

± 0.04 mm (n = 10). Antennal flagellum with basal portion approximately 0.5 x length of pedicel.

Mesonotal wingpads shorter, each covering approximately two-thirds of metanotal wingpad laterally.

Metanotal median length 1.5 x that of mesonotum; wingpad extending to tergite two. Metatibial spur

slightly smaller, with one apical tooth and eight teeth on margin. Metatarsi with two tarsomeres;

tarsomere one with apical transverse row of seven black-tipped spines; tarsomere two subconical with

three black-tipped spines in middle of tarsomere on plantar surface. Abdominal segments four to eight

each with the following number of pits on either side of midline: tergite four with one pit, five with

two, six to eight each with three, segment nine with three.

Third instar nymph (Fig. 12).—Length 2.0 ± 0.12 mm; thoracic length 0.6 ± 0.02 mm; width 0.6

±0.03 mm (n = 10). Mesonotal wingpads shorter, each covering one-third of metanotal wingpad

laterally. Metanotal wingpad extending to tergite one. Metatibial spur smaller; with one apical and

one or two marginal teeth. Metatarsomere one with apical transverse row of six black-tipped spines

on plantar surface.

Second instar nymph (Fig. 11).—Length 1.5 ± 0.06 mm; thoracic length 0.5 ± 0.01 mm; width 0.4

± 0.02 mm (n = 10). Mesonotal median length subequal to that of pronotum; wingpads undeveloped.

Metanotal median length subequal to that of mesonotum; wingpads undeveloped. Metatibia with

apical row of three black-tipped spines; spur small with no marginal teeth, approximately 3.0 x length

of longest metatibial spine; metatarsomere one with four apical black-tipped spines.

First instar nymph (Fig. 10).—Length 1.0 ± 0.06 mm; thoracic length 0.4 ± 0.02 mm; width 0.3

± 0.02 mm (n = 10). Bulbous base of antennal flagellum subequal in length to that of pedicel. Metatibia

lacking spines on shaft; metatibial spur smaller, approximately 1.5 x length of longest metatibial spine.

Egg (Fig. 9).—Length 0.8 ± 0.05 mm; width 0.2 ± 0.02 mm (n = 5). Eggs laid singly; white,

cylindrical, narrower at apical end; chorion translucent, smooth.

Material Examined.— Specimens used for description have the following data: REPUBLIC OF

CHINA. TAIWAN: Taichung, 5 Dec 1989, ex sugarcane, (10 males, 12 females, 5 eggs, 19 first instars,

14 second instars, 15 third instars, 25 fourth instars, 19 fifth instars).

Key to E. flavocapitata Nymphal Instars

1. Metatibial spur with more than five marginal teeth (Figs. 7, 13); meso¬

notal wingpads overlapping more than one-half length of metanotal

wingpads (Figs. 6, 13). 2

- Metatibial spur with fewer than five marginal teeth; mesonotal wingpads

overlap less than one-half length of metanotal wingpads (Figs. 10-12)

. 3

2(1). Metatarsi with three tarsomeres; metatibial spur with more than 10

marginal teeth; mesonotal wingpads extending to or almost to apex

of metanotal wingpads (Figs. 6, 7).fifth instar

Metatarsi with two tarsomeres; metatibial spur with eight marginal teeth;

mesonotal wingpads not extending to apex of metanotal wingpads

(Fig. 13) .fourth instar

3(2). Metatibia with transverse row of four apical spines, spur with one or

two marginal teeth (Fig. 12) .third instar

1992 WILSON ET AL.: IMMATURE EOEURYSA FLAVOCAPITATA 139

Metatibia with transverse row of three apical spines, spur lacking mar¬

ginal teeth (Figs. 10, 11). 4

4(3). Metatibia with lateral spine near middle on outer surface; spur approx¬

imately 3.0 x length of longest metatibial apical spine (Fig. 11) ....

.second instar

Metatibia without lateral spines; spur approximately 2.0 x or less length

of longest metatibial apical spine (Fig. 10).first instar

Acknowledgment

Florida Agricultural Experiment Station Journal Series R-01742.

Literature Cited

Asche, M. 1985. Zur Phylogenie der Delphacidae Leach, 1815 (Homoptera Cicadina Fulgoromor-

pha). Marburger Entomol. Publ., 2(1): 1-910.

Chatterjee, P. G. 1971. Occurrence of Eoeurysa flavocapitata Muir (Fam. Delphacidae) on sugarcane

in India. Indian J. Entomol., 33: 220.

Chatterjee, P. G. & D. K. Choudhuri. 1979. Biology of Eoeurysa flavocapitata—a. delphacid pest on

sugarcane in India. Entomon, 4: 263-267.

Chu, Y. I. & P. H. Chiang. 1975. A new insect pest of sugarcane in Taiwan (Eoeurysa flavocapitata

Muir). Plant Prot. Bull., 17: 355.

Clayton, W. D. & S. A. Renvoize. 1986. Genera Graminum—grasses of the world. Kew Bull. Add.

Ser., 13.

Fennah, R. G. 1969. Damage to sugar cane by Fulgoroidea and related insects in relation to the

metabolic state of the host plant. Chapter 18. pp. 367-389. In Williams, J. R., J. R. Metcalfe,

R. W. Mongomery & R. Mathes. Pests of sugar cane. Elsevier Publishing Co., New York.

Fleady, S. E. & S. W. Wilson. 1990. The planthopper genus Prokelisia (Homoptera: Delphacidae):

morphology of female genitalia and copulatory behavior. J. Kansas Entomol. Soc., 63: 267-

278.

Jiang, B. H. 1976. Studies on Eoeurysa flavocapitata Muir, a new sugarcane planthopper to Taiwan.

Rep. Taiwan Sugar Res. Inst., 74: 53-62.

Metcalf, Z. P. 1943. General catalogue of the Hemiptera. Fasc. IV. Fulgoroidea, Part 3. Araeopidae

(Delphacidae). North Carolina State University, Raleigh, North Carolina.

Metcalfe, J. R. 1969. Studies on the biology of the sugar cane pest Saccharosydne saccharivora

(Westw.) (Horn., Delphacidae). Bull. Entomol. Res., 59: 393-408.

Mirza, R. P. & M. A. H. Qadri. 1964. Black leafhopper of sugarcane of Rajshahi, east Pakistan.

Univ. Stud. Univ. Karachi, 13: 31-34.

Muir, F. 1913. On some new Fulgoroidea. Proc. Hawaiian Entomol. Soc., 2: 237-269.

Qadri, M. A. H. 1963. Sugar cane pest of East Pakistan. Scientist, Karachi, 6: 46.

Wilson, M. R. 1988. A faunistic review of Auchenorrhyncha on sugarcane, pp. 485—492. In Yidano,

C. & A. Arzone (eds.). Proceedings of the 6th Auchenorrhyncha Meeting, Turin, Italy, Sept. 7-

11,1987. Consiglio Nazionale Delle Ricerche, Italy Special Projects, IPRA, University of Turin,

Turin, Italy.

Wu, R. H. & C. T. Yang. 1985. Nymphs of Delphacidae from Taiwan (I) (Homoptera: Fulgoroidea).

J. Taiwan Mus., 38: 95-112.

Yang, C. T. 1989. Delphacidae of Taiwan (II) (Homoptera: Fulgoroidea). Nat. Sci. Council (Taiwan)

Publ., 6.

Received 24 July 1991; accepted 23 October 1991.

PAN-PACIFIC ENTOMOLOGIST

68(2): 140-143, (1992)

A REDESCRIPTION OF ORDOBREVIA NUBIFERA (FALL)

(COLEOPTERA: ELMIDAE)

William D. Shepard 1

Department of Entomology, California Academy of Sciences,

San Francisco, California 94118

Abstract. — Newly found variation in the size, sculpturing and color of Ordobrevia nubifera (Fall)

is described. This variation may be linked to varying larval developmental rates.

Key Words.—Insects, Coleoptera, Elmidae, Ordobrevia, variation, development

The genus Ordobrevia was erected in 1953 by Sanderson, with Stenelmis nu¬

bifera Fall, 1901 as its type species. At that time, S. nubifera was the sole member

of the “ nubifera group” of Stenelmis (Sanderson 1938). Twelve more species of

Ordobrevia have been described since 1953. Four are from the Palaearctic region

(Japan) and eight are from the Oriental region (Brown 1981). It thus appears that

Ordobrevia nubifera represents an intrusion of the Palaearctic/Oriental fauna into

the Nearctic fauna. Zaitzevia (Elmidae) and Eubrianax (Psephenidae) show sim¬

ilar intrusions (Brown 1981).

Ordobrevia nubifera occurs only in North America, and its known range extends

from California to Washington (Brown 1972). More is known about California

populations than those in other areas. Within California, O. nubifera occurs widely

throughout all the various mountain ranges. It inhabits streams of all sizes from

first order to much larger. It seems to prefer microhabitats with faster flows and

coarser substrates.

Several years ago I found what seemed to be a new species of Ordobrevia. It

was larger and more robust, and it had more coarse granulation and a distinctly

different color pattern. I came embarassingly close to describing it as a new species.

Subsequent collections have shown it to be the end of a previously unknown range

of variation within O. nubifera. Recent studies of the elmid fauna of Taiwan by

M. L. Jang and P. S. Yang, and an ongoing revision of Stenelmis by Kurt Schmude

have called into question the status and identity of Ordobrevia. Because of these

two recent studies and discovery of additional intraspecific variation in O. nu¬

bifera, I decided to review what was known about Ordobrevia nubifera.

Existing work that illustrates O. nubifera includes the following: for larvae,

ninth abdominal tergum (Sanderson 1953: fig. 24), mesothorax (Leech & Chandler

1956: fig. 13:5 lh), habitus (Brown 1972: figs. 163 and 164), head (White et al.

1984: fig. 19.246); and for adults, aedeagus (Sanderson 1938: fig. 1), antenna

(Sanderson 1938: fig. 7), elytral pattern (Sanderson 1938: fig. 19), elytron (Leech

& Chandler 1956: fig. 13:52g), and habitus (Brown 1972: fig. 25, White et al.

1984: fig. 19.272).

Ordobrevia nubifera (Fall) 1901

Redescription, —(both sexes, except as indicated).—BODY: Body elongate, slender (Fig. 1) to robust

(Fig. 2), parallel sided; sculpturing and granulation slight to very coarse. Pronotum narrower than

1 Mailing address: 6824 Linda Sue Way, Fair Oaks, California 95628.

1992

SHEPARD: ORDOBREVIA REDESCRIPTION

141

Figures 1-5. Ordobrevia nubifera. Figure 1. Typical morph. Figure 2. Large morph. Figure 3. Right

wing. Figure 4. Aedeagus (a—dorsal view, b—lateral view, c—ventral view). Figure 5. Ovipositor (a—

dorsal view, b—lateral view).

elytra. Body uniformly brown to testaceous, with a transverse yellow band across the middle of the

elytra. Length 2.0-2.6 mm; width 0.8-1.2 mm. HEAD: Head covered with granules. Granules lon¬

gitudinally elongate on the epicranial surface, less elongate elsewhere. Antennal ridges prominent

dorsally. Fronto-clypeal suture absent. Clypeus with coarse setae on apical margin. Labrum dark

brown, shiny, coriaceous; apical margin with thick medially curved setae. Mandibles prominent; tips

trifid. Palpi three-segmented, last segment broad and apically truncate; labial palpi lighter than max¬

illary palpi. Antennae eleven-segmented; segments one to six narrow; segments seven to eleven apically

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

widened forming a weakly defined club, possessing lateral tufts of whitish setae. PRONOTUM: Pro-

notum usually wider than long, width greatest just behind middle. Base of pronotum wider than apex,

lateral margins sinuate and weakly serrate. Entire surface covered with close-set granules and punctures.

Median longitudinal sulcus in basal three-fourths of pronotum; sulcus deeper anteriorly. Base of median

sulcus with two depressions on each side. ELYTRA: Elytra with punctae arranged in striae. Punctures

smaller and less distinct in apical one-third, obsolete at tip. Surface between punctures smooth and

shiny to coarsely granulate. First and accessory striae join in basal one-fourth. Intervals raised; third

intervals carinate in basal third; sixth intervals forming sublateral carinae that reach almost to apex.

Humeri distinct and produced beyond basal pronotal margins. Epipleura extend almost to tip of elytra.

WINGS: Wings entire (Fig. 3); posterior edge with fringe of fine setae. Venation reduced. VENTER:

Hypomera, pro-, meso-, and metastemum covered with granules separated by less than their own

width. Prostemal margin projecting anteriorly under head and labium. Prostemal process parallel¬

sided, projecting beyond coxae, with apex apically rounded. Mesostemum very short, broadly truncate

apically, reaching only to middle of mesocoxae; median longitudinal sulcus present. Metastemum

similar to mesostemum but longer, with apex broadly emarginate. Abdomen with five visible segments;

males with segment V with lateral margins produced into short wide spines and narrowly emarginate

apically; females with lateral margins only weakly spiniform and apex broadly emarginate. Granules

on segment I rounded, separated by own width, those on segments II-V elongate, more widely sep¬

arated. All segments with lateral flanges closely fitting into the epipleura. LEGS: Pro- and mesocoxae

rounded, metacoxae transverse. Metathoracic legs slightly longer than others. Granules on coxae

rounded, close-set. Granules on trochanters 2.0 x as long as wide. Granules on femora and tibiae very

elongate, arranged parallel to the axes of the legs. GENITALIA: Male genitalia (Fig. 4) with parameres

shorter than median piece; accessory sclerites include a longitudinally elongate ventral sclerite and a

short transverse rectangular sclerite located between the tips of the median piece and the ventral piece;

basal piece weakly sclerotized, varying from a flattened U-shape to a complete ring. Female genitalia

(Fig. 5) typical for elmids.

Discussion

This species shows considerably more size variation than previously stated.

However, I think that size ranges given in the literature are often the result of

repetition of earlier measurements that may have been based on relatively few

specimens. The morph that I earlier almost mistook for a new species is now

considered to be the upper range of size for O. nubifera. It is longer and much

wider than “typical” specimens. A more strongly developed granulation, punc-

tation and sculpturing, and a trend toward uniformly dark brown color is cor¬

related with increasing size. Other morphologic characters remain unchanged,

albeit somewhat larger in proportions.

There seem to be higher percentages of larger individuals of Ordobrevia nubifera

in populations in the northern Coastal Mountains and the Warner Mountains

than in other parts of California (unpublished data). These larger individuals of

O. nubifera often occur with “typical” specimens. However, the larger morph is

a larger percentage of the population in rather cold streams, or in areas that

experience rather cold winter weather. I now think that they represent larvae in

which the normal developmental rate was retarded allowing the normal growth

rate to produce a larger-than-normal individual. Larger larvae, of course, mean

larger adults. Differential manipulation of developmental rates has been shown

to cause different sized morphs in the worker caste of ants (Oster & Wilson 1978).

I have found similar, larger-than-normal individuals in several other elmid genera.

Retardation of developmental rates could be a result of intrinsic factors (e.g.,

genetic recombinations, random mutations) or extrinsic factors (e.g., cold tem¬

peratures, limited supplies of specific nutrients). My experiences lead me to favor

the cold temperatures as the major influencing factor. Some described variations

1992

SHEPARD: ORDOBREVIA REDESCRIPTION

143

in Microcylloepus, another elmid, appear to be temperature correlated (Shepard

1990). However, in this case, smaller-than-normal individuals (and species) occur

in warmer habitats. There, I believe, the developmental rate is accelerated while

the growth rate remains unaffected. Thus the individuals mature at a smaller body

size. More study is warranted concerning the influences of varying temperatures

upon life cycle events in natural populations.

My near description of a new species of Ordobrevia should alert other taxon¬

omists to sample extensively and pay more attention to intraspecific variation.

Even though I had previously collected and identified hundreds of the “typical”

O. nubifera, I did not have an accurate idea of the total intraspecific variation

until I collected in streams that were perennially cold, or were very cold during

large parts of the year. I am also reminded of the necessity of obtaining population

samples large enough to contain the rarer morphs within a population. This is

especially a problem when sampling aquatic insects in such an ecologically diverse

area as California. Larger population samples have led me to question the validity

of several California elmid species.

Acknowledgment

Harley P. Brown kindly loaned specimens from his collection for this study,

and generously offered his comments on the manuscript. Part of this study was

supported by a summer research position at the University of Oklahoma Biological

Station.

Literature Cited

Brown, H. P. 1972. Aquatic dryopoid beetles (Coleoptera) of the United States. Biota of freshwater

ecosystems identification manual 6. U.S. Environmental Protection Agency. U.S. Government

Printing Office, Washington, D.C.

Brown, H. P. 1981. A distributional survey of the world genera of aquatic dryopoid beetles (Cole¬

optera: Dryopidae, Elmidae, and Psephenidae sens. lat.). Pan-Pacif. Entomol., 57: 1-6.

Fall, H. C. 1901. List of the Coleoptera of southern California, with notes on habits and distribution

and descriptions of new species. Occ. Papers Calif. Acad. Sci., 8.

Leech, H. B. & H. P. Chandler. 1956. Aquatic Coleoptera. Chapter 13. pp. 293-371. In Usinger, R.

L. (ed.). Aquatic insects of California. University of California Press, Berkeley.

Oster, G. F. & E. O. Wilson. 1978. Caste and ecology in the social insects. Princeton University

Press, Princeton, New Jersey.

Sanderson, M. W. 1938. A monographic revision of the North American species of Stenelmis

(Dryopidae: Coleoptera). U. Kans. Sci. Bull., 25: 635-717.

Sanderson, M. W. 1953. A revision of the Nearctic genera of Elmidae (Coleoptera). J. Kans. Entomol.

Soc., 26: 148-163.

Shepard, W. D. 1990. Microcylloepus formicoideus (Coleoptera: Elmidae), a new riffle beetle from

Death Valley National Monument, California. Entomol. News 101: 147-153.

White, D. S., J. T. Doyen & W. U. Brigham. 1984. Aquatic Coleoptera. Chapter 19. pp. 361-437.

In Merritt, R. W. & K. W. Cummins (eds.). An introduction to the aquatic insects of North

America (2nd ed). Kendall/Hunt Publishing Company, Dubuque, Iowa.

Received 25 June 1991; accepted 1 November 1991.

PAN-PACIFIC ENTOMOLOGIST

68(2): 144-152, (1992)

A REVIEW OF THE SWEETPOTATO WHITEFLY

IN SOUTHERN CALIFORNIA

Raymond J. Gill

Insect Taxonomy Laboratory,

California Department of Food & Agriculture,

Sacramento, California 95814

Abstract.— The sweetpotato whitefly, Bemisia tabaci Gennadius, of Palaearctic origin was orig¬

inally introduced into California in the late 1920s. Since that time it has been restricted to the

state’s southern desert valleys and has, at times, been a significant agricultural problem. In the

mid-1980s, however, a new “strain” of B. tabaci was introduced to southern California and has

wreaked great havoc in the area. This strain, from poinsettia plants, has become known as the

B strain, poinsettia strain or poinsettia whitefly. This paper documents the new introduction,

notes the poinsettia strain’s differences from other B. tabaci, and assesses the possibilities for its

control.

Key Words. — Insecta, Aleyrodidae, Bemisia tabaci, sweetpotato whitefly, California

California has been experiencing serious problems with whiteflies during the

last several years. Of the approximately 1160 described species of whiteflies in

the world, 54 occur in the state along with approximately a dozen undescribed

native species. Of California’s described species, at least 11 were introduced by

man’s activities, and five have been introduced in the last 15 years. Several of

the introduced species have become serious pests and two are currently quite

problematic: the ash whitefly, Siphoninus phillyreae (Haliday), and the sweet¬

potato whitefly, Bemisia tabaci (Gennadius). These species currently have ex¬

tremely large populations in areas of California.

The ash whitefly, an easily recognized species, was introduced into the state in

the late 1980s, and although it spread rapidly with tremendous population ex¬

plosions (Sorensen et al. 1991), a successful parasite was found (Bellows et al.

1991) and effective biological control has progressed rapidly. The sweetpotato

whitefly (SPW), however, has been in California since the 1920s (Russell 1975),

but only in the last two decades, particularly the early 1980s, has it been a serious

agricultural problem (Natwick & Zalom 1984) and a taxonomic and ecological

curiosity. Currently, it is in a disastrous expansion phase in southern California,

which involves the acquisition of many new hosts. This paper documents the

ecological history and potential taxonomic problems with SPW in southern Cal¬

ifornia.

Background

The Bemisia tabaci was originally described as an Aleyrodes from tobacco in

Greece in 1889 (Gennadius 1889). Since then, the species has been redescribed

in synonymy many times (Table 1). The insect has spread to most tropical and

subtropical areas of the globe, occasionally causing serious damage upon colo¬

nization. It was first recorded from India in 1905 (Misra & Lambda 1929, Reddy

Sl Rao 1989, Immaraju 1989), and by 1919 had become a serious pest of cotton

in the Punjab (now Pakistan) (Immaraju 1989). It has been reported as a serious

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GILL: SWEETPOTATO WHITEFLY IN CALIFORNIA

145

Table 1. Taxonomic synonyms of sweetpotato whitefly. 3

Genus

Species

Author

Date

Type locality

Aleyrodes

tabaci

Gennadius

1889

Greece

Aleyrodes

inconspicua

Quaintance

1900

Florida

Bemisia

emiliae

Corbett

1926

Sri Lanka

Bemisia

costa-limai

Bondar

1928

Brazil

Bemisia

signata

Bondar

1928

Brazil

Bemisia

bahiana

Bondar

1928

Brazil

Bemisia

gossypiperda

Misra & Lambda

1929

Pakistan

Bemisia

acyranthes

Singh

1931

Pakistan

Bemisia

hibisci

Takahashi

1933

Taiwan

Bemisia

longispina

Priesner & Hosny

1934

Egypt

Bemisia

gossypiperda var.

mosaicaivectura

Ghesquiere

1934

Zaire

Bemisia

goldingi

Corbett

1935

Nigeria

Bemisia

nigeriensis

Corbett

1935

Nigeria

Bemisia

rhodesiaensis

Corbett

1936

Rhodesia

Bemisia

manihotis

Frappa

1938

Madagascar

Bemisia

vassyierei

Frappa

1939

Madagascar

Bemisia

lonicerae

Takahashi

1957

Japan

Bemisia

minima

Danzig

1964

U.S.S.R.

Bemisia

miniscula

Danzig

1964

U.S.S.R.

3 See Taxonomic Assessment and Biological Control section for comments on B. poinsettiae Hempel,

1923.

pest of various crops in: the West Indies, Nicaragua, Venezuela, Brazil, Turkey,

Israel, Egypt, Sudan, Iran, Thailand, and the Philippines. In addition, it is known

from southern Europe, the Middle East, much of Africa, Madagascar, Sri Lanka,

China, Malaya, Australia, New Guinea, Fiji, and Hawaii, among other locations.

By 1978, SPW was known from at least 420 plant species in 18 families (Mound

& Halsey 1978, Greathead 1986), but new hosts are being continually added as

the current infestation in California and Arizona grows. Currently, SPW is a major

economic pest of cotton, tobacco, cassava, sweetpotato and soy bean in many

areas of the world.

After its introduction to the U.S., SPW was redescribed as Bemisia inconspicua

by A. L. Quaintance (1900) from material collected on okra and sweetpotato in

Florida between 1897 and 1898. Later, museum specimens were found to have

been collected in Pomona, Putnam County, Florida in 1894 (Russell 1975). It

has since spread across the southern part of the U.S. Prior to 1985, it was found

in outdoor environments in Florida, Georgia, Texas, Arizona and California.

Recently, it has been found in extremely high populations in the agricultural areas

of Arizona, California, Texas and northwestern Mexico.

History in California

Specimen records at the U.S. National Museum of Natural History indicate it

had been introduced into California by at least 1928 (Russell 1975), when it was

collected on cotton at Calipatria, Imperial County. Subsequent records of early

spread in California are shown in Table 2. Although SPW was in California in

the late 1920s, it was found outdoors only in the desert valleys of Imperial,

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THE PAN-PACIFIC ENTOMOLOGIST

Yol. 68(2)

Table 2. The earliest records of the spread of Bemisia tabaci within California, after its 1928

introduction (Calipatria, Imperial Co.) on cotton.

Year

County

Location

Host

1947

Riverside Co.

Coachella

sweetpotato

1950-1954

Riverside Co.

Indio

cotton

1951

Imperial Co.

Calexico

cotton

1952

Riverside Co.

Coachella

cotton

1953

Riverside Co.

Thermal

sweetpotato

1953

Riverside Co.

Mecca

sweetpotato

1954

Imperial Co.

Imperial

cotton

1954

Riverside Co.

Riverside

cotton 3

1955

Riverside Co.

Riverside

euphorbia 3

1961

San Bernardino Co.

Yucca Valley

Hibiscus sp.

a In greenhouse.

Riverside, San Bernardino and San Diego Counties. It was seldom, if ever, found

in greenhouses in California, and then usually on plants imported recently from

other states.

In the Imperial Valley of California, a curious and disastrous phenomenon

occurred with SPW in the summer and fall of 1981; its populations exploded on

numerous crops, including cotton, melons and lettuce. D-Vac® monitoring by

University of California Agricultural Extension personnel collected over 60,000

whiteflies per 100 sweeps of the devices (Natwick & Leigh 1984). The large

numbers of whiteflies were severely debilitating the infested crops, and also trans¬

mitting serious viral diseases to the crops. High incidences of squash yellow leaf

curl and lettuce infectious yellows resulted in premature plow-down and total

crop loss in many lettuce and melon fields in fall 1981 (Duffus & Flock 1982,

Natwick & Zalom 1984).

Although 1981 was a disastrous year for the growers in the Imperial, Bard and

Palo Verde Valleys of California, SPW had actually been building up populations

over the preceding several years. University of California extension personnel had

been making routine whitefly counts for many years (Natwick & Leigh 1984)

because SPW and another species, banded-winged whitefly [Trialeurodes abuti-

loneus (Haldeman)] were found on cotton infested with cotton leaf crumple, a

viral disease. Prior to 1975, D-Vac® catches for SPW were running consistently

lower than 300-400 per 100 sweeps. However, in 1975 the number jumped to

nearly 4300 whiteflies per 100 sweeps. Numbers dropped the next year, only to

leap to an incredible 35,000 whiteflies per 100 sweeps in 1977. The populations

dropped again to near zero in 1978, only to be followed by the disastrous rebound

seen in 1981.

There are several possible causes for these population explosions, which prob¬

ably result from several interrelated concurrent events. Starting in 1975, the

southern California desert areas experienced unusually warm winter temperatures,

with a virtual absence of days below freezing (only two years out of nine had

recorded temperatures below 0° C) (Flock & Christopherson 1985). Because SPW

is apparently of tropical origin, cool or cold temperatures appear to prevent normal

development, while high summer temperatures and humidity probably enhance

development. Comparing the warm winter temperature ranges in the Imperial

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GILL: SWEETPOTATO WHITEFLY IN CALIFORNIA

147

Valley with the sudden upsurges observed in SPW populations shows an intriguing,

yet not exactly corresponding, correlation.

A second event in the Imperial Valley area in 1975 involved the first use there

of synthetic pyrethroid insecticides for general pest control (E. T. Natwick, per¬

sonal communication). Such pyrethroids have a devastating effect on the natural

enemies (primarily parasitoids) of SPW. Essentially the lack of cold winter tem¬

peratures allowed SPW to maintain larger than normal populations through the

winter, and a reduced natural enemy population allowed SPW an unencumbered

pathway to the devastating populations that were encountered between 1975 and

1990. By 1986, researchers and growers were discovering ways to deal with the

SPW problem. They observed that the SPW population was building up on cotton

to such large levels that by the time the cotton was ready for the normal fall

defoliation and harvest, it would be heavily covered with honeydew and sooty

mold. When the cotton was defoliated, the whiteflies would move in large numbers

into other crops including squash, melons, lettuce, sugar beets, tomatoes and other

specialty crops, transmitting viral diseases presumably picked up from weeds and

other virus infected hosts. By defoliating cotton early, it was found that SPW did

not have time to develop large populations that could move onto other crops,

and the cotton would be fairly free of honeydew and sooty mold (Meyerdirk et

al. 1986).

By 1990, just when the SPW problem seemed to be under control in the desert

southwest, a second disastrous phenomenon occurred, this time as a result of

events in Florida four years earlier. SPW had maintained a foothold in Florida

for many years, seldom being more than a scientific curiosity. Inexplicably in

1986, growers of greenhouse poinsettias had a devastating outbreak of SPW that

appeared overly resistant to chemical control (Hamon & Salguero 1987). As the

summer of 1986 wore on, these SPW jumped to numerous other greenhouse

bedding plants and nursery stock; they also began infesting outdoor vegetable

crops and gardens with disastrous results. By 1987, the large poinsettia nurseries

of San Diego County were found to be infested, and over the next year or two

SPW was found on poinsettias in many greenhouses throughout California. Shortly

thereafter, in late 1990, SPW moderately infested commercial citrus groves near

Phoenix, Arizona; it had never been found on this crop in economically damaging

populations before (D. N. Byrne, personal communication). SPW was observed

to spend the winter in fairly large numbers on this plant.

Prior to the find of SPW on citrus, researchers in Florida and Arizona were

beginning to evaluate some of the characteristics and effects of the SPW “strain”

(hereafter referred to as poinsettia SPW) that began attacking greenhouse poin¬

settias and other crops in Florida in 1986. Poinsettia SPW was found to cause

virus-like symptoms in cucurbits (Yokomi et al. 1990; Costa & Brown 1990,

1991a, b) that were quickly called “squash silver leaf.” These symptoms probably

are related to a phytotoxin injected into the plant by poinsettia SPW, because the

plants recovered from the effects when the whiteflies were removed.

In contrast, it was found that the original “strain” of SPW (hereafter referred

to as cotton SPW) reared from cotton, squash and other crops in Arizona (Costa

& Brown 1990, 1991) and California (Perring et al. 1991) did not produce these

same symptoms in squash plants. Shortly thereafter, researchers in Arizona (Costa

& Brown 1990, 1991) and California (Perring et al. 1991) investigated the isozymic

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variation in poinsettia versus cotton SPW “strains” using several different elec¬

trophoretic techniques. Populations of poinsettia SPW from poinsettias were found

to show slight, but consistently different, esterase banding patterns from those

cotton SPW populations that had existed in the southwest prior to 1986. Tax¬

onomists, however, have not been able yet to show a morphological difference

between these populations, and they are both currently considered to be B. tabaci.

After SPW was discovered on citrus in Arizona, it was assayed using electro¬

phoretic techniques and found to be poinsettia SPW (D. N. Byrne, personal

communication). By early spring 1991, it became evident that SPW was occurring

in large numbers over the winter on cole crops, particularly broccoli, in the Yuma

and Imperial Valleys; the presence of SPW on cole crops in winter had never been

experienced in these areas before. These whiteflies also were determined to be

poinsettia SPW (T. M. Perring, personal communication).

By July 1991, it was obvious that a major and catastrophic change had taken

place in the SPW situation in the Imperial and Palo Verde Valleys (Weddle &

Carson 1991, Perring et al. 1991). Observers in the field made many startling

discoveries. Some cotton was covered completely by adult whiteflies before the

plants could produce more than three or four leaves. The first leaves of squash

plants were being devastated before the plants could send out three or four inches

of runners. Many fields were disced under. The cotton that did mature was hope¬

lessly sticky with honeydew before the bolls could open. Fields of alfalfa were so

sticky they could not be baled. In late August, table grape vineyards on the north

shore of the Salton Sea in Riverside County were found heavily infested and sticky

with SPW. The same was found on new growth of grapefruit plantings and on

many weed species in the immediate vicinity. Some of this infestation apparently

originated from clouds of SPW that have been observed flying across the Salton

Sea from breeding grounds in the Imperial Valley. These whiteflies are generally

considered to be the poinsettia SPW.

After just one season, cotton SPW is now believed to be practically nonexistent

in California (T. M. Perring, personal communication), due either to interbreeding,

competition between the two strains, the extreme cold temperatures of December

1990, or possibly all of these reasons. Cross breeding experiments that are now

being conducted in Arizona and California may shed light on this phenomenon.

However, work that had been done on the two strains prior to this summer has

also produced some other interesting differences between the two SPW strains.

Poinsettia SPW is more cold tolerant. The time required to complete a generation

has been found to be slightly shorter in poinsettia SPW, or identical in the two

strains (usually 16-23 days), but poinsettia SPW is considered to be five times as

prolific (T. M. Perring, personal communication). Poinsettia SPW has been found

to extract five times as much nutrient material from plants and, therefore, produces

five times as much honeydew as cotton SPW. Although cotton SPW is thought

to be a better virus disease vector, at least with lettuce infectious yellows (J. E.

Duffus, personal communication), poinsettia SPW has produced such large pop¬

ulations that plants die before virus symptoms appear (F. Laemmlin, personal

communication), so its effectiveness in virus transmission is unknown. Further¬

more, poinsettia SPW severely attacks more crops, including some not previously

utilized by cotton SPW.

By the first week in October 1991, SPW had been found in moderate numbers

in dooryard vegetable gardens in the city of San Bernardino. This is the first

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149

important record for any SPW outdoors in California outside of the desert valleys.

One week later, SPW was found on established, outdoor poinsettia bushes in

Riverside, Riverside County. The owners of these bushes said that the whiteflies

had been a problem since the previous year. By December, SPW had been found

in three southern San Joaquin Valley counties in held situations not associated

with nurseries.

Taxonomic Assessments and Biological Control

As was done with ash whitehy, the first step that should be taken to find an

effective biological control for SPW is to identify the native home of the insect,

so that natural enemies can be found. In the case of SPW, however, this creates

an immediate dilemma. Up until recently, the native home of SPW was thought

to be either the Orient or Africa/the Middle East (Mound 1963, Lopez-Avila

1986, Anonymous 1987). Other Bemisia are prevalent in southern Russia and

are also known from mainland Asia, southeast Asia along the Pacific rim, Africa,

and one species each from South America and the western United States (Mound

& Halsey 1978). Certainly, the area to the north and west of Pakistan shows the

greatest diversity in parasitoids of Bemisia (Mound & Halsey 1978, N. Mills

1992), reputedly an indication of a genus epicenter.

SPW was probably moved around the world at a very early date, but was not

described until 1889. Because SPW has probably been reintroduced into many

countries numerous times, it becomes extremely difficult to trace the origin of the

whitefly. Because poinsettia and cotton SPW cannot presently be separated mor¬

phologically, we cannot effectively access pre-1986 museum specimens to ascer¬

tain where poinsettia SPW occurred prior to 1986. Lacking adequate surveys

using electrophoretic analysis to separate the strains, we so far have very limited

knowledge of where poinsettia SPW presently occurs in the world. We know only

that it has been transported over most of the U.S. and the Caribbean on poinsettia

and other nursery crops (J. K. Brown, personal communication). It has also been

transported to Canadian greenhouses (Broadbent et al. 1989), from where it es¬

caped to the field but probably could not survive the Canadian winters.

Recently, however, evidence is emerging that indicates B. t abaci may be of New

World origin. For example, it seems to do best on hosts that are of New World

origin (unpublished data), such as sweetpotato, poinsettia, tomato, common bean,

squash, peppers, and tobacco. Further, in Puerto Rico (Bird 1957), a strain of B.

tabaci was identified that feed solely on Jatropha gossypifolia L., a plant of New

World origin, despite numerous trials on other hosts; a feeding pattern that seems

highly unlikely if B. tabaci were of Old World origin. A New World origin hy¬

pothesis for B. tabaci would have important ramifications for searching for natural

enemies, switching the search emphasis to the Neotropics.

New studies of genetic variance may also suggest a New World origin for B.

tabaci. Wool et al. (1991) examined isozymes of B. tabaci populations in Israel

and found genetic uniformity, with no geographical races existing there. However,

in examining B. tabaci from Columbia, they found differing esterase patterns

among populations from various Columbian regions. In fact, the esterase pattern

found in samples from the Valle, near Cali, were “very similar to the Israeli

pattern” (Wool et al. 1991: 228). Similar circumstances exist in other homopter-

ans, suggesting that centers of origin for a species probably have higher genetic

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variability than do invaded areas. For example, among now cosmopolitan aphids,

such as Myzus persicae (Sulzer) and Macrosiphum euphorbiae (Thomas), electro¬

phoretic surveys of variance in North America indicate that the former, with

zero variability in the Nearctic, probably had a limited introduction to that con¬

tinent, whereas the latter, with a higher heterozygosity level, is probably a Nearctic

endemic (May & Holbrook 1978).

This limited “founder effect” variance appears contrary to implied increasing

genetic variance in other invasive whiteflies, such as Siphoninusphillyreae recently

in California, where Sorensen et al. (1991) proposed a mutation-driven expansion

of feeding-range, caused by explosive invasive populations in the absence of

population controls. (A situation also similar to poinsettia SPW there.) Clearly,

electrophoretic surveys of S. phillyreae in California should be (or should have

been ) conducted to monitor its heterozygosity during the geographical expansion.

If limited genetic variability were maintained for S. phillyreae during its California

explosion, then theories of expansions in the range of host-feeding during invasions

might require modification (J. T. Sorensen, personal communication).

SPW, like several other whiteflies and scale insects, tends to be morphologically

variable depending on both its host and on its location on the plant (Mound 1963).

In SPW, the last stage nymph (“pupa”) usually has a smooth dorsal surface if the

host leaf is smooth. Alternatively, if the underside of the host leaf is covered with

stiff hairs or spines, the pupa usually possesses very long (usually two to eight)

dorsal setae arising on the head, thorax and abdominal areas. The pupa also tends

to develop other unique characteristics on given hosts, as has been demonstrated

by cross-rearing various populations on different hosts. Before interhost mor¬

phological variability was realized, numerous Bemisia synonyms were described

as distinct species, but are now considered to be B. tabaci (Russell 1957) (Ta¬

ble 1).

Partly because of host induced morphological variation, conventional taxon¬

omists have not been able to find characters in any of the life stages of SPW that

would indicate that more than one species is present. Current diagnostic methods

require either live insects to test for the ability to induce squash silver leaf symp¬

toms, or adults that have been adequately preserved for electrophoretic analysis.

What poinsettia SPW actually represents remains in question. Because no dif¬

ferentiating morphological traits have been found it must currently be considered

to be the same species as cotton SPW, B. tabaci. Yet its explosive population

growth and host acquisitions in the presence of cotton SPW suggest that it probably

represents something more than a simple biotype, perhaps a sibling species. (In¬

terestingly, type material of Bemisia poinsettiae Hempel, 1923, described from

Brazil on Poinsettia [obtained by E. Delfosse], shows no conventionally used

morphological characters that can be used to separate it from B. tabaci, with which

it thus may be synonymous.) Although there are a few taxonomic tools that are

still available to use (e.g., morphometric multivariate analyses), it may take a

while before they can be adequately developed on this problem. However, even

if we can satisfactorily determine the relationships between the two SPW “strains,”

we will still require satisfactory control measures. Cotton SPW caused as much

as $100 million in agricultural losses in southeastern California in 1981 (Duffus

& Flock 1982). With recent developments, losses in 1991 may go well beyond

that mark, because now crops are being attacked that were not infested previously.

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151

Acknowledgment

I thank J. K. Brown, D. N. Byrne, E. Delfosse, J. E. Duffus, F. Laemmlin, N.

Mills, E. T. Natwick, and T. M. Perring for providing information used in this

review. John T. Sorensen also provided information, and expanded parts of the

last section of the article in galley.

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in populations of the whitefly Bemesia tabaci (Genn.) in Israel and Columbia: an interim report.

Insect Sci. Applic., 12( l/2/3):225—230.

Yokomi, R. K., K. A. Hoelmer & L. S. Osborne. 1990. Relationships between the sweetpotato

whitefly and the squash silverleaf disorder. Phytopathology, 80: 895-900.

Received 1 November 1991; accepted 26 November 1991.

PAN-PACIFIC ENTOMOLOGIST

68(2): 153-154, (1992)

Scientific Note

A NEW ANT INTRODUCTION FOR NORTH AMERICA:

PHEIDOLE TENERIFFANA (FOREL)

(HYMENOPTERA: FORMICIDAE)

During the spring of 1989, while spraying weeds at Admiral Kidd Park in

western Long Beach, California, I discovered several foraging columns of small

brown ants. The ants were nesting in the southeastern comer of the park, in sandy

soil. I was able to identify the ants to the genus Pheidole; later, I sent samples to

the Departments of Agriculture in Orange and Los Angeles counties, the California

Department of Food & Agriculture, and the Los Angeles County Natural History

Museum. Ultimately, the ants were identified to species as Pheidole tenerijfana

(Forel) on 20 Feb 1990 by E. O. Wilson of Harvard University. This species, a

native of north Africa and the Canary Islands, has never been recorded from

North America, although it had been previously found in Cuba in 1932 (Aguayo,

T. 1932. Bull. Brooklyn Entomol. Soc., 22: 219). Between 1989 and 1991, this

ant had spread to infest about five acres of the seven acre park site where it was

discovered.

The workers of P. tenerijfana are 2.5 mm long, with a black-brown head and

gaster, and a lighter brown thorax. Soldiers of the species have oversized heads

with powerful mandibles, and are the same colors as the workers, but larger and

3.75 mm long. The queen is entirely a shining dark brown, and 5.5 to 6.0 mm

long, while males are dull light brown to medium brown and 4.0 to 4.5 mm long.

The main function of the soldiers is to defend the nest, although both they and

the workers will fight with other ants over food or when they are invading new

territory.

In Admiral Kidd Park, I have observed on several occasions P. tenerijfana

displacing the Argentine ant, Iridomyrex humilis Mayr, another introduced pest

species. Between 13 Mar and 5 Jun 1990, and on 9 Sep 1991, P. tenerijfana

advanced into Argentine ant territories, attacking and destroying colonies and

taking over their nest sites. Similar interactions have been observed between

Iridomyrex humilis, and another Pheidole sp., P. megacephala (Fabr.), in Hawaii

and Bermuda (Haskins, C. P. & E. F. Haskins. 1965. Ecology, 46: 736-740;

Crowell, K. L. 1968. Ecology, 49: 551-555; Fluker, S. S. & J. W. Beardsley. 1970.

Ann. Entomol. Soc. Am., 63: 1290-1296).

In contrast, in the park, a native fire ant, Solenopsis xyloni McCook, often raids

the nests ofP. tenerijfana and may annihilate whole colonies. Curiously, however,

S. xyloni is itself displaced, at least partially, by I. humilis, so that a repetitious

cycle of displacement might occur. It may be possible that I. humilis is repelled

by a kariomone produced by P. tenerijfana, but which does not repel S. xyloni;

whereas, S. xyloni might be repelled by a kariomone produced by I. humilisl

Pheidole tenerijfana seems to have few “conflicts” with less aggressive native

ants in the park, but I have observed it attacking workers of the California red

harvester ant, Pogonomyrmex californicus Buckley. Other ant species present in

154

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 68(2)

the park are: Conomyrma bicolor Wheeler, Conomymra insana Buckley, Tapi-

noma sessile Say, Formica pilicornis Emery, Monomorium minimum Buckley and

Cardiocondyla ectopia Snelling.

Pheidole tenerijfana nests have many inseminated queens; 23 were observed in

one colony that was changing its nest site. The nests occur as large colonies with

low mounds in the soil, along curbs or sidewalks, at the edges of lawns, in cracks

in pavement, and at the bases of trees. New colonies are started by budding, with

new queens mating in the nest and moving with part of the existing colony to

form new nests in adjacent territory. The workers forage night and day, unless it

gets too hot (>26° C). However, if the nest is in a shady location, they will remain

active on the hottest days. Colony members are predacious on live insects, such

as noctuid or beetle larvae. They may also harvest seeds and scavenge dead or

dying insects. I have not observed them tending aphids, but they do feed on sweet

or greasy materials.

Record.-CALIFORNIA. LOS ANGELES Co.: Long Beach, Admiral Kidd Park, 7 Mar 1989, M.

J. Martinez.

Acknowledgment.—1 thank Edward O. Wilson, Harvard University, for the

determination of this ant to species, and Roy Snelling, Los Angeles County Natural

History Museum, for his assistance and support in identifying the ants. I also

thank Phil Hester and his staff of the Recreation and Marine Department, Park

Bureau, Long Beach Parks, for their cooperation, and my wife, Charlean, for her

help and patience.

Michael J. Martinez, Department of Parks and Recreation, City of Long Beach,

Long Beach, California 90815.

Received 30 September 1991; accepted 10 October 1991.

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