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PAN-PACIFIC ENTOMOLOGIST

76(1): 1-11, (2000)

THE AUSTRALIAN FRUIT FLY PARASITOID

DIACHASMIMORPHA KRAUSSIT (FULLAWAY): LIFE

HISTORY, OVIPOSITIONAL PATTERNS, DISTRIBUTION

AND HOSTS (HYMENOPTERA: BRACONIDAE: OPITNAE)

K. RUNGROJWANICH! AND G. H. WALTER

Department of Zoology and Entomology, The University of Queensland,

Brisbane, Queensland 4072, Australia

Abstract—Diachasmimorpha kraussii is a \arval-pupal parasitoid of tephritid fruit flies in Aus-

tralia. It is currently being considered for release against fruit fly pests in Hawaii. Virgin D.

kraussii females lived longer (mean = 31.4 days; n = 10) than mated females (mean = 27.6

days; n = 10) by a factor of about 12%. The rate of offspring production per day by virgins

(about four emerging adults per day) was the same as that of mated females, so virgins tended

to produce more offspring in total (mean = 125) than did mated females (mean = 112), but the

difference was not statistically significant. The time between egg deposition and emergence of

the resultant adult varied from 16 days to more than 300 days, and males achieved maximum

emergence before females. Adult wasps emerged at any time of the photophase, both under

laboratory and field conditions, but the rate declined towards the end of the daylight period.

Adult females oviposited more actively during the day than at night (30.8 vs 19 adults), and the

pattern tended to be stronger when wasps were exposed to hosts initially during the scotophase

(37.4 vs 18.4 adults). Mated females produced female-biased brood sex ratios of about 0.28

(proportion of males) on average, and the older the mother wasps the greater the proportion of

female offspring produced. Diachasmimorpha kraussii is distributed only in northern and eastern

Australia, as far south as New South Wales. It has been recorded from 13 host fly species and

in association with 18 host plant species.

Key Words.—Insecta, Diachasmimorpha kraussii, parasitoid, Braconidae, Tephritidae, oviposi-

tion, fecundity, sex ratio, delayed emergence, distribution, hosts.

Braconid parasitoids are the major natural enemies of tephritid fruit flies and

have been used for biological control in various parts of the world. Although

several species have been used in this capacity, fewer than a dozen are relatively

well known, e.g., Diachasmimorpha longicaudata (Ashmead), D. tryoni (Cam-

eron), Fopius arisanus (Sonan), F. vandenboschi (Fullaway) and Psyttalia fletch-

eri (Silvestri) (Willard 1920; Leyva et al. 1991; Ramadan et al., 1991, 1992,

1994a, b; Messing & Jang 1992; Messing et al. 1996; Purcell 1998). Little in-

formation is available about the other species, and for some not even basic in-

formation on life history has been published. This is the case with the Australian

species D. kraussii, despite it having been described almost 50 years ago (Ful-

laway 1951), when it was used in biological control efforts. Diachasmimorpha

kraussii was introduced into Hawaii between 1947 and 1952, cultured on Ceratitis

capitata Wiedemann and released in areas infested with this pest species. It did

not establish permanently (Clausen et al. 1965), but the species is currently being

cultured and considered for further releases against C. capitata (R. Messing, per-

sonal communication). The hosts recorded for D. kraussii to date include Bactro-

cera barringtoniae (Tryon), B. cacuminata (Hering), B. dorsalis (Hendel), B. jar-

' School of Agricultural Extension and Cooperatives, Sukothai Thammathirat Open University, Pak-

kred, Nonthaburi 11120, Thailand.

2 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

visi (Tryon), B. kraussii (Hardy), B. murrayi (Perkins) and B. pallida (Perkins &

May) (Clausen et al. 1965). Recent results show, however, that D. kraussii cannot

develop successfully|in Hawaiian B. dorsalis flies (R. Messing, personal com-

munication).

To facilitate the use of D. kraussii in biological control, we investigated several

aspects of its biology and ecology, namely: (i) adult life span and fecundity, (ii)

diurnal and nocturnal levels of oviposition and (i11) developmental duration and

emergence patterns. The Australian distribution, host flies and host plants of D.

kraussii were evaluated from museum specimens and from personal collections.

MATERIALS AND METHODS

Insect Cultures.—A Bactrocera tryoni (Froggatt) culture was initiated from a

colony held at the Queensland Department of Primary Industries, Long Pocket,

Brisbane. We used the culture technique described by Heather & Corcoran (1985),

except for providing|Vegemite® (concentrated yeast extract) as a protein source

for adult flies, instead of protein hydrolysate.

The D. kraussii colony was started with about 35 pairs reared from B. tryoni

puparia derived from) Brazilian cherries (Eugenia uniflora L.) collected in St Lu-

cia, Brisbane, in November 1990. The adult D. kraussii were exposed, in a 15 X

15 X 30 cm perspex! cage, to 3rd instar (eight days old since oviposition) fruit

fly larvae that were restrained in an “‘oviposition unit’? (see below). Honey was

always available to adult wasps, both in culture and in all experiments. Hosts and

parasitoids were reared at 25 + 1° C, 60 + 5% R.H. and 12:12 L:D. Experiments

were also conducted under these conditions.

‘“‘“Oviposition units’’ were prepared from a 7.5 cm diameter plastic lid with a

raised outer rim (0.6 cm high). The inside cavity of the lid was filled with larval

medium into which the B. tryoni larvae had been placed. The whole unit was

wrapped with tightly-stretched Parafilm®, through which the wasps readily ovi-

posited.

For culturing purposes, oviposition units were exposed to D. kraussii adults for

24 h. Units were then unwrapped and placed in a 1000 mI plastic container lined

with sawdust. No additional larval medium was needed because the fruit flies

always pupated within 24 h of their removal from the oviposition cage. An excess

of larval medium, which would encourage fungal growth, was thus avoided. Fun-

gi, if present, made it difficult for fruit fly larvae to spring into the sawdust to

pupate. The puparia were sieved from the sawdust and held in a 30 ml plastic

cup for adult emergence.

Life Span, Developmental Duration and Reproductive Capacity.—Ten D.

kraussii females, three days old, were exposed to conspecific males, one day old,

for mating (one pair/mating unit (Rungrojwanich & Walter 1999)). After each

female had mated (all mated within 10 min), she was held alone in an inverted

125 ml plastic container and exposed to hosts (i.e., early on the fourth day after

eclosion). Each container had a 3 cm diameter hole in the bottom with fine muslin

glued over it. The containers were kept upside down so that the lids could serve

aS Oviposition units. Each oviposition unit contained 20 B. tryoni larvae. After

each 24 h of exposure, the oviposition unit was replaced and the old one trans-

ferred to its own 1000 ml plastic container for pupation of the larvae. The life

span of each wasp was monitored.

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BIOLOGY 3

The pupae were sieved and placed in a 30 ml plastic cup for emergence. Each

plastic cup was cross-labelled to its adult female and day of exposure. Pupae in

each plastic cup were checked daily for emergence over 12 months, to establish

the minimum fecundity of each female and the duration between oviposition and

adult emergence. Emerged wasps were sexed and counted.

The above procedures were followed at the same time, using 10 virgin D.

kraussii females (three days old) for comparative purposes. Ten oviposition units

(20 B. tryoni larvae/unit) were also prepared daily as a control, to check the

survival rate of unparasitized B. tryoni.

Diel Patterns of Emergence and Ovipositional Activity—To determine the diel

pattern of D. kraussii emergence, a set of parasitized B. tryoni puparia, on the

verge of eclosing, were placed in a 250 ml plastic container. At the anticipated

time of peak emergence, the puparia were checked hourly through three consec-

utive days and the number and sex of D. kraussii that emerged each hour was

recorded. Preliminary tests had shown no emergence during the scotophase.

To check emergence patterns in the field, laboratory-parasitized B. tryoni pu-

paria (prepared as above) were held in 30 ml plastic cups (about 50 puparia/cup)

each of which was placed on the bottom of a 1000 ml plastic container. The lid

of each large container was cut to allow another 30 ml plastic “‘catching”’ cup to

be inserted through the hole, inverted, so its base protruded to the outside. The

large outer containers were covered with aluminium foil to reflect light. Only the

30 ml plastic “‘catching’’ cups received light, to which the D. kraussii adults were

attracted. The apparatus was placed in the shade of a Brazilian cherry tree at St

Lucia, Brisbane, on 2 Feb 1993 at 23:00 h and removed three days later at 20:00 h.

The “catching”? cups were checked hourly between 04:30 h and 18:30 h and the

D. kraussii that had emerged were removed, sexed and counted.

To assess the influence of day and night conditions on levels of ovipositional

behaviour, 15 D. kraussii females, three days old, were each exposed to a different

conspecific male, one day old, for mating (one pair/mating unit). After mating,

females were transferred to perspex cages (five females/10 <X 10 X 15 cm cage).

An oviposition unit (100 B. tryoni larvae/unit) was placed in each cage for the

light period 07:00 h to 17:00 h and another was substituted between 19:00 h and

05:00 h (dark period), in a constant environment room (conditions above). The

procedure was repeated with new oviposition units for three consecutive photo-

phases and scotophases. After exposure, each oviposition unit was transferred to

its own 1000 ml plastic container to await pupation of the flies. The numbers of

B. tryoni and D. kraussii adults that resulted from oviposition during the light

and dark exposures were sexed and counted.

To establish whether mated females that had never oviposited during the pho-

tophase would oviposit during the scotophase, a 2nd test was conducted following

the above procedures, but the times of exposure were reversed i.e., wasps were

initially exposed from 19:00 h to 05:00 h and only after that between 07:00 h to

17:00 h.

RESULTS AND DISCUSSION

Life Span, Developmental Duration and Reproductive Capacity—The mean

lifetime production of adult offspring by mated D. kraussii females (+ SE) was

111.7 (4 11.29, n = 10), and was not significantly different (¢ = —0.85, P =

4 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

ae FA Virgin

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AAA AAAS AAA Waa Ae AOD

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4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33

Days after eclosion

Figure 1. Number of offspring produced by virgin and mated Diachasmimorpha kraussii females

during each day of their lives. Their first day of exposure to hosts was their 4th day after eclosion.

The number (x + SE) of adults that emerged from the eggs deposited each day of exposure is given.

Ten replicates were run, but that dropped in the virgin treatment to 9 (day 30), 8 (d 31), 7 (d 33), 5

(d 34) and none were alive on day 35. The equivalent data for mated females is 8 (d 25), 7 (d 27), 6

(d 32), 3 (d 34) and O on day 35.

0.4141) from that of virgins (124.5 (+ 10.34, n = 10)). Although mated females

had a life span (27.6 + 4.55 days) significantly shorter than that of virgins (31.4

+ 1.96 days) (Wilcoxon two-sample test t = 0.0742, P = 0.0315), the mated

females produced more adult offspring per day (4.19 + 0.51, n = 10) than did

virgin females (3.97 = 0.33, n = 10) (Wilcoxon two-sample test t = 0.0514, P

= 0.0376). Offspring production peaked early (mean of 15—20/d on days 7-8 of

adult life), declined abruptly but somewhat erratically to steady low levels (3/day

or less) by day 20, and ceased on the last few days of adult life (Fig. 1).

The mean (+ SE) number of dead fruit fly puparia in the controls (10.7 +

0.48) was not significantly different from the number dying (without yielding a

parasitoid) after exposure to mated (11.2 + 0.56) or virgin (11.1 + 0.27) females

(Kruskal-Wallis x? = 0.96086, P = 0.6185, df = 2). Thus, ovipositor probing by

the wasps probably was not responsible for additional mortality.

Male offspring started to emerge (and achieved the maximum rate of emergence

per day) one day before their female siblings (Table 1). The male offspring pro-

duced by virgin females achieved maximum emergence on the same day as the

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BIOLOGY 5

Table 1. Duration until first emergence and maximum rate of emergence (days since parasitoid

oviposition) of offspring of mated (MF) and virgin females (VF) of Diachasmimorpha kraussii. The

number of offspring that emerged during the relevant time period is given in brackets. The number

of days taken to achieve 50%, 75% and 100% cumulative emergence is given.

Day of Cumulative emergence

(days)

Offspring First Maximum

Parent sex emergence emergence 50% 75% 100%

VF m 16 (1) 19 (408) 19 25 335

MF m 18 (10) 19 (53) 50 135 335

MF f 19 (3) 22 (122) 25 63 297

male offspring of mated females, but their first emergence was two days sooner

(Table 1). The overall emergence pattern (Fig. 2) of the male offspring produced

by mated females was not significantly different from that of their sisters (Kol-

mogorov-Smirnov two-sample test, D(332735) = 0.037, x? = 1.278, P > 0.05), but

ie males, virgin mother (n=1240)

70 males, mated mother (n=332)

L] females, mated mother (n=785)

Frequency (%)

16 - 25 —WNAAAAAAAAAAAANSAAAAAA A

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ee Pipe all ee lCl6[UlU LUC ell eC lle COt~*«C YD:

ae 8 ¢ 8 8 8 #8 © 6 © @

~_ , jF

Emergence time (days after oviposition)

Figure 2. Frequency diagram to illustrate the temporal pattern of adult emergence in Diachasmi-

morpha kraussii, calculated from the time of oviposition of each individual. Data are presented sep-

arately by sex and according to the mating status of their mother.

6 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

was so in comparison with the males produced by virgin females (Kolmogorov-

Smimov two-sample test, Di249332) = 0.393, x? = 0.084, P < 0.05). Specifically,

many more of the males from virgin mothers emerged early (days 16—25), and

relatively more males from mated mothers emerged late (126 days or more). The

reason for this large disparity in emergence patterns between males produced by

virgin and mated females is unclear, and we know of no other case of such a

discrepancy in the literature. The analysis of cumulative emergence showed that

male offspring in general took longer than their female siblings in achieving

cumulative emergences of 50, 75 and 100% (Table 1).

Extended emergence by some brood members of opiine species has been re-

corded previously. In D. longicaudata cultured at 25° C, emergence time varied

from 18 days to a year or more (Snowball et al. 1962, Clausen et al. 1965), which

suggests that the staggered and protracted emergence times within broods ensures

that at least some offspring of each female survives unfavourable conditions that

might arise unpredictably.

Mated females produced significantly more females than males. The average

sex ratio (+ SE) of all offspring produced over the entire life of mated females

was 0.28 (+ 0.03, n = 10) (proportion males). The overall sex ratio of all progeny

was 0.297 (n = 1117). Brood sex ratio was positively correlated with time of

offspring emergence. Based on cumulative emergence, brood sex ratio increased

with time (Pearson and Spearman Correlation Coefficients, r = 0.85297 and

0.90885, respectively, P = 0.0001).

Virgin females (50%, n = 10) also produced female offspring, but in very low

numbers (1 female offspring/virgin female producing a female).

Diel Patterns of Emergence and Ovipositional Activity.—In the laboratory,

most male and female wasps emerged during the first three to four hours after

the lights came on and the number emerging gradually declined towards the end

of the light period (Fig. 3). No wasps emerged during the scotophase (from 18:00

h to 06:00 h).

Under field conditions, both male and female wasps started to emerge from

O500h. The emergence rate gradually increased towards midmorning and declined

in the afternoon (Fig. 4). No wasps emerged between 18:00 h and 05:00 h. Emer-

gence times for parasitoids of both sexes under laboratory conditions were not

significantly different from those recorded in the field (Kolmogorov-Smimov two-

sample test on data for males: Dis5560) = 0.252, x? = 10.988, P > 0.05, and for

females: Dyo490) = 0.208, x? = 11.535, P > 0.05). The asynchronous pattern of

adult emergence may be related to individual physiology.

We found no published records on the time of day that other opiine parasitoids

emerge, but another braconid, Bracon hebetor Say, also emerged asynchronously

under constant temperature and light intensity (Antolin & Strand 1992). Emer-

gence time may not be constrained by strong natural selection, as these solitary

braconids are not geared to mating at their natal site (see Antolin & Strand 1992,

Rungrojwanich & Walter 1999), even though their hosts may be somewhat

clumped, as in quasigregarious parasitoid species (Nadel & Luck 1992).

Mated female D. kraussii oviposited under both light and dark conditions in

the laboratory (Table 2). The pattern of offspring production was influenced pri-

marily by whether photophase or scotophase conditions prevailed (Tables 2 and

3). More than 60% of offspring were produced during the photophase, regardless

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BIOLOGY 7

6 7 8 9 1

[] Females

Hi Males

Number of adults

1 1

Time of day (hours)

1 1

Figure 3. Total number of Diachasmimorpha kraussii that emerged hourly during the photophase

under laboratory conditions.

1 6 17 18

3 4 5

of whether the parent females were exposed first to scotophase conditions or

initially to photophase conditions.

The diel emergence data (Figs. 3 and 4) suggest D. kraussii is diurnal in the

field, but the oviposition data suggests that oviposition may also take place at

night, but at lower levels than in the day (Table 2). This has yet to be confirmed.

Field evidence from other opiines is contradictory, but since most of it derives

from light trapping, its implications for interpreting patterns of ovipositional ac-

tivity are still unclear. Light traps, operated continuously for four years in Ma-

laysia (1986-1989), attracted no fruit flies nor any opiine parasitoids (S. Vijay-

segaran, personal communication), which does support our contention. Both sexes

of another opiine, Psyttalia incisi (Silvestri), were attracted to a light trap set up

in India, with most individuals caught between 19:00 h and 22:00 h (Banerjee

1989), so the species may be active nocturnally, or both nocturnally and diurnally.

Different fruit fly parasitoid species may therefore respond differentially to light

traps and may be active at different times of the day or night.

The only information on nocturnal oviposition by opiines in the field involves

unquantified observations on F.. arisanus. Females have been recorded ovipositing

at night under laboratory conditions (van den Bosch & Haramoto 1951; G. M.

Quimio, personal communication). Fopius arisanus females were seen ovipositing

8 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

[] Females

Hi Males

Number of adults

Time of day (hours)

Figure 4. Total number of Diachasmimorpha kraussii that emerged hourly under field conditions.

Table 2. Numbers of adult offspring of Diachasmimorpha kraussii produced by six groups (=

replicates) of mated D. kraussii females. The females in three of the groups were first presented with

hosts during the photophase (labelled “‘photophase first’’) and the other three groups were first exposed

to hosts during the scotophase (“‘scotophase first”’). All wasps were exposed to new hosts at the change

of the light phase, and this continued for three days.

Photophase first Scotophase first

Rep Day Ph Sc Ph Sc

1 1 37 25 48 15

2D 31 16 45 17

3 34 16 39 18

Z 1 2d: 15 39 2a

Z 38 14 45 20

3 26 19 38 21

3 1 25 27 28 14

Zz 31 19 31 19

3 28 20 24 15

Total 277 171 337 166

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BIOLOGY 9

Table 3. Results of a three-way ANOVA testing the influence of day of exposure, light sequence

(scotophase first treatment or photophase first treatment) and light conditions at the time of oviposition

(scotophase or photophase). The test was performed on the untransformed raw data in Table 2. Trans-

formation of the data did not affect the outcome.

Sum of Mean

Factors df squares square F P

Day (A) 2 45.167 22.583 0.635 0.5385

Light sequence (B) 1 84.028 84.028 2.363 0.1373

AXB 2 12.056 6.028 0.170 0.8451

Light condition (C) ] 2131.361 2131.361 59.945 <0.0001

AXC 2 70.056 35.028 0.985 0.3880

BXC 1 117.361 117.361 3.301 0.0818

AXBXC 2 29.389 14.694 0.413 0.6661

in the field during nocturnal observations on fruit fly activity (R. A. I. Drew,

personal communication).

Australian Distribution, Host Flies and Host Plants—Diachasmimorpha

kraussii is distributed in the Northern Territory, Queensland and New South Wales

(Fig. 5). There are no records further south than New South Wales although its

best-known host, B. tryoni, occurs regularly in Victoria and sporadically in South

Australia (White & Elson-Harris 1992). The list of host plants and fruit flies with

NORTHERN

TERRITORY

WESTERN

AUSTRALIA

SOUTH

AUSTRALIA

Figure 5. Distribution of Diachasmimorpha kraussii in Australia.

10 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Table 4. List of host plants and host fruit flies (all in the genus Bactrocera) with which Diachas-

mimorpha kraussii has been associated. The number of records, from museum specimens, for each

association is also given. The fly names are from the labels and all are still current (Norrbom 1998).

Host plants and host fruit flies are not correlated across the table.

Host plant Host fly

Species No. records Species No. records

Eugenia uniflora regular* tryoni 15; regular*

Psidium guajava 6 Jarvisi 5

Solanum mauritianum neohumeralis 3

Eriobotrya japonica cacuminata 2

Mangifera indica aquilonis 1

Terminalia catappa halfordiae 1

Juglans regia kraussii 1

Planchonia careya melas 1

Prunus persica murrayi 1

Solanum seaforthianum visenda 1

Morus nigra

Musa spp.

Nauclea orientalis

Persea gratissima

Pyrus communis

Terminalia melanocarpa

Semecar pus australiensis

Cherry guava

Myer lemon

RS SR eB ee eS ONW WW ARAL A

* Diachasmimorpha kraussii was found in association with Bactrocera tryoni in Eugenia uniflora

fruits regularly and in large numbers, when E. uniflora was fruiting in Brisbane (1991-1994).

which D. kraussii has been associated in the field is shown in Table 4 (which

includes the records of May & Kleinschmidt (1954)). Six species of flies recorded

here as hosts had not been listed by Clausen et al. (1965). The date and place of

collection and the collectors are documented in full by Rungrojwanich (1994).

ACKNOWLEDGMENT

We thank Bob Wharton who kindly identified the parasitoid specimens. Sincere

thanks for the kind favours of Eddie Hamacek who supplied us with fruit flies,

Laurie Jessup who identified plant specimens, Joan Hendrikz, Laraine Law and

Andrew Loch who advised on statistics and Tony Clarke, Jenny Beard and Stefan

Schmidt who commented on the manuscript. The research was supported by an

AusAID scholarship to K.R. and by the Department of Entomology, The Uni-

versity of Queensland.

LITERATURE CITED

Antolin, M. F & M. R. Strand. 1992. Mating system of Bracon hebetor (Hymenoptera: Braconidae).

Ecol. Entomol., 17: 1-7.

Banerjee, T. C. 1989. Nocturnal periodicity of natural population of a braconid fly, Opius incisi: size

and distribution pattern. Indian J. Entomol., 51: 440-449.

Clausen, C. P, D. W. Clancy & Q. C. Chock. 1965. Biological control of the oriental fruit fly (Dacus

dorsalis Hendel) and other fruit flies in Hawaii. USDA Tech. Bull., 1322: 1-102.

Fullaway, D. T. 1951. Review of the Indo-Australiasian parasites of the fruit flies (Tephritidae). Proc.

Hawaiian Entomol. Soc., 14: 243-250.

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BIOLOGY 11

Heather, N. W. & R. J. Corcoran. 1985. D. tryoni. pp. 41-48. In Singh, P. & R. E Moore (eds.).

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438.

Received 29 Nov 1998; Accepted 10 Jul 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 12-23, (2000)

THE AUSTRALIAN FRUIT FLY PARASITOID

DIACHASMIMORPHA KRAUSSHI (FULLAWAY): MATING

BEHAVIOR, MODES OF SEXUAL COMMUNICATION

AND CROSSING TESTS WITH D. LONGICAUDATA

(ASHMEAD) (HYMENOPTERA: BRACONIDAE: OPIINAE)

K. RUNGROJWANICH! AND G. H. WALTER

Department of Zoology and Entomology, The University of Queensland,

Brisbane, Queensland 4072, Australia

Abstract—We describe the mating behavior of Diachasmimorpha kraussii for the first time, and

confirm with cross-mating tests the separate species status of D. kraussii and D. longicaudata.

Flight cage experiments suggest that mating takes place on foliage and that a distance attractant

pheromone is secreted by the females, and perhaps also by the males. The most obvious aspect

of the sexual interaction between males and females is the wing vibration performed by males

in the nearby presence (about 1 cm) of a conspecific virgin female. Wing vibration produces an

acoustic signal critical to mating success, for wingless males could seldom mate. Experimental

manipulations demonstrate that males vibrate their wings in response to a chemical associated

with the female, but not present in males. The chemical appears to be associated with the cuticle,

as it is present (as demonstrated by male behavior) in recently-killed females, and it can be

stripped from these females with acetone. The interaction proceeds only if the female is receptive

(starting 6-48 h after emergence) and when she adopts a particular stance. Receptive females

stand still, fold both pairs of wings over the abdomen, hold their antennae back together over

their wings and allow males to mount. Males continue tapping their antennae on the females’

thoraces while intromission takes place. The mating sequence of D. longicaudata is generally

similar to that of D. kraussii, but individuals of the two species did not mate in small cages,

which confirms their species status. In crossing tests all males vibrated their wings, indicating

that the female’s cuticular chemicals are similar across species. No females in mixed pairs

assumed the receptive stance, suggesting the acoustic signals differ across species.

Key Words.—Diachasmimorpha kraussii, Diachasmimorpha longicaudata, parasitoid, Braconi-

dae, Tephritidae, acoustic signal, wing vibration, monandry, pheromone.

Understanding mating behaviour is central to accurate interpretation of the spe-

cies limits of sexual organisms. This is particularly true when cryptic (sibling)

species compelexes are suspected (Fernando & Walter 1997). The sexual com-

munication mechanism, or Specific-Mate Recognition System (SMRS), comprises

several steps, each of which serves a function subservient to the ultimate function

of achieving fertilization (Paterson 1985). Although particular aspects of the

SMRS of many insect species are well known, in only very few species have

attempts been made to identify each step in the sequence (see Matthews 1975,

Field & Keller 1993, Abeeluck & Walter 1997). An understanding of all steps is

critical to assessment of species limits and the species status of different popu-

lations (Fernando & Walter 1997). Here we describe research on the parasitic

wasp Diachasmimorpha kraussii (Fullaway) that allows us to develop a diagram-

matic model of the communication modes associated with each step in the entire

mating sequence of this species.

' School of Agricultural Extension and Cooperatives, Sukothai Thammathirat Open University, Pak-

kred, Nonthaburi 11120, Thailand.

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BEHAVIOR 13

Wasps in the genus Diachasmimorpha parasitize tephritid fruit flies. Several of

the species resemble one another so closely that the morphological features used

to distinguish them tend to grade into one another. For example, the Australian

species D. kraussii resembles D. longicaudata (Ashmead), an Asian species, and

is distinguished only by the combination of pale coloration and an unsculptured

second abdominal tergite (Wharton & Gilstrap 1983). Such characters are often

subtle in that they may be open to subjective interpretation, especially when the

named species have allopatric distributions. In such cases, behavioral confirmation

of species limits helps to provide confidence in the designated morphological

features actually being diagnostic.

Understanding the mating behavior of opiine parasitoids may also yield prac-

tical benefits because of their value to biological control. Several species of opiines

were transported to Hawaii for rearing and release against Bactrocera dorsalis

(Hendel) and Ceratitis capitata Wiedemann, between 1947 and 1953, but not all

could be successfully reared (Clausen et al. 1965). A problem with some of the

Species was the low incidence of successful mating, so a preponderance of male

offspring was produced prior to those cultures dying out. Understanding the re-

quirements for mating of these species is therefore important for successful mass

rearing (Purcell 1998). Although D. kraussii was ‘‘eventually propagated’”’ on C.

capitata and released prior to 1953, it did not establish (Clausen et al. 1965).

Whether there were problems in getting wasps of this species to mate was not,

however, mentioned.

To date, nothing has been published on the mating behavior of D. kraussii. Our

aim in investigating the SMRS of D. kraussii is to provide a first model of the

complete mating sequence of an opiine parasitoid. We thus provide a basis for

comparison of mating behavior across host associated populations and across al-

lopatric populations of Diachasmimorpha. We also use the information obtained

to investigate intersexual interactions between D. kraussii and D. longicaudata

individuals.

MATERIALS AND METHODS

The D. kraussii colony was established with about 35 pairs reared from B.

tryoni (Froggatt) puparia derived from Brazilian cherries (Eugenia uniflora L.)

collected in St Lucia, Brisbane, in November 1990. The identification of the wasps

was confirmed by R. A. Wharton and specimens have been deposited in the

collections of Texas A&M University, College Station and The University of

Queensland. Diachasmimorpha longicaudata was obtained from the culture held

at the Tropical Fruit and Vegetable Research Laboratory of the United States

Department of Agriculture, Honolulu, Hawaii. Both parasitoids were reared on B.

tryoni and the maintenance of host and parasitoid colonies is described elsewhere

(Rungrojwanich & Walter 1999).

Unless otherwise stated, the males and females used in experiments were one

and three day old virgins, respectively, and observations were made in a “‘mating

unit” made up of two glass tubes (2.5 cm diam. X 4.5 cm), each with one end

covered with fine muslin and the other end left open. Only one pair/mating unit

was observed at a time, for no more than 10 min.

Description and Duration of Mating Behavior.—Observations were conducted

to establish which sex displays observable signalling behaviors, the nature of

14 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

those behaviors and the duration of signalling and copulation. Before the wasps

were brought together, each insect was allowed to settle for ten min in its own

tube. The open ends of the tubes were then brought together to allow the male

and female to approach one another. Time spent on wing vibration by males and

in copulation were recorded (in seconds) with a stop-watch (m = 20 pairs). Ob-

servations were conducted at 25 + 1°C and 60 + 5% R.H.

Mate Location: Flight Cage Observations.—The site where mating partners

usually meet in nature was assessed indirectly. Nine flowering and/or fruiting

guava trees (2 m tall, each in a 30 cm diameter pot, all with young fruits and

some with honeydew-secreting scale insects) were placed 45 cm apart in a 180

x 180 X 200 cm fly-screen cage in a glasshouse. Conditions varied from 22° C

to 29° C and 60% to 96% R.H.., typical of summer weather conditions in Brisbane.

All observations were conducted between 09:00 h and 17:30 h.

In the first set of observations, six D. kraussii females were released into the

flight cage and their behavior was observed and recorded for 15 min before a

single male was introduced. The time taken before a mating pair came together

was measured and all behaviors and interactions were recorded. Once mating had

taken place, the mating pair was removed and a new female released to maintain

numbers. Another male was released 15 min later. Nineteen females and 14 males

were ultimately tested.

In a second set of identical observations six virgin males were released first,

followed by a single female. Similar observations to those described above were

made and the wasps were treated in the same way. Totals of 25 males and 20

females were observed.

Stimulus for Male Wing Vibration.—A series of experiments was conducted

with dead females (at 25 + 1° C and 50 + 2% R.H.) to determine (i) whether

wing vibration by males is stimulated by chemicals associated with the female,

(ii) if stimulatory chemicals are cuticular or are under active control from within

the insect and (i111) whether visual cues are involved. Unless otherwise stated, a

new set of one day old males was used in each test. Each replicate was observed

for no longer than 10 min. The occurrence of male wing vibration and copulation

attempts were noted and the duration of intromission was recorded.

Initially, a set of controls was run to determine whether the mating units and

acetone (used later as a solvent: see below) would, in themselves, have any effect

on the behavior of D. kraussii males. Nine males were placed in clean mating

units (one/unit) for 10 min of observation, then each one was placed for 10 min

in a mating unit to which had been added 0.5 ml acetone that had been allowed

to evaporate. Males were also tested for their responses to other males.

The distance between the wasps of each living pair (n = 22) was measured

when the male started vibrating his wings (see below) in response to the female’s

presence. Males were similarly assessed against virgin females that had recently

been killed by freezing (— 10° C for 15 min). These same dead females were then

soaked in 15 ml of acetone for 15 min and specimens were air dried for 1 h

before again being exposed to males.

To test whether males would recognize females by their visual appearance, the

Ovipositors of females were removed before exposure to males (as ovipositor

protrusion is the only general observable way in which females differ from con-

specific males). The females with ovipositors removed were soaked in acetone

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BEHAVIOR 15

after the experiment and air dried for another exposure to males. Females recently

killed by freezing were cut into two parts (head + thorax and abdomen) to in-

vestigate whether the factor that influences the behavior of conspecific males is

specific to body part.

To investigate whether an internally-derived chemical (e.g., from a pheromone

gland reservoir) influences male behavior, dead females were crushed on the side

of mating units (two/unit), with a glass rod. The carcasses were then removed

and a male was placed in each mating unit.

The duration for which dead females would remain attractive to conspecific

males was investigated by exposing the same set of dead females (n = 10) toa

new set of males daily, until no male responded to them.

Role of Male Wing Vibration.—The functional significance of male wing vi-

bration was assessed by exposure of females to wingless males (at 27° C and 65

+ 2% R.H.). Males (n = 16) were immobilized at —10° C for five min before

each wing was cut off, just above the base. To establish whether the cold treatment

alone would affect the behavior of males negatively, a set of controls was run

first. Ten males were cold immobilized as above, but their wings were not cut off

before exposure to females.

Female Premating Period and Polyandry.—The age of females (all virgins)

used to measure the premating period varied from six h after emergence to 25

days old, whereas all males were one day old. Individuals in each pair (n = 208

pairs altogether) were tested only once even if they did not mate on first exposure.

To establish whether D. kraussii females would mate more than once, they (n

= 17) were each exposed to a different conspecific male. When mated, each

female was transferred to a 10 X 10 X 15 cm perspex cage. Honey solution was

provided as food. An “oviposition unit’? (Rungrojwanich & Walter 1999) con-

taining about 100 B. tryoni larvae was provided daily. Each of these previously-

mated females was later exposed to another one day old conspecific male at seven,

15 and 20 days after the first mating. Because of the early death of some females,

only six and three were alive on days 15 and 20, respectively.

Is Mating Strictly Diurnal ?—To assess whether mating by D. kraussii is con-

fined to daylight, the mating success of laboratory-reared wasps (n = 14) was

observed at night under field conditions (Sir John Chandler Park, Indooroopilly,

Brisbane, 21:00 h, 3 Feb 1993). Light intensity was measured with a system

exposure meter (LUNASIX 3). During the experiment it was equal to 0.35 lux

or 0.032 fc, and ambient conditions were 25.5° C and 77% R.H. Males in pairs

that did not mate were separated from the females for immediate transfer to an

illuminated laboratory for re-testing with each other.

Cross-mating Experiment.—The sexual interactions between D. kraussii and D.

longicaudata individuals were tested in mating units. The number of mixed pairs

that mated successfully (out of 20 pairs in each reciprocal cross) was recorded,

as was the level of interaction between the sexes. A set of control crosses (n =

20 pairs of each species) was also observed. Each pair was placed together in a

mating unit for a maximum of 10 min, or until mating was completed. Diachas-

mimorpha kraussii males and females used in the experiments were one and three

days old, respectively, whereas both sexes of D. longicaudata were three days

old. Each wasp was virgin and used only once, and experiments were conducted

at 25 + 1° C and 60 + 5% R.H.

16 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

RESULTS

Description and Duration of Mating Behavior.—All D. kraussii males (n =

16) showed a distinct behavioral sequence in the nearby presence (about 1 cm)

of a conspecific virgin female. Each male made a protracted series of wing vi-

brations, which was continued after he mounted a receptive female. Males also

tapped their antennae on the female’s thorax during intromission and insemination.

A few males (12.5%, n = 16), stopped wing vibration and antennal tapping before

intromission was completed.

Receptive D. kraussii females stood still while the male vibrated his wings.

Simultaneously, the females folded both pairs of wings over their abdomens and

held their antennae back together over the wings (100%, n = 16). Other obser-

vations (see below) showed that if a female was not ready to mate, she continued

to walk around and sometimes vibrated her wings. Her antennae were kept apart

and pointed forward. Intromission was never successful when a male tried to

copulate with a female that had not assumed the appropriate posture.

The mean (+ SE) time spent in copula was 23.7 (+ 3.19) sec (nm = 20), and

in precopulatory courtship it was much less, at 8.8 (= 1.29) sec (n = 20).

Mate Location: Flight Cage Observations——When virgin females were re-

leased into the cage before males, they flew directly to the plants and each landed

on the upper surface of a leaf (1-2 m above ground). They fed on honeydew,

preened and rested near the leaf apex. Only five females (26.3%; n = 19) took

short flights to nearby leaves (8—10 cm away).

When males were then released into the cage singly, they also flew to the upper

surface of a leaf, near the apex, and fed, preened and rested. When males did fly

again, which they did sporadically, they flew in a zig-zag pattern at a height of

1—2 m above the ground and they flew around the tree canopy. In most matings

that ensued (64%, n = 14) males thus approached females. In only four cases did

females (29%, n = 14) land on a male’s leaf, where mating took place. The male

and female of the remaining pair landed simultaneously on a leaf before mating.

In those cases in which males flew to a female, the time from release of the male

to contact with a mate varied from 18 to 90 min (mean + SE = 43.7 + 7.77, n

= 9), which was not significantly different (¢ = —0.051, P = 0.96) from the time

between release of a male and a female approaching that male (42.8 (+ 9.68, n

= 4) min)). Most matings (86%, n = 14) took place on the upper leaf surface,

and mating never took place below a height of 1 m.

When males were released before females they also flew to the upper surface

of a leaf about 1.5—2 m above the ground and fed on honeydew before preening,

resting and flying. Males flew much longer distances than females and they flew

in a zig-zag pattern around the tree canopy. Two males landed on leaves where

other males were resting and performed typical premating wing vibrations. They

did not try to copulate and soon flew away.

On being released individually into the cage with males, females landed on the

upper surface of a leaf 1-2 m above the ground, where they fed, preened and

rested. Some females (35%, n = 20) flew in a zig-zag pattern around the tree

canopy and landed on leaves where males were resting. The males immediately

vibrated their wings and all of these pairs then copulated. The time females spent

in flying before landing alongside a mate varied from 5 to 25 min (mean + SE

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BEHAVIOR 17

Table 1. Response of Diachasmimorpha kraussii males to females recently killed by freezing (con-

trol) and to freeze-killed females soaked in acetone to remove cuticular lipids and with their ovipositor

removed to alter their visual appearance.

Wing Mounting

n vibration % attempted %o

Control 22 22 100 22 100

Acetone soak (Ac) 22 15 68 11 50

Ovipositor off (Oo) 26 24 92 15 58

Ac + Oo 26 6 23 3 12

= 12.9 + 3.05, n = 7). Most females (65%, n = 20) did not fly again after their

initial landing, and they were located by males in zig zag flight. The time between

female release and a male landing alongside a female varied from 3 to 129 min

(mean + SE = 36.6 + 9.36, n = 13), and was not significantly different (tf =

—1.813, P = 0.087) from the time preceding mate finding by females. Again,

mating usually took place on the upper surface of a leaf (95%, n = 20), and

never took place lower than 1 m from the ground.

Combining the results across experiments shows that males located females

much more frequently than females located males (67% vs 33%, n = 33). The

average time for males to locate a female did not differ across experiments (t =

—0.54, P = 0.59). Those females that located males did so significantly faster

when females were released after males (¢t = —3.68, P = 0.005), but sample sizes

were low.

The Stimulus for Male Wing Vibration—No males showed courtship behavior

when left alone in clean mating units (7 = 9) or when exposed to a container

that had held 0.5 ml acetone (n = 9). Also, no males vibrated their wings in

response to males recently killed by freezing (7 = 10) or to males soaked in

acetone after being killed by freezing (n = 10). By contrast, males presented with

females recently killed by freezing, all showed the usual pattern of wing vibration

(i.e., several series of wing vibrations) and each mounted the female (n = 22).

Whereas males vibrated their wings in response to living females (n = 22) ata

mean (+ SE) distance of 10.2 (+ 0.064) mm for living females, they did so at a

significantly greater distance from dead females (mean + SE = 13.2 + 0.094, n

= 20) (Wilcoxon two-sample test, t = 0.0153, P = 0.011).

When males were provided with recently frozen females that had been soaked

in acetone, 68% of them vibrated their wings, but only 50% mounted the females

(Table 1). Almost all males vibrated their wings in response to females recently

killed by freezing and with their ovipositor removed, but almost half of them did

not mount the females (Table 1). After the females without ovipositors had been

soaked in acetone, the number of males that responded was reduced drastically;

only a quarter of them vibrating their wings and 12% mounting (Table 1).

Twenty per cent of males vibrated their wings when placed in vials containing

extracts from crushed females (n = 10), but they made only one brief burst of

wing vibration. All males vibrated their wings and all mounted the female if only

her head and thorax was provided (n = 10). Although all males vibrated their

wings when presented with just the abdomen of the female, only two tried to

mount (n = 10).

18 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

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123 45 6 7 8 9 10111213 14 15 16 17 18 19 20 21 22 23 24 25

Days

Figure 1. Positive behavioural response of Diachasmimorpha kraussii males to dead females. The

same set of dead females was used throughout whereas a new set of males (nm = 10) was exposed to

the females each day. Only one pair of wasps was held in each vial.

When a set of dead females (one per vial) was exposed each day to a newly-

emerged male (one per female), all males vibrated their wings each day for the

first 18 days of exposure (Fig. 1), but the number responding declined from day

19, and by day 25 no males responded. The number of males attempting to mate

varied considerably (Fig. 1).

Role of Male Wing Vibration—When males’ wings were removed, the per-

centage that achieved intromission was reduced drastically. No females showed

receptive behavior in response to wingless males, but just walked and preened.

Nevertheless, three of the wingless males (18.8%, n = 16) were able to mate.

The other 13 walked around, but did not attempt copulation. By contrast, most

control males (80%, n = 10) mated successfully, despite their prior immobilization

in a freezer. The two males that did not mate successfully nevertheless displayed

typical wing vibration in response to the presence of the virgin females.

Female Premating Period and Polyandry.—About a quarter of females mated

as soon as six hours after eclosion, and all had mated by the time they were two

days old (Fig. 2). Regardless of age, all virgin females mated when exposed to

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BEHAVIOR 19

HM Proportion mating (%)

30 28 1010 10 10 7 10 9 2 19 10

100

Proportion mating (%)

0123 4 5 6 7 8 9 1011 12 13 14 15 16 17 18 19 20 21 22 23 24 25

Age (days)

Figure 2. The percentage of virgin female Diachasmimorpha kraussii of different ages that mated

within 10 minutes on first exposure to conspecific males. The number of wasps observed on each

occasion is presented above each bar.

males (Fig. 2). None of the females that were mated one day after emergence

mated again, whether at seven days (n = 17), 15 days (n = 6) or 20 days (n = 3).

Is Mating Strictly Diurnal?—Under low light intensity (0.35 lux) only four

pairs (28.5%) of D. kraussii mated (n = 14). The behavior of these males and

females was normal in all respects. The males in the ten pairs that did not mate

initially in the field did not vibrate their wings in response to female presence.

However, they did so in the laboratory, where light intensity was 11 lux, and all

of them mated.

Cross-mating Experiment.—Superficially, the mating behavior of D. longicau-

data resembles that of D. kraussii, but durations were not measured. In the cross-

mating tests, when D. kraussii males (n = 20) were exposed to D. longicaudata

females, they vibrated their wings in the same way as when exposed to D. kraussii

females, but none attempted to mate (Table 2). When D. longicaudata males (n

= 20) were in the same mating unit as D. kraussii females, all of the males

vibrated their wings and 60% of them attempted copulation despite the females

20 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Table 2. Results of a cross-mating test between Diachasmimorpha kraussii (K) and D. longicaudata

(L). Control crosses were conducted simultaneously. All wasps were virgin, males were one day old

and females two days old. n = 20 pairs in each test.

% Males involved in:

Wing Mounting

Test crosses vibration attempts Intromission

@ KX 9K 100 100 100

(oe ae Ca! I 100 100 100

e-Kex OL 100 0 0

Co hs Xee K 100 60 0

not taking up the usual posture that indicates receptivity. No females allowed a

non-conspecific male to copulate. When D. longicaudata males tried to copulate

with D. kraussii females, the females did not stand still but walked around and

sometimes vibrated their wings. Conspecific males and females in the control

crosses mated normally when they were placed together in a mating unit (Table 2).

DISCUSSION

The SMRS of D. kraussii is a sequence of steps involving olfactory, visual and

acoustic signals, as summarized diagrammatically in Fig. 3. This synthesis is

hypothetical, and is designed to help visualise the relationship among components

of the overall interaction, to identify areas for further research and to aid the

design of comparisons among populations.

Presumably the wasps do not mate at their emergence site because most males

emerge at least one day before females (Rungrojwanich & Walter 1999). Although

males could wait for female emergence, this is not usual in solitary species, or

even quasi-gregarious ones (Myint & Walter 1990, Nadel & Luck 1992). Also, a

considerable proportion (about 30%) of the brood emerged at irregular intervals

after the main emergence (Rungrojwanich & Walter 1999) and they would be less

likely to find mates at the emergence site. Possibly there is a particular part of

the environment to which both sexes would be attracted and where mating would

take place, as proposed for the pteromalid parasitoids Spalangia cameroni (Per-

kins) (Myint & Walter 1990) and Pachycrepoideus vindemiae (Rondani) (Nadel

& Luck 1992). In D. kraussii one or more particular host plant species may attract

males and females, or the environmental cue may be more general, perhaps any

tree with fruit attacked by fruit fly larvae. Observations in the field have shown

that males, sometimes in considerable numbers, may fly around the canopy of

fruiting host trees (brazilian cherry) during the day (M. K. Ross, personal com-

munication), presumably waiting for females to arrive.

A combination of visual and chemical signals seems to be used once the sexes

are in the same general area around a host plant. This is supported by the low

frequency of mating recorded at night. In the flight cage experiment, the most

common means of the sexes meeting was through males flying around the tree

canopy and landing on the leaf on which a female was resting, which implies that

visual and chemical cues are used for mate finding. Our behavioral observations

suggest the females release an attractant volatile. Because females sometimes

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BEHAVIOR 21

ADULT EMERGENCE

MALE FEMALE

ATTRACTION TO HOST TREE

RANDOM FLIGHT AROUND TAKES UP

CANOPY AND/OR »_ » POSITION

SETTLES ONLEAF ». ~~ ONLEAF

~~ Pheromones | -

_-~ & visual cues~ ~~

TOLEAF «4-7 ~s, STANDS

OF FEMALE y STILL

Cuticular chemicals my

APPROACHES FEMALE € -- - — - — & visual cues ¢ - - -— -’”

~s 4 Sound from

wing vibration- --------------- > STILL ON

LEAF

RECEPTIVE NOT RECEPTIVE

MOUNT FEMALE,

WING VIBRATING STANDS STILL,

CONTINUED, WINGS FOLDED

ANTENNAL TAPPING OVER ABDOMEN, VIBRATES

ON FEMALE’S ANTENNAE POINTED WINGS,

THORAX BACKWARDS WALKS AWAY

COPULATION

Figure 3. Postulated structure of the Specific-Mate Recognition System of Diachasmimorpha

Kraussii, an opiine parasitoid of fruit flies. See text for details of the type of evidence supporting each

interpretation. Solid lines indicate transitional steps in behaviour, with arrows indicating direction.

Dashed lines indicate signals between males and females. Arrows indicate the direction in which the

signal acts. Bidirectional arrows imply uncertainty of signal direction.

ae, THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

homed in on males, both sexes may do so. Possible sex pheromones in opiines

have been investigated only in males (Williams et al. 1988).

Once the sexes are in close proximity (about 1 cm) the male responds to the

female with acoustic signals associated with wing vibration, as recorded for the

congeneric D. longicaudata (Sivinski & Webb 1989). Whether the signals are air-

borne or vibrational (see Field & Keller 1993) has yet to be established for these

opiines. The stimuli for wing vibration are relatively stable (for about 25 days:

Fig. 1) chemicals associated with the female cuticle. At this stage the female, if

receptive, stands still and adopts a characteristic posture while the male approach-

es her. Again, visual stimuli seem to be a cue for the male to locate the mating

partner with whom he is interacting. The typical posture that the receptive female

adopts is not critical to the recognition process itself, because the male tries to

copulate even with severed parts of a female (head + thorax), and also with D.

longicaudata females that do not adopt such a posture in the presence of D.

kraussii males (Table 1).

No information is available on the role of tactile stimuli during courtship,

mounting and copulation. Presumably tactile stimuli come into play only after

males have actually mounted their mating partners. Some tactile stimuli are as-

sociated with overt behaviour. For example, the male taps his antennae on the

thorax of the female during copulation. Other signals may be less obvious. For

example, the position of the male’s legs during intromission may be important in

keeping the female passive and willing to copulate.

Finally, we confirm with behavioral evidence that the separation of D. kraussii

from D. longicaudata on morphological grounds is accurate, and we predict from

the behavioral observations, reported in Table 2, that the cuticular chemicals of

the females of the two species will resemble one another closely, but the acoustics

of the males will be different across the species.

ACKNOWLEDGMENT

Special thanks to Bob Wharton for identifying specimens, Joan Hendrikz and

Laraine Law for statistical advice, staff of the Honolulu USDA who arranged the

D. longicaudata colony for us, and Tony Clarke, Andrew Loch and Stefan

Schmidt for comments on the manuscript. The research was supported by an

AusAID scholarship to K.R. and by the Department of Entomology, The Uni-

versity of Queensland.

LITERATURE CITED

Abeeluck, D. & G. H. Walter. 1997. Mating behaviour of an undescribed species of Coccophagus,

near C. gurneyi (Hymenopera: Aphelinidae). J. Hym. Res., 6: 92-98.

Clausen, C. P, D. W. Clancy & Q. C. Chock. 1965. Biological control of the Oriental fruit fly (Dacus

dorsalis Hendel) and other fruit flies in Hawaii. USDA Tech. Bull., 1322: 1-102.

Fernando, L. C. P. & G. H. Walter. 1997. Species status of two host-associated populations of Aphytis

lingnanensis (Hymenoptera: Aphelinidae) in citrus. Bull. Entomol. Res., 87: 137-144.

Field, S. A. & M. A. Keller. 1993. Courtship and intersexual signaling in the parasitic wasp Cotesia

rebecula (Hymenoptera: Braconidae). J. Insect Behav., 6: 737-750.

Matthews, R. W. 1975. Courtship in parasitic wasps. pp. 66-86. Jn Price, P. W. (ed.). Evolutionary

Strategies of parasitic insects and mites. Plenum Press, New York.

Myint, W. W. & G. H. Walter. 1990. Behaviour of Spalangia cameroni males and sex ratio theory.

Oikos, 59: 163-174.

2000 RUNGROJWANICH & WALTER: FRUIT FLY PARASITOID BEHAVIOR 23

Nadel, H. & R. E Luck. 1992. Dispersal and mating structure of a parasitoid with a female-biased

sex ratio: implications for theory. Evol. Ecol., 6: 270-278.

Paterson, H. E. H. 1985. The recognition concept of species. pp. 136-157. In McEvey, S. E (ed.).

Evolution and the recognition concept of species. The John Hopkins University Press, Balti-

more.

Purcell, M. F 1998. Contribution of biological control to integrated pest management of tephritid fruit

flies in the tropics and subtropics. Integ. Pest Manage. Rev., 3: 63-83.

Rungrojwanich, K. & G. H. Walter. 1999. The Australian fruit fly parasitoid Diachasmimorpha kraussii

(Fullaway): life history, ovipositional patterns, distribution and hosts (Hymenoptera: Braconi-

dae: Opiinae). Pan. Pac. Entomol., 75: 1-11.

Sivinski, J. & J. C. Webb. 1989. Acoustic signals produced during courtship in Diachasmimorpha (=

Biosteres) longicaudata (Hymenoptera: Braconidae) and other Braconidae. Ann. Entomol. Soc.

Am., 82: 116-120.

Wharton, R. A. & FE E. Gilstrap. 1983. Key to and status of opiine braconid (Hymenoptera) parasitoids

used in biological control of Ceratitis and Dacus s.]. (Diptera: Tephritidae). Ann. Entomol. Soc.

Am., 76: 721-742.

Williams, H. J.. M. Wong, R. A. Wharton & S. B. Vinson. 1988. Hagen’s gland morphology and

chemical content analysis for three species of parasitic wasps (Hymenoptera: Braconidae). J.

Chem. Ecol., 14: 1727-1736.

Received 29 Nov 1998; Accepted 10 Jul 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 24-27, (2000)

FORMOSOZOROS NEWI, A NEW GENUS AND SPECIES

OF ZORAPTERA (INSECTA) FROM TAIWAN

REN-FANG CHAO AND CHIN-SHENG CHEN*

Department of Biology, Tunghai University, Taichung, Taiwan 407, R.O.C.

e-mail: cschen @ mail.thu.edu.tw

Abstract—A new genus and species of Zoraptera (Insecta), Formosozoros newi NEW GENUS

and NEW SPECIES, is described from female specimens collected on Taiwan. Zoraptera is newly

recorded from Taiwan.

Key Words.—Insecta, Formosozoros, Zorotypidae, Zoraptera, Taxonomy, Taiwan.

The zorapterans are minute insects that live in moist tropical and warm tem-

perate forested habitats. Since the order was established by Silvestri (1913), only

30 living and 1 amber fossil species have been described (Poinar 1988, Hubbard

1990, New 1995). All these species are traditionally placed in the single genus

Zorotypus belonging to the single family Zorotypidae. Kukalova-Peck & Peck

(1993) erected 6 new genera, i.e., Brazilozoros, Centrozoros, Floridazoros, La-

tinozoros, Meridozoros, and Usazoros, for New World zorapterans on the basis

of characters of wing venation. When Silvestri (1913) first described and proposed

the name for the order Zoraptera, he thought that they were apterous insects.

However, adults may develop either with or without wings (Caudell 1920, Gurney

1938, Riegel 1987). Although we can treat the taxonomy of Zoraptera with the

characters of wing venation, these characters are known for very few species and

apterous individuals often cannot be placed in a genus (New 1995). All Old World

species of the order are currently retained in Zorotypus, pending discovered and

appraised of winged forms.

Recently, five specimens of Zoraptera were found in Taiwan. It is the first

record of the order in Taiwan. The tarsi are different from any other recorded

species in the world. The Taiwan species is therefore considered to be a new

genus of Zoraptera, which is described in this paper. Any additional records of

this poorly understood insect order are valuable in documenting its distribution

and diversity. All specimens are deposited in the Department of Biology, Tunghai

University, Taiwan.

FORMOSOZOROS NEW GENUS

Type species.—Formosozoros newi NEW SPECIES

Description.—Apterous female, similar in general appearance to Zorotypus. Epicranial suture weak-

ly developed. Antenna 9-segmented. Apical segment of labial palpus and maxillary palpus with elon-

gate, narrowed apex respectively. Hind tarsi 2-segmented; first segment and second segment almost

equal in length; first-segment with a row of 5-6 pairs of short thickened setae. Abdomen 1 1-segmented.

Cerci very long; apical seta absent.

Etymology.—A combination from the words Formosa and Zorotypus; mascu-

line.

Remarks.—The major diagnostic characters of this new genus to Zorotypus are:

* Correspondence/reprint request address.

2000 CHAO & CHEN: NEW ZORAPTERA FROM TAIWAN 25

Figure 1. Formosozoros newi, n. sp., dorsal view.

(1) the strongly contracted apex of the apical segment of labial palpus and max-

illary palpus; (2) first segment and second segment of hind tarsus almost subequal

in length; (3) the 5—6 pairs of short thickened setae in first-segment of tarsus; and

(4) the greatly enlarged cerci. Additionally, all known zorapterans have an apical

seta in cerci except Zorotypus longicercatus Caudell, 1927 (Caudell 1927) and Z.

palaeus Poinar, 1988 (Poinar 1988); the new genus also lacks apical seta in cerci.

A small first-segment of the tarsus is common in other living zorapterans and in

fossil species (Poinar 1988). The longer first-segment of the tarsus of Formoso-

zoros may be an advanced character for Zoraptera. As the specimens of Formo-

sozoros \ack winged individuals, we cannot discuss the evolutionary relationship

among genera by wing venation.

FORMOSOZOROS NEWI NEW SPECIES

(Figs. 1—6)

Apterous female.—Body length 2.88—3.14 mm (n = 3). Head: pear-shaped; length 0.57—0.58 mm;

width 0.50—0.52 mm. Eyes and ocelli none. Frons with 6 macrochaetae. Antenna 9-segmented; second

segment short; segment length (I-IX, mm) 0.12, 0.05, 0.18, 0.16, 0.16, 0.17, 0.18, 0.18, 0.19. Labial

3-segmented; apical segment with a nipple seta. Maxillary 5-segmented; apical segment with a nipple

seta. Thorax: pronotum breadth longer than length (0.34 X 0.25 mm). Hind legs: femur strongly

expanded, length 0.37—0.38 mm, width 0.25—0.26 mm, with 3 posterior spines on the inner margin;

tibia with a row of 5 short thickened setae and 1 short apical spur; first-segment of tarsus 0.18 mm

in length, with a row of 5—6 pairs short thickened setae; second-segment length 0.17 mm, without

any thickened setae but with 2 claws. Abdomen: tergite I-X with 2 medial posterior macrochaetae

26 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Figures 2-6. Formosozoros newi, n. sp. 2. Antenna; 3. Hind leg; 4. Mandible; 5. Maxilla; 6.

Cercus.

and 2 lateral posterior macrochaetae; tergite XI small. Sternite I-IV with 4 posterior macrochaetae;

sternite V-IX with 6 posterior macrochaetae; sternite X—XI macrochaetae absent. Cerci long, length

0.41—0.43 mm, with 3—4 preapical long setae, apical seta absent. 2 spermathecae, connected by a long

slender duct.

Male and larva—Unknown.

Material examined.—Holotype ¢, Taiwan, Hualien Hsien, Nanan 300 m, 23°19' N, 121°15'E, 25

August 1994, R. FE Chao, leaf and humus. Paratypes: 2 ¢ 2, same data as holotype; 2 ? 2, 28 August

1996, same habitat as holotype.

Etymology.—The name is dedicated to the entomologist, Dr. T. R. New who

has worked extensively on Zoraptera.

Remarks.—Because the most distinct species characters of many Zoraptera oc-

cur in the male genitalia, specific diagnosis of female Zoraptera is often difficult

(New 1978). The hind femur of the new species resembles Zorotypus lawrencei

New, 1995 from Christmas Island, Indian Ocean (New 1995), but can be differ-

entiated in the number of posterior spines. In this case, Z. palaeus has 4 spines

on the inner hind femora, but Formosozoros newi has only 3, whereas all other

species of Zoraptera have more than 5 posterior spines in the hind femur. How-

ever, the full extent of variability in this feature cannot yet be determined. The

cerci of the new species resemble Z. longicercatus Caudell, 1927 (Caudell 1927),

but lack the apical seta. There are no posterior spines on the hind femur of Z.

longicercatus. The other characters are in agreement with the description of the

genus Zorotypus.

ACKNOWLEDGMENT

The authors thank Dr. T. R. New, Department of Zoology, LaTrobe University,

Bundoora, Vic, Australia, for providing some references and correcting the man-

2000 CHAO & CHEN: NEW ZORAPTERA FROM TAIWAN 29

uscript, and Dr. S. B. Peck, Department of biology, Carleton University, Ottawa,

Canada, for correcting the manuscript.

LITERATURE CITED

Caudell, A. N. 1920. Zoraptera not an apterous order. Proc. Entomol. Soc. Wash., 22: 84-97.

Caudell, A. N. 1927. Zoroty pus longiceratus, a new species of Zoraptera from Jamaica. Proc. Entomol.

Soc. Wash., 29: 144-145.

Gurney, A. B. 1938. A synopsis of the order Zoraptera, with notes on the biology of Zorotypus

hubbardi Caudell. Proc. Entomol. Soc. Wash., 40: 57-87.

Hubbard, M. D. 1990. A catalog of the order Zoraptera (Insecta). Insecta Mundi, 4: 49-66.

Kukalova-Peck, J., & Peck, S. B. 1993. Zoraptera wing structures: evidence for new genera and

relationship with the blattoid orders (Insecta: Blattoneoptera). Syst. Entomol., 18: 333-350.

New, T. R. 1978. Notes on Neotropical Zoraptera, with descriptions of two new species. Syst. Ento-

mol., 3: 361-370.

New, T. R. 1995. The order Zoraptera (Insecta) from Christmas Island, Indian Ocean. Invertebr. Taxon.,

9: 243-246.

Poinar, G. O., Jr. 1988. Zorotypus palaeus, new species, a fossil Zoraptera (Insecta) in Dominican

amber. J. N. Y. Entomol. Soc., 96: 253-259.

Riegel, G. T. 1987. Order Zoraptera. pp. 184-185. In E. W. Stehr (ed.). Immature Insects. Kendall/

Hunt, Dubuque, Iowa.

Silvestri, E 1913. Descrizione di un nuovo ordine di insetti. Boll. Lab. Zool. Portici, 7: 193-209.

Received 4 Apr 1999; Accepted 27 Oct 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 28-48, (2000)

OBSERVATIONS ON THE NESTING BIOLOGY AND

BEHAVIOR OF TRYPOXYLON (TRYPARGILUM) VAGULUM

(HYMENOPTERA: SPHECIDAE) IN COSTA RICA

ROLLIN E. COVILLE!, CHARLES GRISWOLD’, & PAMELA L. COVILLE!

16201 Tehama Ave., Richmond, California 94804

*Department of Entomology, California Academy of Sciences, Golden Gate

Park, San Francisco, California 94118-4599

Abstract.—Trypoxylon (Trypargilum) vagulum Richards, a spider-hunting sphecid wasp, was

studied at the Organization for Tropical Studies’ field station at La Selva in Costa Rica. The

wasps constructed nests in trap-nests with tube diameters of 3.2, 4.8, and rarely 6.4 mm. Nest

structure, cocoon morphology, and differences between male and female provisions are dis-

cussed. Brood cells were provisioned with 7 to 35 spiders comprised mostly of juvenile snare-

building spiders of the families Araneidae, Tetragnathidae, Theridiidae, and Uloboridae. Prey

also included a few juvenile Clubionidae, Ctenidae, and Pisauridae. Natural enemies were Lep-

idophora trypoxylona Hall (Diptera: Bombyliidae), Amobia erythura (Wulp) (Diptera: Sarco-

phagidae), Phalacrotophora punctiapex Borgmeier (Diptera: Phoridae), Macrosiagon lineare (Le

Conte) (Coleoptera: Rhipiphoridae), Trichrysis nigropolita (Bishoff) (Hymenoptera: Chrysidi-

dae), and ants (Hymenoptera: Formicidae).

Key Words.—Insecta, Hymenoptera, Sphecidae, Trypoxylon, Biology, Behavior, Costa Rica.

During 1980 and 1981 we conducted a trap-nest survey in the atlantic lowlands

of Costa Rica to obtain comparative biological information on poorly known

neotropical species of Trypoxylon. Two earlier papers described the nests of T.

(Trypargilum) xanthandrum Richards (Coville & Griswold 1983) and T. (Try-

pargilum) superbum Smith (Coville & Griswold 1984); both species are uncom-

mon wasps. In this paper we describe the nesting biology and behavior of T.

vagulum Richards, the most frequently trap-nested wasp or bee in the study.

Trypoxylon (Trypargilum) vagulum is a solitary wasp, first described by Rich-

ards (1934) from a single male taken at Magdalena, Colombia. The female has

only recently been described and the known range of the species shown to extend

from Veracruz, Mexico, to Colombia (Coville 1982). Its biology is poorly known.

Rau (1933, 1935) reported that 7. vagulum built its larval cells within old mud

cells from nests of T. (Trypoxylon) fabricator Smith. Griswold & Coville (1986),

using a sample of spider prey from nests of 7. vagulum built in trap-nests, ex-

amined the diurnal habits of the types of spiders taken.

STUDY SITE

The trap-nest study conducted in 1980 and 1981 was at La Selva, a field station

of the Organization for Tropical Studies, located (84°00—02’ W, 10°24—26' N)

near the town of Puerto Viejo de Sarapiqui, Heredia Prov. (see Coville & Gris-

wold, 1983, 1984). Nests of 7. vagulum were obtained in trap-nests in four hab-

itats.

Successional strips.—Trap-nests were placed along the edge of newly cut strips

that bordered an old cacao plantation in 1980 and lowland tropical rainforest in

1981 (see Coville & Griswold 1983, 1984).

Arboretum.—Trap-nests were placed throughout a 3.5 ha area circumscribed

2000 COVILLE ET AL.: TRYPOXYLON NEST 29

by undisturbed lowland tropical rainforest and an abandoned cacao plantation (see

Coville & Griswold 1983: fig. 2). In the arboretum the undergrowth was peri-

odically cleared with machete to provide easy access to numerous trees that pos-

sessed identification numbers.

Rafael’s house.—Coville & Griswold (1984) described this residence. Trap-

nests were placed in trees about the yard in 1981 only.

Living quarters——The main structure at La Selva during the study was also the

principal nesting area of T. vagulum in 1980 and 1981. Trap-nests were attached

to beams, railings and other parts of the building.

MATERIALS AND METHODS

Coville & Griswold (1983) provide details of the general trap-nest and rearing

techniques used in the study. We used three types of trap-nests. Standard trap-

nests were 2 X 2 X 16.5 cm blocks of straight grain pine or fir. An 11.0, 9.5,

8.0, 6.4, or 4.8 X 155 mm hole was drilled in each block. Trap-nests with 3.2 X

85 mm holes drilled in them were made from smaller blocks, 1 X 1 X 10-12

cm. Short trap-nests had holes 9.5, 6.4, and 4.8 mm drilled to a depth of only 75

mm. Bundles containing one or two trap-nests (either standard or short trap-nests,

but not both types) with different sized holes were attached to various objects.

Bundles with standard trap-nests were used in all habitats, but bundles with short

trap-nests were used only at the living quarters. Each bundle in the field was

subsequently examined 1-3 times per week. Bundles at the living quarters were

examined daily. During these examinations, we collected trap-nests containing

completed wasp nests and replaced them with trap-nests of the same type. This

generally ensured that wasps had a choice of trap-nest diameters to use in building

nests.

To facilitate behavioral studies, special observation trap-nests were also used

at the living quarters. Observation trap-nests had clear or red transparent plastic

taped over a U-shaped groove or grooves routed in blocks of wood. The grooves

were 9.5, 6.4, 4.8 or 3.2 mm in diameter and their length approximated the depth

of the corresponding diameter holes in standard trap-nests. We sectioned standard

trap-nests and glued a piece to the front of each observation trap. As a result, the

entrance to each observation trap resembled the circular entrance of a standard

trap-nest. Another thin piece of wood was lain over the plastic and loosely wired

into place. This cover created within the grooves a dark environment necessary

to induce the wasps to build their nests. Once a nest had been initiated the cover

could be removed so that the wasp’s nesting activity could be directly observed.

The red plastic caused the least disturbance of the wasps, but was so dark that

we had to use a flashlight to clearly observe their activity. Our later observation

traps all had clear plastic windows. When making routine observations of the

wasp’ behavior we laid a piece of red plexiglass on top of the window so as to

cause as little disturbance of the wasps as necessary. For detailed observation or

photography we would carefully remove the red plexiglass for short periods.

Adult wasps, individual spider prey and the contents of newly provisioned cells

were weighed on a Mettler balance. After weighing, cell contents were carefully

placed in artificial cells within grooves of certain observation trap-nests for rearing

the wasp egg to the adult stage. These rearing traps were stored in a tightly sealed

plastic box provided with a few balls of moist cotton to maintain a high humidity.

30 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Table 1. Summary of trap-nest study at La Selva during 1980 and 1981 at locations frequented by

Trypoxylon vagulum.

No. of

bundles

No. of with T. No. of

trap-nest Bundle Date bundles Date study vagulum T. vagulum

Location bundles pattern® set out ended nests nests

Successional strips 5 1 9-IT-80 5-IV-80 0 0

10 1 30-VII-80 13-IX-80 6 16

10 Z 4-IX-81 22-X-81 1 1

Arboretum 13-15 1 12/13-IV-80 13-IX-80 8 28

20 2 3-IX-81 24-X-81 2 8

4 3 9-IX-81 24-X-81 0 0

Rafael’s house i) 3 5-IX-81 22-X-81 2 2

Living quarters 5 1 15-II-80 15-IX-80 5 72

5 +} 6-VITII-80 9/18-IX-80 5 13

10 4 3-IX-81 26-X-81 8 26

2 3 7-IX-81 26-X-8 1 2 10

4Patterns 1, 2, and 3 had standard trap-nests; pattern 4 had short trap-nests. In pattern 1 bundles

contained one 9.5, two 6.4, two 4.8, and two 3.2 mm diameter trap-nests. In pattern 2 bundles con-

tained two each of 11.0, 9.5, 8.0, 6.4, 4.8, and 3.2 mm diameter trap-nests. In pattern 3 bundles had

one each of 11.0, 9.5, 8.0, 6.4, 4.8, and 3.2 mm diameter trap-nests. In pattern 4 bundles had two

each of 9.5, 6.4, and 4.8 mm diameter trap-nests.

The box was left outside in a screened enclosure. It was examined once or twice

per day to determine how the larvae were developing and to make sure no ants

had succeeded in entering the container. Once the larvae had completed their

cocoons they were transferred to gelatin capsules, where they remained until

emergence.

Voucher specimens of the spider prey are labelled ‘Coville & Griswold Try-

poxylon study’ and are deposited at the California Academy of Sciences. Speci-

mens of the wasps are deposited in the Essig Museum of Entomology at the

University of California, Berkeley.

RESULTS

Trypoxylon vagulum nested in 176 standard and short trap-nests (Table 1) and

accepted trap-nests with 3.2, 4.8, and 6.4 mm diameter holes. In bundles con-

taining equal numbers of all three diameter traps (configurations 1—3 in Table 1),

T. vagulum showed a greater preference for those with 3.2 mm diameter holes

(85) over those with 4.8 (50) and 6.4 (2) mm diameter holes.

We also obtained 10 nests of T. vagulum from observation trap-nests; 3 nests

were in 4.8 mm diameter grooves and 7 were in 3.2 mm diameter grooves. The

discussion on nest characteristics, prey, and enemies pertain to information gath-

ered from the standard and short trap-nests.

Trap-nests set out at the living quarters produced most nests (Table 1). Dwell-

ings and other human structures seem to be a favorable habitat for these wasps,

perhaps because they provide numerous suitable nest sites. Trypoxylon vagulum

readily utilized nail holes and abandoned cells of mud-daubing wasps such as T.

(Trypoxylon) fabricator. One even built its nest in the handle of one of our spare

insect nets. Generally the overhanging roof protected the nest sites from rain.

2000 COVILLE ET AL.: TRYPOXYLON NEST 31

Table 2. General structural characteristics of 155 completed nests of Trypoxylon vagulum in stan-

dard and short trap-nests.

Trap-nest tube sizes in mm

48 X 155 4.8 X 75 3.2 X 85

Total no. of completed nests 41 36 78

No. of nests:

With vestibular cells 34 23 47

Without vestibular cells 7 13 31

With intercalary cells 12 4 4

Without intercalary cells 29 © 32; 74

With closure plug at entrance 35 27 72

With recessed plug 6 2 6

With first cells at inner end of tube 36 35 56

With first cells at distance from end of tube 5 1 22

Provisioned cells:

Total no. 230 159 Zt 2

No. of cells per nest

range 1-10 1-6 1-6

mean + SD 5.6 + 2.4 44+ 1.4 Sy emate! les

Average no. of cells per cm of nest tube 0.36 0.59 0.75

Thus trap-nests used at the living quarters were set out in an area with many

wasps already nesting. The wasps quickly discovered the trap-nests. At times, 4

pairs of T. vagulum were found nesting in a single bundle of trap-nests at the

living quarters.

The other sites (Table 1) were more complex environments in which suitable

nest sites were more scattered. The wasps were probably more dispersed than at

the living quarters and consequently only rarely discovered the trap-nests. Once

a female of T. vagulum discovered a bundle of trap-nests she would build many

successive nests in the same bundle, for as long as suitable diameter traps were

available.

Nest Structure.—Nests in all diameter tubes were similar. The wasps initiated

nests by first depositing a preliminary mud plug. The preliminary plug was usually

a small bit of mud placed at the inner end of the trap-nest tube. Occasionally, the

preliminary plug was a mud wall or partition placed a variable distance from the

inner end (see Table 2). The distance in 4.8 X 155 mm tubes was 26.4—112.0

mm (x = 60.5 mm, n = 4 of 5 instances reported in Table 2). In one 4.8 X 75

mm tube the distance was 3.0 mm. Finally, in 3.2 * 85 mm tubes the distance

was 7.9—45.6 mm (* = 25.8 mm, n = 12 of 22 instances reported in Table 2).

Following the preliminary plug the wasps usually built a series of brood cells

arranged end to end along the length of the trap-nest tube. An empty cell preceded

the first brood cell in one nest in a 4.8 X 155 mm trap and 2 nests in 3.2 X 85

mm traps. Two empty cells preceded the first brood cell in one 3.2 X 85 mm

trap.

Each brood cell was separated from the neighboring cells by a mud cell par-

tition. The inner surface of the cell partitions were convex with a rough lumpy

surface. The outer surfaces were smoothly concave without mud globules ap-

32 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

pressed to the surface. The wasps did not line the trap-nest tubes with mud, so

the exposed wood lining the tubes formed the lateral walls of the brood cells.

Occasionally one or more empty cells (intercalary cells of Krombein 1967b) were

placed between adjacent brood cells. Intercalary cells occurred most frequently

in 4.8 X 155 mm trap-nests. We are uncertain as to their significance (see dis-

cussion by Krombein 1967b). Many nests had one to 4 empty cells (vestibular

cells of Krombein 1967b) placed between the last provisioned cell and the mud

closure plug that sealed the entrance to completed nests (Table 2). Vestibular cells

are commonly found in nests of may kinds of wasps and bees and presumably

discourage enemies from penetrating far enough into completed nests to reach the

larval cells (See Krombein 1967b).

All completed nests possessed a closure plug (Table 2). In only one case was

the closure plug recessed by more than 5 mm (the closure plug was recessed by

15.2 mm in one 4.8 X 155 mm nest). Detailed structure of the closure plug was

examined in thirty-nine 4.8 X 155 mm nests, thirty-five 4.8 X 75 mm nests, and

seventy-six 3.2 X 85 mm nests (Table 3). Among nests in which the closure plugs

were examined, plugs consisted of 1 or 2 mud elements, except one 4.8 X 155

mm nest had 3 elements in its closure plug. Most nests in 4.8 mm diameter tubes

had a single element (26 vs. 12 nests in 155 mm tubes and 25 vs. 10 nests in 75

mm tubes). One and two element plugs were found in an equal number of 3.2 X

85 mm nests (38 nests each). The inner elements were similar to cell partitions.

The outer element, which was appressed to the inner elements or separated from

it by less than 1.0 mm, was generally thicker. When placed at the nest entrances

the outer surface of the closure plug was always smooth and flat or only slightly

concave. In addition, the plugs were clearly defined in that the mud was not spread

onto the wood of the trap-nest.

Small partial mud rings were found in some cells and appeared to represent

sites where a female had initiated and then aborted construction of a cell partition.

In an observation trap-nest we also observed a female apparently accidentally

drop part of the mud she was carrying for constructing a cell partition. This

dropped mud remained in the cell. Similar deposits were occasionally found in

other trap-nests.

For nests opened after all wasp larvae had formed cocoons we took paired

measurements of cell lengths and head widths of the reared wasps (Table 4). The

relationship of cell length and wasp size (head width) was determined with Pear-

son’s product moment correlation coefficient (7). In 3.2 mm diameter nests among

male cells r = 0.01 (n = 64) and among female cells r = 0.31 (n = 18). In 4.8

xX 155 mm nests among male cells r = —0.11 (nm = 25) and among female cells

r = —0.17 (n = 40). None of the r values were significant at P < 0.05, so we

concluded that among wasps of the same sex cell length is not correlated with

wasp SiZe.

Sex ratios and arrangement of male and female cells.—The effects of nest

length and diameter on sex ratios were examined from data presented in Table 5.

Nest length in 4.8 mm diameter trap-nests did not have any apparent effect on

sex ratios. The proportion of males and females reared from conventional and

short trap-nests with tube dimensions of 4.8 X 155 mm and 4.8 X 75 mm, re-

spectively, were not significantly different (x? = 0.32, df = 1). Nest diameter did

have an effect. Trap-nests with 4.8 mm diameter tubes produced more than 2

2000

COVILLE ET AL.: TRYPOXYLON NEST

Table 3. Dimensions of nests of Trypoxylon vagulum in trap-nests (measurements in mm).

3.2 X 85

Nest tube diameter X length

4.8 X 75 48 X 155 6.4 X 155

Female cell length?

range 9.4—32.2 9.9-27.9>« 8.3—27.62> 13.2-14.4

mean (n) 16.4 (19) 12.5 (46) 14.6 (54) 13.8 (3)

Male cell length?

range 9.8—32.9a¢ 8.3—19.7°4 10.6—18.92°¢ —

mean (n) 16.5 (108) 11.5 (20) 13.0 (28) —

Vestibular cell length

range 1.3-53.7 1.4—27.4 2.8-90.4 —

mean (7) 10.9 (33) 8.9 (12) 22.0 (23) —

Intercalary cell length

range 5.0 3.7 5.1-34.8 —

mean (n) 5.0 (1) SC k) 20.0 (2) —

Cell partition thickness

range 0.4-1.6 0.3—1.4 0.4-0.8 —

mean (7) 0.7 (27) 0.6 (56) 0.7 (4) —

Closure plug thickness

range 0.4—4.3 0.6-3.8 1.4-1.5 —

mean (7) 2.1 (34) 1.7 (29) 152) —

4 Measurements do not include brood cells considered abnormally long. For 3.2 and 4.8 mm diameter

nests cells greater than 2 standard deviations from the mean for all (male and female) cells were

deleted. Deleted cells included 4 male cells (42.7, 43.9, 53.8, and 72.9 mm long) from 3.2 mm diameter

nests, 2 female cells (40.2 and 107.4 mm long) and 1 male cell (97.1 mm long) from 4.8 X 155 mm

nests.

> Female cell length in 4.8 X 155 mm nests was significantly greater than in 4.8 X 75 mm nests (t

= 2.95, df = 98, P < 0.05).

¢ Male cell length in 3.2 X 85 mm nests was significantly greater than in 4.8 X 155 mm nests (t =

4.07, df = 134, P < 0.05), and in 4.8 X 155 mm nests male cell length was significantly greater than

in 4.8 X 75 mm nests (t = 1.75, df = 46, P < 0.05). |

4 Female cells were significantly longer than male cells in 4.8 X 75 mm nests (t = 2.04, df = 80,

P < 0.05).

females for every male (45 ¢ 46:91 2 2), whereas 3.2 mm diameter nests produced

more than 8 males for every female (166 66:20 °°). The difference in sex

ratios between 3.2 and 4.8 mm diameter nests was significant (x? = 109.69, df

= 1, P < 0.05).

The arrangement of male and female cells within the nests was nonrandom.

Table 5 shows that cells at the inner end of the nests produced mostly males with

a progressively greater proportion of females reared from cells closer to the en-

trance. The trend was most apparent in the 3.2 mm diameter nests. In all 3 trap-

nest sizes, the observed frequencies of males and females in each cell position

were significantly different from frequencies expected if the probability of a male

or female being produced was the same for each cell position (3.2 X 85 mm trap-

nests— x” = 28.48, df = 4, P < 0.01; 4.8 X 75 mm trap-nests— y? = 9.48, df =

4, P < 0.01; 4.8 X 155 mm trap-nests—y? = 10.33, df = 4, P < 0.01). However,

there was no significant difference between the first 6 cell positions in 4.8 X 75

34 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Table 4. Head widths and cocoon dimensions of Trypoxylon vagulum reared from trap-nests. Mea-

surements are in mm.

Trap-nest tube diameter

3.2 4.8

Males Females Males Females

Head width?

Range 1.9-2.3 2.0—2.3 2.0—2.3 2.1-2.4

Mean (n) 2.1 (86) 2.2 (18) 2.2 (31) 2.3 (56)

Cocoons

Length

Range 7.7-9.8 8.8-10.8 8.1-10.1 8.1-10.3

Mean (n) 8.8 (82)? 9.7 (16) 8.9 (27) 9.5 (49)°

Width at center

Range 2.3-2.9 2.4-3.1 2.7-3.6 3.0-3.8

Mean (n) 2.6 (82)> 2.7 (16) SAME2T IE 3.4 (49)°

Greatest width (near anterior)

Range 2.3-3.1 2.43.2 3.0—4.1 3,.2-4.4

Mean (n) 2.7 (82) 2.9 (16) 3.5 (27) 4.0 (49)

Correlation of head width and cocoon length

r (n) 0.75 (56) 0.84 (16) 0.36 (21) 0.67 (45)

4 Mean head widths of both sexes reared from 3.2 mm diameter nests were significantly smaller than

than those of the same sex reared from 4.8 mm nests (males—t = —3.56, df = 115, P < 0.05;

females—t = —3.51, df = 72, P < 0.05).

>In 3.2 mm diameter nests female cocoons were significantly longer (t = 6.88, df = 96, P < 0.05)

and wider at the center (t = 4.89, df = 96, P < 0.05).

‘In 4.8 mm diameter nests female cocoons were significantly longer (t = 6.53, df = 74, P < 0.05)

and wider at the center (t = 6.92, df = 74, P < 0.05) than male cocoons.

mm and 4.8 X 155 mm trap-nests. Therefore, for a given cell position the prob-

ability of a male or female being produced is the same in the 4.8 X 75 mm and

4.8 X 155 mm diameter nests. There were significant differences in the frequen-

cies of males and females produced at each cell position in 4.8 mm and 3.2 mm

diameter trap-nests (cell position 1, x? = 28.49; pos. 2, x? = 28.88; pos. 3, x? =

29.83; pos. 4, x? = 14.53; pos. 5-6, x? = 9.00, df = 1, and P < 0.01 for all

tests).

Growth and development of the immature stages.—Female wasps always glued

the egg onto the abdomen of one of the largest spiders in the nest, but the egg’s

position varied. The egg is pearly white, sausage-shaped, 1.8 mm long (n = 3),

and 0.5—0.8 mg in weight (n = 2). Under the unnaturally fluctuating but generally

cool conditions in the air conditioned laboratory, elapsed time from oviposition

to emergence of the adult was 26—64 days. Development may take only 21-28

days under natural conditions. In a typical sequence, the embryo required less

than 2 days to develop. The first instar larva punctured the surface of the egg

and the cuticle of the attached spider’s abdomen. For the first 2 days, it fed upon

that spider, completely consuming it. By that time, its growth had become so

rapid that in only another 2—4 days it had devoured all of the remaining spiders.

Within another 3 days, it had completed its cocoon and entered the prepupal stage.

2000 COVILLE ET AL.: TRYPOXYLON NEST 35

Table 5. Distribution of male and female cells in nests of Trypoxylon vagulum from trap-nests.*

Cell position numbered from inner end of the nest

toward the entrance

Trap-nest No. of

size in mm nests 1 2 3 4 5-6 7-9

Fe 2 OR SS" 54 No. males 42 36 31 22 12 —

No. females 0 2 3 6 9 —

4.8 X 75° 30 No. males 12 4 5 a5 2 —

No. females 10 8 11 8 18 —

4.8 X 155° 35 No. males 13 6 4 fi 4 3

No. females 10 12 14 12 15 15

4Nests used to compile this table all began at the inner end of the trap-nest tube.

> Observed frequencies of males and females at each cell position were significantly different at P

< 0.01 from expected frequencies for all trap-nest sizes (3.2 X 85 mm traps— x? = 28.5, df = 4; 4.8

x 75 mm—y? = 9.48, df = 4; 4.8 X 155mm—y? = 10.33, df = 5).

The prepupal stage lasted for 6 or more days and the pupal stage for another 16

or more days. After the wasp moulted to the adult stage, it remained quiescent

for several days, before attempting to cut its way out of its cocoon and leave the

nest.

The fragile cocoons (Figs. 1 and 2) were thin and brittle. They had a varnished,

reddish-brown appearance with a gray anterior end. Their shape was influenced

by the diameter of the nest. In 4.8 and 6.4 mm diameter nests, the cocoons were

robust (Fig. 2) with bulbous anterior ends. In 3.2 mm diameter nests, the cocoons

were slender (Fig. 1) with the anterior ends flaring outward to the to the walls of

LETT EPP PEP PEP PEPE ETP eee aes

Figure 1. Cocoons of Trypoxylon vagulum taken from a 3.2 mm diameter trap-nest. The scale at

the bottom is in mm.

36 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

LEE eee

Figure 2. Cocoons of Trypoxylon vagulum taken from a 4.8 mm diameter trap-nest. The scale at

the bottom is in mm.

the trap-nest tubes. Female cocoons averaged longer and wider at the center than

male cocoons (Table 4), but were otherwise indistinguishable from those of males.

Parental Investment in male and female progeny.—Larval cells were provi-

sioned with 7 to 35 (n = 51) spider prey and the wet weight biomass of all spiders

in a cell ranged from 33.4 to 65.4 mg (nm = 41). We reared 16 females and 11

males of T. vagulum from cells in which the number of prey and total wet weight

biomass of the prey were determined (Table 7). The difference in number of

spiders between male and female cells was not significant but female cells did

have a significantly greater total wet weight biomass of prey than male cells (t =

6.65, df = 25, P < .05).

Besides the difference in biomass of provisions for male and female cells, there

is probably a difference in the biomass of provisions in cells from 3.2 mm and

4.8 mm diameter nests. The evidence is based upon head widths (HW) of adult

wasps produced from cells in these two sizes of nests (Table 4). Males reared

from 3.2 mm diameter nests were significantly smaller than males from 4.8 mm

diameter nests (Table 4). Likewise, females produced from 3.2 mm diameter nests

were significantly smaller than females produced from 4.8 mm diameter nests

(Table 4). Because the developing larvae normally consume all the provisions in

their cells, the difference in size of adults produced from 3.2 and 4.8 mm diameter

nests is probably related to the amount of provisions available to them as larvae.

Prey preferences.—Trypoxylon vagulum preyed upon several families of spi-

ders (Table 9). Most prey were snarebuilding spiders, among which Araneidae

predominated, but occasional Tetragnathidae, Theridiidae and Uloboridae were

also found. Wandering spiders consisted of a juvenile Ctenidae, juvenile Clu-

2000 COVILLE ET AL.: TRYPOXYLON NEST Si

Table 6. Mortality in brood cells of Trypoxylon vagulum obtained from trap-nests.?

Trap-nest tube size in mm

3.2 X 85 4.8 X 155 4.8 X 75 6.4 X 155

No. of nests 68 33 13 2

No. of cells 245 167 54 8

No. of males reared 166 37 8 —

No. of females reared 20 71 20 3

Losses due to unknown factors

Moldy cells fs — —

Dead egg or egg not found 5 11 2 —

Dead larva 2 — — ==

Dead prepupa or pupa 16 8 2, —_

Losses due to enemies

Bombyliidae (Diptera)

Lepidophora try poxylona

Hall 4 4 2 AL.

Anthrax sp. — 1 — —-

Sarcophagidae (Diptera)

Amobia erythrura (Wulp) — 10 16 --

Phoridae (Diptera)

Phalacrotophora punctiapex

Borgmeier 13 19 4 5

Rhipiphoridae (Coleoptera)

Macrosiagon lineare

(Le Conte) 2 aoe, a,

Chrysididae (Hymenoptera)

Trichrysis nigro polita

(Bischoff) 7 2

Ichneumonidae (Hymenoptera)

Poly phaga sp. (?) -=

Formicidae (Hymenoptera) 2 3

Total Losses 59 55 26 5

4 Nests used to compile this table were ones in which all wasp larvae and-parasites had completed

their larval stages before the nests were opened.

> Ants were such a problem that during most of the study we tried to exclude them by coating the

trap-nest wires with Tanglefoot®. Hence, the mortality due to ants is not representative.

bioniidae, and 2 juvenile Pisauridae. Although T. vagulum primarily takes snare-

building spiders, we could not determine if the prey were taken from webs or

other situations, because the diurnal behavior of the prey is variable (Griswold &

Coville 1986). For example, the prey could have been hiding near a web or hiding

without a web. Only for Cyclosa and Micrathena can we be sure that they were

taken from webs, as we always observed these genera in webs during the day.

Mostly small immature spiders comprised the prey of T: vagulum. The distri-

bution of size classes of the prey by number and biomass is skewed toward the

small sized prey (Fig 3). Although individual weights of spider prey ranged from

0.2 to 21.3 mg, 64% of the prey weighed 2.0 mg or less.

38 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Table 7. Amount of provisions in male and female cells of Trypoxylon vagulum.

Male Cells Female Cells

(n = 11) (n = 16)

No. of spiders

range 10-28 11-35

mean + SD 13:3 +, 5.3 19.6 + 5.9

Total wet weight biomass of provisions?

range 33.4—47.9 41.7-65.4

mean + SD 41.5 + 5.0? 51.6 + 5.68

4 Wet weight biomass of provisions was significantly greater in female cells than male cells (t =

6.65, df = 25, P < 0.05).

Mortality and Natural Enemies.—We have no information on mortality of adult

wasps, but Table 6 summarizes the mortality occurring in brood cells. All of the

nests used to compile the table were collected from the field within a few days

of their completion. A greater mortality probably would have been evident if the

nests had been left in place, because many would probably have been discovered

by enemies capable of digging their way through the closure plugs or ovipositing

through walls of the trap-nests.

The greatest losses (48% of the cells) occurred in short (4.8 X 7.5 mm) trap-

nests, but this is more related to their being only set out at the living quarters

where natural enemies were especially abundant. The 3.2 X 85 mm traps and 4.8

x 155 mm traps were distributed in equal numbers among all habitats. The brood

cells from 3.2 X 85 mm traps suffered significantly less mortality (24% of the

Table 8 Mortality of immature stages of Trypoxylon vagulum in the first 2 and last 2 cells in

completed nests.?

First 2 Last 2

cells of nest cells of nest

3.2 X 85 mm nests (n = 38)>

No. of cells 76 76

No. of adults reared 63 61

Losses‘ 13 15

4.8 X 155 mm nests (n = 2)°

No. of cells 44 44

No. of adults reared 28 29

Losses* 16 15

4.8 X 75 mm nests (n = 9)>

No. of cells 18 18

No. of adults reared 7 6

Losses* 11 12

4 Nests used in this study were selected by the following criteria: 1) All wasp larvae and parsasites

had completed their larval stages before the nests were opened, and 2) the nests contained 4 or more

cells.

> Number of nests.

° Losses in the first 2 and last 2 cells were not significantly different.

2000 COVILLE ET AL.: TRYPOXYLON NEST 39

Table 9. Prey preferences of Trypoxylon vagulum.

Spider Prey Number of Prey

Araneidae

Eustala sp. #1 54

Eustala sp. #2 4

Eustala sp. #3 6

Eustala miscellaneous juveniles 93

Verrucosa sp. 89

Acacesia sp. 74

Wagneriana tauricornis v5

Wagneriana sp. #1 21

Wagneriana sp. #2 4

Wixia or Parawixia sp. 9

Micrathena sp. 47

Gasteracantha sp. 5

Scoloderus sp. 1

Cyclosa 2

undetermined sp. #1 6

undetermined sp. #2 5

undetermined sp. #3 78

Clubionidae

Clubionia sp. 1

Ctenidae

undetermined sp. #1 1

Pisauridae

undertermined sp. 2

Tetragnathidae

undetermined sp. #1 1

undetermined sp. #2 1

undetermined sp. #3 1

undetermined sp. #4 (Tetragnatha) 1

undetermined sp. #5 1

Theridiidae

undetermined sp. #1 1

Uloboridae

Miagrammopes 9

undetermined sp. #1 5

undetermined sp. #2 1

cells) than did the brood cells from 4.8 X 155 mm traps (33% of the cells) (X?

= 3.89, df = 1, P < 0.05).

We suspected that the longer a nest was being constructed the greater the prob-

ability of its discovery by natural enemies. If this is the case, we could expect a

greater mortality in the last cells to be provisioned than in the first cells provi-

sioned. Nevertheless, our data did not support this contention. In completed nests

with a total of 4 or more cells, losses in the first 2 and last 2 cells were not

significantly different (Table 8).

The natural enemies listed in Table 6, exhibit diverse modes of attacking nests

of T. vagulum. Adults of the bombyliid flies Lepidophora trypoxylona Hall and

Number of Prey

Weight of Prey in mg

THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Trypoxylon vagulum prey size

Total Number of Prey = 333

0 5 10 15 20 25

—_

(=)

Total Wet Weight of All Prey = 792 mg

100

Size Classes of Spider Prey in mg

Figure 3. Size classes of prey taken from nests of Trypoxylon vagulum. The top bar graph breaks

down the size classes by the number of spiders occuring in each class. The bottom bar graph breaks

down the size classes by the total wet weight of all spiders in each class.

Anthrax sp. hovered in front of nests while presumably ejecting microscopic first

instar larvae at the entrance opening. The larva of Anthrax developed as a para-

sitoid upon a single wasp larva. Before attacking its host the Anthrax larva pre-

sumably waited until the host larva had completed its feeding, because the An-

thrax adult emerged from a completed cocoon of T. vagulum. Lepidophora try-

poxylona was more destructive. Its larva first killed the host egg or young larva

and then consumed the spider prey by sucking the contents out of their bodies.

2000 COVILLE ET AL.: TRYPOXYLON NEST 41

Larvae of L. trypoxylona commonly invaded additional cells if there were insuf-

ficient food in the initially infested cell. Pupae of both bombyliids possessed

powerful cephalic teeth that enabled them to dig their way out of nests shortly

before emergence of the adult fly.

The sarcophagid fly, Amobia erythrura (Wulp) was only found at the living

quarters, but was very destructive in 4.8 mm diameter nests of T. vagulum at that

site. None of the 3.2 mm diameter nests were attacked: Adults of A. erythrura

stationed themselves near bundles of trap-nests and attempted to follow prey laden

females of 7. vagulum into their nests. Successful attacks resulted in a cell being

infested by 1 to 7 maggots. The maggots appeared to work in unison, first at-

tacking the wasp egg, and then consuming the spider prey. Amobia maggots

always worked their way toward the nest entrance and destroyed every cell in

their path. Once at the nest entrance they bored their way partially through the

closure plug and then pupated. The adult Amobia usually could punch their way

through the remaining part of the closure plug and escape the nest. In one nest

the maggot failed to work its way to the entrance and the adult fly being unable

to punch its way out of the nest died next to its puparium.

The phorid fly, Phalacrotophora punctiapex Borgmeier is a small (33-4 mm

long), fast running insect that frequented the entrances of wasp nests. Adults of

P. punctiapex entered nests by evading the host wasps. Once in the nest the flies

hid among the spider prey and proceeded to lay many eggs on the walls and mud

partition at the inner end of the cell. The newly emerged maggots first attacked

and consumed the host wasp egg and then fed upon the spider prey. Often the

maggots invaded additional cells by burrowing through the cell partitions. Nev-

ertheless, the maggots seldom made their way to the nest entrance. Instead, they

pupated within a brood cell. Their puparia were strongly glued to the walls of

the trap-nest tube. It is unclear to us as to how P. punctiapex escape the nests.

The adults have no obvious digging structures for penetrating cell partitions, and

they generally seem to emerge before the host wasps. Cells of 7. vagulum were

infested by 1-15 maggots and one nest contained 34 maggots.

An adult of the rhipiphorid beetle, Macrosiagon lineare (Le Conte) was reared

from each of 2 cocoons of 7. vagulum. The cocoons were from different nests.

The chrysidid wasp, Trichrysis nigropolita (Bishoff), was reared from cells of

T. vagulum, although we did not observe any successful attacks. Adult chrysidids

waited at the nest entrances for an opportunity to enter an unguarded nest and

Oviposit into a newly provisioned cell. Coville & Coville (1980) observed this

chrysidid species chewing holes through outer cell partitions and suspected that

they can oviposit into a cell through such a hole.

An occasional spider that was brought into the nest was parasitized by an

ichneumonid (Polyphaga sp.). In one cell the ichneumonid completed its devel-

opment and spun a cocoon. On emerging the ichneumonid in attempting to escape

from the nest killed a T. vagulum pupa.

Probably the most severe enemies of wasps at La Selva were the numerous

species of ants. At the living quarters Monomorium floricula (Jerdon) and Tetra-

morium bicarinatum (Nylander) were the principal nest raiders. At other sites

several other ant species nested in trap-nests or raided them. These included Cam-

ponotus abdominalis Mayr, Camponotus planatus Roger, Crematogaster limata

palans Forel, Pachycondyla unidentata Mayr, Paratrechina caeciliae Forel, So-

42 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

lenopsis picea Fomery, and Solenopsis sp. Many of these species and especially

Tetramorium and Monomorium were capable of digging through closure plugs

and looting completed nests. Male wasps were usually able to thwart individual

foraging ants but if ants were able to gain entrance to the nest through temporary

absence of the male or by overwhelming him, the nest was lost. As noted in Table

6, we coated the wires supporting trap-nest bundles with Tanglefoot® to discour-

age ants from attacking nests. Otherwise, ants would have destroyed many more

nests.

Brockmann (1992) reported chalcid wasps in the genus Mellitobia attacking

nests of T. monteverdeae Coville from Monteverde, Costa Rica. Nevertheless, we

encountered no chalcids in nests 7. vgulum or any other Trypoxylon species from

La Selva.

Nest supercedure.—Four traps from which T. vagulum were reared also con-

tained cells of other species of wasps and bees. In the arboretum a 4.8 X 155

mm trap had one cell of T. vagulum at the inner end of the tube. On top of the

cell were two empty cells and then a 4-celled nest of an undetermined species of

Trypoxylon. At the living quarters a 4.8 X 155 mm nest had 1 cell of JT. (Try-

pargilum) majus Richards followed by a 3-celled nest of 7. vagulum. Another 4.8

x 155 mm nest at the living quarters had 1 cell of T. vagulum 49.4 mm from the

inner end of the tube followed by 2 cells of a resin bee (Megachilidae). Also at

the living quarters a 3.2 X 85 mm trap had two cells of 7. vagulum at the inner

end of the tube followed by a 2-celled nest of an eumenid wasp.

Behavioral observations.—Our observations took place at the living quarters.

Trypoxylon vagulum was one of several species in the subgenus 7rypargilum that

nested in trap-nests at the living quarters. Trypoxylon lactitarse Saussure and an

undetermined sibling species (C, corresponds to Sp. C in Griswold & Coville

1986) were almost as abundant as 7. vagulum. Another undetermined species (A,

corresponds to Sp. A in Griswold & Coville 1986), related to T. nitidum Smith,

was also abundant. Trypoxylon saussurei Rohwer and T. majus were occasional

trap-nest occupants in 1980.

Trypoxylon vagulum was the only species to use 3.2 mm diameter trap-nests,

but it did compete with T. saussurei, T. majus, and species A for 4.8 mm diameter

nests. There was little direct competition, however, because we insured that sev-

eral empty trap-nests were continuously available to the wasps. Trypoxylon lac-

titarse and species C only used trap-nests with diameters of 6.4 mm or larger.

During fair weather, 7. vagulum were active from 1 to 2 hours after sunrise

until 1.5 to 2.5 hours before sunset. They ceased their foraging and nest construc-

tion activities during rainy spells.

Like most Trypoxylon wasps in the subgenus Trypargilum, a male of T. va-

gulum would pair with a female that was initiating a nest. Thereafter he would

guard the nest while the female hunted for spiders or foraged for mud. During

the day the male usually remained face outward at the nest entrance and would

snap his mandibles at potential intruders, such as ants. The male seldom left the

nest unguarded for more than a few minutes. Occasionally they did leave for 5—

15 minutes, when the female was present. The male would often leave his nest

to pursue or rarely butt unpaired males of 7. vagulum that alighted on or hovered

near his nest. Nevertheless, we never observed males of T. vagulum grappling

with one another or physically trying to usurp another male’s nest. In contrast,

2000 COVILLE ET AL.: TRYPOXYLON NEST 43

males of T. lactitarse and species C often engaged in fights and attempted to

invade one another’s nest.

Active nests of T. vagulum always had a male associated with them. In fact, a

female of JT. vagulum would not leave to forage unless a male was in the nest.

At night the female would remain in her nest, whereas a male would be present

or absent.

One or two males would follow females that were searching for new nest sites.

So pair formation usually took place even before a new nest had been initiated.

When searching for a new nest, a female of T. vagulum in a slow hovering

flight would move along the surface of objects such as beams, railings, posts, etc.

She would frequently alight to examine dark spots and small holes, which she

would attempt to enter headfirst. If the tubular cavity within a hole was potentially

suitable for a new nest, she would begin to clean it out. Otherwise, she would

resume her search.

In preparing a new nest site the female would remove any material blocking

or constricting the tube. The material included loose debris as well as mud and

frass that adhered to the walls of the tube. She would use her mandibles to scrape

the surface of the tube. With a load of debris cradled with her mandibles and

front legs she would back out of the nest and either immediately drop the load

or take flight and drop the load 3—50 cm from the nest entrance. Once the nest

had been cleaned, she would then leave the nest and return with a load of mud.

She used the mud for the preliminary plug at the inner end of the nest. Several

loads of mud were required for the plug.

Females of T. vagulum were observed obtaining mud from nests of mud-daub-

ing wasps, especially those of Trypoxylon (Trypargilum) species in the Abitarse

Group (see Coville 1982, for classification) and 7. (Trypoxylon) fabricator. Fe-

males apparently carried water that they regurgitated to soften the hard mud on

nests of the host wasps. They then removed a small ball of mud with their man-

dibles and transported it to their own nest. We were unable to determine if T.

vagulum uses other sources of mud. We did not observe them at muddy spots

frequented by other species of Trypoxylon.

Once the preliminary plug was completed, the female began to forage for spi-

ders. Females usually returned to their nests with spiders within 6—15 minutes,

but foraging flights as short as 2 minutes and longer than 1 hour were also ob-

served. On returning to the nest with a spider the female would alight at the

entrance. The male, after tapping the female’s antennae with his own, would leave

the nest and climb upon the female’s back. The female would then enter the nest

with the male following her. Once inside the nest the male would often invert

himself, climb beneath the female and grip the base of her petiole with his man-

dibles. In this manner the pair would proceed further into the tube, until the female

dropped her spider.

During the early stages of provisioning, the female often merely deposited the

spider, groomed herself and then left on the next foraging trip. After the female’s

departure the male would pack the spiders tightly into the mass of provisions by

butting the spider with his head. When the cell was almost fully provisioned, the

female would usually pack the spiders with her head as well.

Within nests in 3.2 mm diameter grooves in observation trap-nests there was

insufficient space for the wasps to turn around. They would first have to leave

44 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

the nest in order to reverse their position. Within 4.8 mm diameter grooves the

wasps were able to turn around.

After the final spider had been obtained the female would usually leave the

nest for a few minutes. On returning she would inspect the prey, firmly pack them

with her head, and then groom herself. When she was ready to oviposit, she would

back out of the nest, reverse her position, back into the nest, and begin probing

the mass of spiders for a suitable abdominal surface upon which to oviposit.

During oviposition, she remained motionless.

We observed 10 cells of 7. vagulum continuously from initiation through ovi-

position. In addition, we made numerous more fragmentary observations on pro-

visioning and oviposition in other cells. Nevertheless, mating was never observed,

although a male was present at all times. This was in direct contrast to T. lactitarse

and species C in which mating took place immediately before oviposition in

almost every case. In fact, unless mating had occurred, males of the latter two

species would disrupt their mate’s attempt to oviposit.

Of the 10 ovipositions observed in T. vagulum, 7 males were reared from the

resulting eggs (the other 3 rearing attempts failed). In 7. lactitarse and species C,

eggs resulting from ovipositions preceded by mating produced males and females;

in the 1 case in which mating did not precede oviposition, the egg failed to

develop. '

After oviposition, the female would begin construction of the cell partition to

seal the cell. She spread her first load of mud in a thin band across the bottom

of the tube in front of the provisions. She spread the mud with her mandibles

and simultaneously vibrated her flight muscles, which emitted a buzzing sound

audible for about 1 meter from the nest. Several loads of mud were required to

complete a partition. During the final stages her body would rotate so that she

could spread mud across the roof of the tube and complete the closure of the cell.

The male watched the female or held onto her petiole with their mandibles while

she spread the mud. We did not observe males assisting females in construction

of partitions or closure plugs.

DISCUSSION

We chose to follow the classification provided by Coville (1982), in which the

genus Trypoxylon is divided into the subgenera Trypoxylon and Trypargilum. The

two subgenera are distinguished on the basis of a several of morphological char-

acters. Trypar gilum, confined to the Western Hemisphere, also differs in the wide-

spread occurrence of male guarding of nests and the generally darker and harder

cocoons. Unless specifically stated, the following discussion is restricted to species

of the Trypoxylon species in the subgenus Trypargilum.

Most authors have informally recognized several species groups and complexes

of Trypargilum (Richards 1934, Krombein 1967a, Krombein 1979, Coville 1982).

Three groups are fairly well defined morphologically: the Albitarse Group, the

Superbum Group, and the Nitidum Group (Bohart & Menke 1976, Coville 1982).

The Albitarse Group contains all species that build mud nests, generally in the

form of a series of vertical mud tubes. The other groups nest in pre-existing

tubular cavities. The Superbum Group, recognized by a transverse carina on the

frons, only contains a few tropical species. The Nitidum Group encompasses the

majority of species. It is a heterogeneous collection divided, not always success-

2000 COVILLE ET AL.: TRYPOXYLON NEST 45

fully, into several species complexes. Trypoxylon vagulum is a member of the

vagum complex (see Coville 1982). Krombein (1967b) and Matthews & Matthews

(1968) found that nest architecture, cocoon structure and prey preferences seem

to vary according to the species groups and complexes. Subsequent studies have

continued to build upon that framework.

Overall, the nest structure of T. vagulum resembles that of other species of the

Nitidum and Superbum groups, although because of its small size it is able to

use smaller diameter nests than any species studied so far. As with most species,

T. vagulum built the closure plugs at the nest entrance. Two species diverge from

this pattern by having the closure plugs recessed from the entrance. One of these

species is T. superbum (Coville & Griswold 1984) of the Superbum Group. The

other is T. xanthandrum (Coville & Griswold 1983) of the Nitidum Group’s fugax

complex. Among all species of 7rypargilum observed at La Selva, the outer sur-

face of plugs of T. vagulum were distinctive in that they were smoothly concave

and their mud was never spread onto the outer surface of the trap-nest itself.

Other species were not so neat.

Trypoxylon vagulum seems to obtain mud for its nests from the mud nests of

other Trypoxylon wasps, particularly T. ([rypoxylon) fabricator. At typical sites

where many species of Trypoxylon of both subgenera mine mud, such as muddy

spots along trails, J. vagulum was absent in our observations, despite being the

most abundant species trap-nested. Nevertheless, we do not feel that we can totally

discount the possibility that T. vagulum may occasionally use such sites.

Genaro (1996) found that the inner walls of cell partitions of 7. (Trypargilum)

subimpressum Smith (excavatum complex) generally have a glob of mud ap-

pressed to them which is mined by the larva for incorporation into its cocoon. In

freshly provisioned cells of T. vagulum, the inner walls of cell partitions were

generally smoothly concave, and we noticed no mud globs adhering to them.

Nevertheless, the larvae did mine the inner partitions for mud during cocoon

spinning, as reported by Genaro (1996). This is probably typical of the wasps in

the subgenus. In our own observations of larvae spinning cocoons, the larvae

sometimes appear to be regurgitating material into the cocoon matrix.

Cocoons of J. vagulum are distinct from all other species in the subgenus

Trypargilum, not only by their small size, reflecting the small size of T. vagulum

compared to other Trypargilum (Coville 1982), but also by their shape. In this

species, as the nest diameter becomes larger, the cocoons become more globular

with the anterior end distinctly swollen (Fig. 2) but not flaring outward. This

outward flare is frequently seen in species of the punctulatum complex (see Krom-

bein 1967b, Camillo et al. 1993, 1994; Coville 1981, 1982), fugax complex (Co-

ville & Coville 1980, Coville & Griswold 1983), and Superbum group (Coville

& Griswold 1984). In these and other species, the cocoon diameter at the middle

does not increase greatly with nest diameter increase, as it does in T. vagulum.

Compared with other species of Trypargilum we observed at La Selva including

T. lactitarse, T. superbum, T. xanthandrum, T. vagum Richards, T. agamemnon

Richards, T. saussurei, T. majus, and a couple of undetermined species the co-

coons of T. vagulum appeared to be somewhat lighter in color and more fragile.

Among TJrypargilum species that normally nest in pre-existing cavities (Super-

bum and Nitidum Groups), nests generally have a non-random distribution of

male and female cells. There are two trends. First, for a given species, the smallest

46 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

acceptable diameter nests produce mostly males. The proportion of females in-

creases in larger diameter nests. When cells of both sexes are present, male cells,

with few exceptions, occur more frequently at the inner end of the nests. In some

species, such as T. tenoctitlan Richards (Coville & Coville 1980), 7. tridentatatum

Packard (Krombein 1967b, personal observation) and T. lactitarse (Krombein

1967b, under the name striatum) in North America, male and female cells are

clearly segregated with male cells at the inner end and female cells toward the

entrance. In other species, such as T. vagulum, T. clavatum Say (Krombein

1967b), T. collinum collinum Smith (Krombein 1967b), T. lactitarse (Camillo et

al. 1993) in Brazil, and T. rogenhoferi Kohl (Camillo et al. 1994), male and female

cells are frequently intermixed. Nevertheless, the probability of a given cell being

a male or female cell still relates to its position in the nest. One exception, may

be JT. collinum rubrocinctum (Packard) in which Krombein (1967b) reported fe-

male cells to be at the inner end and males at the outer end. In contrast, species

that build mud nests (Albitarse Group), such as T. politum Say (Brockmann &

Grafen 1989) and T. monteverdeae (Brockmann 1992), appear to have a random

distribution of male and female cells in the mud tubes.

In the present study, we also found that males and females reared from small

3.2 mm diameter nests were significantly smaller than individuals of the same

sex from 4.8 mm diameter nests. Among wasps reared from the same diameter

nests, we found no correlation between size of the wasps and cell length.

Prey preferences of Trypargilum vary widely (Matthews & Matthews 1968,

Coville 1982, Griswold & Coville, 1986). Among species of the Albitarse Group

studied, prey consist entirely of orbweaving spiders of the family Araneidae, prin-

cipally among the genera Eustala, Neoscona, and, less frequently, Araneus (see

review of Coville 1982, Brockmann & Grafen 1992 on T. politum, Brockmann

1992 on T. monteverdeae). The only species of the Superbum Group so far stud-

ied, T. superbum (Coville & Griswold 1984, Griswold & Coville 1986), special-

izes on Salticidae. Among species in the Nitidum Group, prey preferences vary

from extremely narrow to broad. For example, T. xanthandrum of the fugax com-

plex appears to specialize on spiders of the family Senoculidae, a poorly known

group of wandering spiders (Coville & Griswold 1983, Griswold & Coville 1986).

Trypoxylon tenoctitlan of the fugax complex (Coville & Coville 1980), and spe-

cies in the spinosum complex normally take a wide variety snarebuilding and

wandering spiders (see review of Coville 1982) found on vegetation and manmade

structures. Data of Genaro et al. (1989) indicate that Trypoxylon subimpressum

of the excavatum complex may also fall into this later category.

Trypoxylon vagulum preys almost entirely on snarebuilding spiders. This strong

preference for snarebuilding spiders is shared with species of the nitidum com-

plex, T. orizabense Richards, and T. tridentatum (see review of Coville 1982,

O’Brian 1982, Coville 1986, Jiménez & Tejas 1994). Species in the punctulatum

complex also show a preference for snare building spiders (see review of Coville

1982, Camillo et al. 1993 on T. lactitarse, 1994 on T. rogenhoferi). A preference

for snare-building spiders does not necessarily mean that the wasps hunt for spi-

ders that are in webs. Griswold & Coville (1986, Table 2) noted that the diurnal

habits of prey of 7. vagulum at La Selva fall into 5 patterns, all but the last pertain

to snarebuilding species: 1) on intact web (eg., Cyclosa, Micrathena, and Ver-

rucosa),; 2) at the edge of intact webs (eg., araneid undetermined sp. #3); 3)

2000 COVILLE ET AL.: TRYPOXYLON NEST 47

cryptic on substrate, no retreat, intact web present (eg., Eustala); 4) cryptic on

substrate, retreat and web absent (eg., Acacesia); 5) motionless, exposed on fo-

liage (Clubionidae and Ctenidae).

Brockmann & Grafen (1989) showed that females of T. politum provision male

cells with less biomass of spiders than female cells. This is also the case with T.

vagulum, even though we found no significant difference in the number of prey

in male and female cells.

Camillo et al. (1993) found that brood cells nearest the entrance in nests of T.

lactitarse suffered the greatest mortality. This seems logical for a couple of rea-

sons. First, some enemies such as the bombyliid fly Lepidophora, phorid fly

Phalacrotophora, and sarcophagid flies may destroy many or all of the cells

between the initially infested one and the nest entrance. Second, some enemies

attack completed nests. Among these, some can oviposit through the closure plug

while others such as certain mutillid wasps, and some ants, burrow through the

closure plug. Although we found no significant difference in the mortality rates

between the first two and last two cells in 7. vagulum nests (Table 8), our data

are irrelevant with regard to ants and enemies that attack completed nests. We

used Tanglefoot® to discourage attacks by ants before and after nest completion.

In addition, we collected nests from the field within a few of days of completion.

This artificially reduced their vulnerability to enemies that attack competed nests.

Studies by Brockmann & Grafen (1989) on T. politum, Brockmann (1992) on

T. monteverdeae, Coville & Coville (1980) on T. tenoctitlan, and our own un-

published observations on T. lactitarse and species C indicate that mating takes

place in the nest. In particular, mating seems to take place just before oviposition,

when the male becomes particularly sexually aggressive. Nevertheless, with T.

vagulum we observed no matings inside or outside of nests. Do they mate away

from the nests or before nesting begins? Did the transparent windows in our

observation trap-nests disturb them more than other species we watched? What-

ever possible reasons, the mating behavior of 7. vagulum requires further study.

ACKNOWLEDGMENT

We gratefully acknowledge the logistic support of the Organization for Tropical

Studies. The following individuals kindly identified natural enemies of Trypoxy-

lon: R. M. Bohart (Chrysididae), R. J. Gagne (Sarcophagidae), J. C. Hall (Bom-

byliidae), R. R. Snelling (Formicidae), W. W. Wirth (Phoridae). This research was

supported by a grant from the National Geographic Society (R. E. Coville, prin-

cipal investigator).

LITERATURE CITED

Brockmann, H. J. 1992. Male behavior, courtship and nesting in Trypoxylon (Trypargilum) montev-

erdeae (Hymenoptera: Sphecidae). J. Kans. Entomol. Soc., 65: 66-84.

Brockmann, H. J. & A. Grafen. 1989. Mate conflict and male behavior in a solitary wasp, Trypoxylon

(Trypar gilum) politum (Hymenoptera: Sphecidae). Anim. Behav., 37: 232-255.

Camillo, E., C. A. Garofalo, G. Muccillo & J. C. Serrano. 1993. Biological observations on Trypoxylon

(Trypargilum) lactitarse Saussure in southeastern Brazil (Hymenoptera: Sphecidae). Rev. Bras-

ileira de Entomol., 37: 769-778.

Camillo, E., C. A. Garofalo & J. C. Serrano. 1994. ObservacGdes sobre a biologia de Trypoxylon

(Trypar gilum) rogenhoferi Kohl (Hymenoptera: Sphecidae). Anais Soc. Entomol. Brasil, 23:

299-310.

48 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Coville, R. E. 1979. Biological observations on Trypoxylon (Trypargilum) orizabense Richards in

Arizona (Hymenoptera: Sphecidae). J. Kansas Entomol. Soc., 52: 613-620.

Coville, R. E. 1981. Biological observations on three Trypoxylon wasps in the subgenus Trypar gilum

from Costa Rica: T. nitidum schulthessi, T. saussurei, and T. lactitarse (Hymenoptera: Sphe-

cidae). Pan-Pacific Entomol., 57: 332-340.

Coville, R. E. 1986. Spider Prey of Trypoxylon tridentatum (Hymenoptera: Sphecidae) from Arizona

and California. Pan-Pacific Entomol., 62: 119-120.

Coville, R. E. & P. L. Coville. 1980. Nesting biology and male behavior of Trypoxylon (Trypar gilum)

tenoctitlan in Costa Rica (Hymenoptera: Sphecidae). Ann. Entomol. Soc. Am., 73: 110-119.

Coville, R. E. & C. Griswold. 1983. Nesting biology and male behavior of Trypoxylon (Trypar gilum)

xanthandrum in Costa Rica with observations on its spider prey (Hymenoptera: Sphecidae;

Ananeae: Senoculidae). J. Kans. Entomol. Soc., 56: 205-216.

Coville, R. E. & C. Griswold. 1984. Biology of Trypoxylon (Trypargilum) superbum (Hymenoptera:

Sphecidae), a spider-hunting wasp with extended guarding of the brood by males. J. Kans.

Entomol. Soc., 57: 365-376.

Genaro, J. A. 1996. Estructura del nido y capullo de Trypoxylon (Trypargilum) subimpressum (Hy-

menoptera: Sphecidae). Caribbean J. Sci., 32: 240-243.

Genaro, J. A., A. Sanchez & G. A. Garcia. 1989. Notas sobre la conducta de la nidificacién de

Trypoxylon (Trypargilum) subimpressum Smith (Hymenoptera: Sphecidae). Caribbean J. Sci.,

25: 228-229.

Griswold, C. E. & R. E. Coville. 1986. Observations on the prey and nesting biology of spider-hunting

wasps of the Genus Trypoxylon (Hymenoptera: Sphecidae) in Costa Rica. Proceedings of the

IX International Congress of Arachnology (Panama 1983), pp. 113-116.

Jiménez, M. L. & A. Tejas. 1994. Las Arafias presa de la avispa lodera Trypoxylon (Trypargilum)

tridentatum tridentatum en Baja California Sur, Mexico. Southwestern Entomol., 19: 173-180.

Krombein, K. V. 1967a. Jn Krombein, K. V. & B. D. Burks (eds.). Hymenoptera of America north of

Mexico, synoptic catalogue. U.S. Dept. Agr. Monogr. 2, 2nd suppl.

Krombein, K. V. 1967b. Trap-nesting wasps and bees: life histories, nests, and associates. Smithsonian

Inst. Press, Washington, D.C.

Krombein, K. V. 1979. pp. 1199-2209. Jn Krombein, K. V. et al. Catalogue of Hymenoptera in

America north of Mexico, vol. 2, Apocrita (Aculeata). Smithsonian Inst. Press, Washington,

De:

Matthews, R. W. & J. R. Matthews. 1968. A note on Trypargilum arizonense in trap nests from

Arizona, with a review of prey preferences and cocoon structure in the genus (Hymenoptera:

Sphecidae). Psyche, 75: 285-293.

O’Brien, M. F 1982. Trypargilum tridentatum (Packard) in trap nests in Oregon (Hymenoptera: Sphe-

cidae). Pan-Pacific Entomo., 58: 288—290.

Rau, P. 1933. The jungle bees and wasps of Barro Colorado Island. Phil Rau, Kirkwood, Mo., 324 pp.

Rau, P. 1935. Additional Trypoxylon names in ‘‘Jungle Bees and Wasps of Barro Colorado Island.”’

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R. Entomol. Soc. Lond., 82: 173-362.

Received 19 Mar 1999; Accepted 27 Oct 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 49-51, (2000)

A NEW SPECIES OF ENCARSIA

(HYMENOPTERA: APHELINIDAE), A PARASITOID OF

WHITEFLY ALEURODICUS SP.

(HOMOPTERA: ALEYRODIDAE) IN MEXICO

JAIME GOMEZ! AND OSWALDO GARCIA2

'F] Colegio de la Frontera Sur, Apartado Postal 36,

Tapachula, Chiapas, 30700 México

Universidad Auténoma Agraria “‘Antonio Narro” Buenavista,

Saltillo Coahuila, 25315 México

Abstract.—Encarsia narroi, NEW SPECIES, is described from Mexico. Females of this species

were collected from fourth instar nymphs of an Aleurodicus sp. on the host plants Bauhinia

variegata and Hibiscus sp. at Parras, Cohuila State, México. The new species is similar to

Encarsia coquilletii.

Key Words.—Insecta, Hymenoptera, Aphelinidae, Encarsia, Mexico.

About 100 species of parasitoids have been identified among the natural ene-

mies of whiteflies (van Lenteren et al. 1996). Most of these parasitoids belong to

the family Aphelinidae (Hymenoptera: Chalcidoidea), although Scelionidae, Ce-

raphronidae, Encyrtidae, Eulophidae and Platygasteridae species have also been

reported (Gerling 1990, Myartseva & Yasnosh 1994, Polaszek et al. 1992). The

most important whitefly parasitoids belong to the generia Encarsia and Eretmo-

cerus (Hennessey et al. 1995, Polaszek et al. 1992, Schauff et al. 1996). In the

present paper we report and describe an Encarsia species from Mexico, it was

identify as undescribed species by R. C. Williams, Agricultural Research Service

Laboratory (ARS-USDA), which is described here as Encarsia narroi G6mez &

Garcia species nova, named in honor of Antonio Narro founder of the Agricultural

University “‘Antonio Narro” Saltillo, Coahuila, Mexico.

ENCARSIA NARROI GOmez & Garcia, NEW SPECIES

(Figs. 1-3)

Diagnosis.—Female, brown coloured; antennal club quite wide (0.06 mm) and

spindle shaped, with the terminal of the 3rd segment more clearly apically pointed

than the base of the Ist segment (Fig. 1); mesoscutum with 42 pairs of long setae

(Fig. 2); forewing uniformly setose, except below the submarginal vein which

bears 10 setae and one small clear area below the stigmal vein (Fig. 3).

Description—Female. Body length 1.37—1.4 mm. Brown coloured, except scutellum which is pale

yellow. Funicular segments, front legs, middle legs, and tibiae, tarsus hind legs also pale yellow

coloured. Vertex and part of face striated, with short stout setae; setae surrounding the eyes softer and

shorter than those on the vertex. Eye colour orange. Antenna densely pilose, with 3 funicle and 3 club

segments, is about 0.48—0.52 mm long; antennal scape slender; pedicel conical subequal in length to

funicular segments; funicular segments of equal size, 0.066—0.073 mm long and about 1.6 times longer

than wide; antennal club wide and spindle shaped, with 3rd club segment more apically pointed than

the Ist segment (Fig. 1), measures 0.16—0.17 mm long and about 2.8 times longer than its widest

point.

Mesoscutum, scutellum and axillae strongly reticulated, with pentagonal and hexagonal cells with

a few distinct ridges; mesoscutum shows 42 pairs of long setae (Fig. 2); scutellum with 4 long setae,

50 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Figures 1-3. Encarsia narroi. Figure 1. Antenna. Figure 2. Mesoscutum in dorsal view, setae and

cells. Figure 3. Forewing.

longer than mesoscutum setae; axillae with 1 short seta on each one; scutellar sensilla widely separated,

more than twice their diameter. Abdominal tergites smooth at the center and finely reticulated laterally,

with 4—5 setae on each side starting from 4th tergites. Ovipositor shorter than metasoma, originating

between 2nd and 3rd tergite.

All tarsi 5-segmented, tibial spur is about 0.066—0.073 mm long and middle basitarsus is 1.4 times

more long than tibial spur. Forewing hyaline, varies between 0.80—0.89 mm long and about 2.3 times

longer than its widest point; disk area uniformly setose except for the area below the submarginal

vein which bears 10 setae (Fig. 3), and with a small asetose area below the stigmal vein.

Male.——Unknown.

Biology.—This species is a parasitoid of the pupa of an Aleurodicus sp. (Ho-

moptera: Aleyrodidae) collected on Bauhinia variegata and Hibiscus sp., A max-

imum of 64% parasitism was observed in one sample. A Signiphora sp., is also

associated with this species, which is probably a hyperparasitoid of whitefly (Po-

laszek 1992).

Distribution.—Mexico, Coahuila State.

Material examined.—Holotype 2 MEXICO, Parras, Coahuila, 1500 m a.s.1., 11 Feb 1995 and 2

paratypes are deposited in the entomological collection of the National Reference Centre for Biological

Control in Colima, Mexico.

DISCUSSION

Encarsia narroi is most similar to Encarsia coquilletii (Schauff et al. 1996),

but can be differentiated from EF. coquilletii by the following caracteristics: me-

2000 GOMEZ & GARCIA: NEW ENCARSIA SPECIES 51

soscutum with 5 pairs of setae in E. coquilletti, whereas E. narroi have 42 pairs

of setae; both species with body color brown, but scutellum in E. narroi is pale

yellow.

ACKNOWLEDGMENT

We thank the Universidad Autonoma Agraria “Antonio Narro”’ and CONACyT

for financial support and ECOSUR facilities allowing the completion of this paper,

Roishene C. Williams for identification the biological material and to Trevor Wil-

liams for comments on the manuscript.

LITERATURE CITED

Arredondo, H. C. 1995. Los parasitoides en el control bioldgico de mosquita blanca (Homoptera:

Aleyrodidae) en México. pp. 4-15. En Simposio Sobre Control Biol6gico de Mosquita Blanca,

organizado por SARH-DGSV-CNRF-CNCB-SMCB-ECOSUR, 9 Noviembre 1995. Tapachula,

Chiapas, México.

Gerling, D. 1990. Natural enemies of whiteflies: predator and parasitoids. pp. 147-185. In Gerling,

D. (ed.). Whiteflies: their bionomics, pest status and management. Intercept, Andover, Hants,

U.K.

Hennessey, R. D., H. C. Arredondo & L. A. Rodriguez. 1995. DistribuciOn geografica y huéspedes

alternos de parasitoides afelinidos de Bemisia tabaci (Homoptera: Aleyrodidae). Vedalia, 2: 61-75.

Myartseva, S. N. & V. A. Yasnosh. 1994. Parasites of Greenhouse and Cotton Whiterflies (Homoptera:

Aleyrodidae) in Central Asia. Entomol. Rev., 73: 1-11.

Polaszek, A.,G. A. Evans & E D. Bennett. 1992. Encarsia parasitoids of Bemisia tabaci (Hymenop-

tera: Aphelinidae, Homoptera: Aleyrodidae): a preliminary guide to identification. Bull. Ento-

mol. Res., 82: 375-392.

Rivnay, T. & D. Gerling. 1987. Aphelinidae parasitoids (Hymenoptera: Chalcidoidea) of whiteflies

(Hemiptera: Aleyrodidae) in Israel, with description of three new species. Entomophaga, 32:

463-475.

Schauff, M. E., G. A. Evans & J. M. Heraty. 1996. A pictorial guide to the species of Encarsia

(Hymenoptera: Aphelinidae) parasitic on whiteflies (Homoptera: Aleyrodidae) in North Amer-

ica. Proc. Entomol. Soc. Wash., 98: 1-35.

van Lenteren, J. C., H. J. W. van Roermund & S. Siitterlin. 1996. Biological control of greenhouse

whitefly (Trialeurodes vaporariorum) with the parasitoid Encarsia formosa: how does it work?

Biol. Cont., 6: 1-10.

Received 6 Jun 1998; Accepted 24 Aug 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 52-54, (2000)

A NEW SPECIES OF HOCKERIA WALKER FROM

MEXICO (HYMENOPTERA: CHALCIDIDAE)

JEFFREY A. HALSTEAD

296 Burgan Avenue, Clovis, California 93611

Abstract—Hockeria burdicki Halstead, NEW SPECIES, is described and illustrated based on

material from Mexico, and is compared to closely related congeneric species. This wasp is the

tenth species of Hockeria described from the Nearctic region.

Key Words.—Insecta, Hymenoptera, Chalcididae, Hockeria, Mexico.

Wasps of the genus Hockeria Walker are distributed worldwide and the genus

contains about thirty-five described species. Hosts are summarized by Halstead

(1990) and include antlion and owlfly larvae (Neuroptera), elasmid and tenthre-

dinid pupae (Hymenoptera), free-living Strepsiptera, dipteran pupae and, com-

monly, lepidopteran larvae and pupae. In the Nearctic region, three economically

important lepidopterous pests are parasitized by Hockeria spp.: the Nantucket Pine

Tip Moth (Rhyacionia frustrana (Comstock)), Ponderosa Pine Tip Moth (Rhva-

cionia zozana (Kearfott)), and the Western Grapeleaf Skeletonizer (Harrisina bril-

lians Barnes and McDunnough).

A new species of Hockeria is described from Mexico; the tenth for the Nearctic

region—exclusive of Neotropical Mexico (Peck 1963, Burks 1979, De Santis

1979, Halstead 1990). No biological or host information is known and its potential

as a biological control agent is, therefore, unassessed.

MATERIALS AND METHODS

Specimens were discovered while sorting and identifying material for biosys-

tematic studies on chalcidid wasps. The specimens were compared to reference

material of the other Nearctic Hockeria species. Drawings were made using a

microscope drawing-tube. Measurements were made with a micrometer grid. Mor-

phological terminology follows Gibson, Huber, & Woolley (1997). Types are de-

posited in California Academy of Sciences, San Francisco (CAS) and the United

States Museum of Natural History, Washington, D.C. (USNM).

HOCKERIA BURDICKI HALSTEAD, NEW SPECIES

(Figs. 1—3)

Types.—Holotype female: MEXICO. JALISCO: Chamela Research Station, 20

Aug 1986, M. Sanchez, Malaise trap (CAS). Allotype male: as above but 26 Sep—

8 Oct 1985, Parker & Griswold (CAS). Paratypes: 1 female and 7 males, same

locale as holotype; 1 female, 20 Aug 1986, M. Sanchez, Malaise trap (USNM);

3 males, 26 Sep—8 Oct 1985, Parker & Griswold (CAS); 1 male, 15—24 Apr 1986,

E D. Parker (CAS); 2 males, 13 May 1986, M. Sanchez, Malaise trap (CAS); 1

male, 17 Oct 1985, Malaise trap (USNM).

Description—Female (holotype, Fig. 1). Length. 5.0 mm. Color. Black with scape, pedicel, pro-

notum, mesoscutum except anterior margin and lateral lobe, axilla, tegula, scutellum, fore- and middle

coxa, fore- and middle femur, apices of tibia, tarsi, and ventral margin of gastral terga, orange-red.

Head. Sculptured with dense umbilicate setigerous punctures; setae pale; integument polished. Scrobal

2000 HALSTEAD: NEW HOCKERIA SPECIES 53

Figures 1-3. Hockeria burdicki, NEW SPECIES (lateral views). Figure 1. Female habitus. Figure

2. Antenna, male. Figure 3. Scutellum, male. Scale lines 1.0 mm.

cavity deeply concave, transversely carinate. Scape reaching anterior ocellus. Malar sulcus reaching

to near ventral margin of eye. Postorbital carina curving along posterior margin of eye to near apex

of occiput. Mesosoma. Thorax slightly convex in lateral view, sculpture and integument like head.

Mesopleuron, anterior to femoral depression, transversely carinate. Metapleuron with dorsal half punc-

tate; ventral half rugose, densely setose. Scutellum with posterior margin rounded, with two minute

upturned teeth, and the punctures separated from each other by 0.16-0.25 X puncture diameter. Pro-

podeum with submedian longitudinal carinae and a couple of vague transverse carinae between these,

elsewhere carinately reticulate with surface between carinae shiny and slightly rugose. Hind femur.

Large and oval, 1.6 X as long as high (lateral view); integument coriaceous; setose. Forewing. Clouded

from near apex of submarginal vein to apex of wing except for clear rectangular area below stigma.

Gaster. In lateral view oval, 2.5 X as long as high, apex subaccuminate. Tergum 1 0.40 X length of

gaster in dorsal view, smooth and polished except for lateral coriaceous and setose area. Other terga

coriaceous except for polished medial area of tergum 2.

Male (Allotype).—Length. 4.5 mm. Color. Black with apices of tibia, tarsi, and ventral margins of

metasomal terga, orange. Forewing. Evenly clouded, and with an orangish tint. Body. Like female

except as noted for color and the following: antenna robustly filiform (Fig. 2), flagellomeres 1.5 X as

long as wide; apex of scape not reaching anterior ocellus, separated from it by 1 X ocellar diameter;

scutellum convex (Fig. 3); tergum 1 sublaterally polished, punctate medially, remainder and other

terga coriaceous.

Diagnosis.—Hockeria burdicki (both males and females) is distinguished from

54 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

other New World Hockeria by the large, oval hind femur which is 1.6 X as long

as high. In other Nearctic Hockera, the hind femur is at least 2 X as long as high.

Females most closely resemble H. tenuicornis (Girault), and will key out to that

species in Halstead’s (1990) key to species. Females of H. burdicki are distin-

guished from those of H. tenuicornis by their large, oval hind femur, and the

forewing disc with a rectangular-shaped, unclouded area rather than an elliptical-

shaped area. Other distinguishing characters for H. burdicki females include the

black body color, the elongate abdomen with the apex pointed, tergum 1 polished

dorsally, a clouded pattern in the forewing, and the relatively large body size.

Males most closely resemble H. unipunctatipennis (Girault), and will key out to

that species in Halstead’s (1990) key to species. Males of H. burdicki are distin-

guished from those of H. unipunctatipennis by their large, oval femur, their black

rather than orange femora, and their small rather than large and protruding inter-

antennal lobe. Other distinguishing characters for H. burdicki males include the

black body color, the punctate and coriaceous tergum 1 (dorsal view), the rela-

tively large body size, flagellomeres 1.5 X as long as wide (Fig. 2), convex

scutellum (Fig. 3), and the clouded forewing which has an orangish tint. Only

the male of H. unipunctatipennis has a similarly colored forewing.

Variation.—Male. Length 4.5 to 5.4 mm. Female. One paratype with flagel-

lomere 1, mesepimeron, and dorsal half of mesopleuron, orange.

Distribution.—Mexico, Jalisco.

Host—Unknown.

Etymology.—The specific name, a noun in the genitive case from a modern

personal name, is in honor of Donald J. Burdick—a friend and mentor in ento-

mology.

Material Examined.—This species is known only from the type specimens.

ACKNOWLEDGMENT

I thank D. J. Burdick and K. J. Woodwich, at California State University,

Fresno, for mentoring and the use of laboratory facilities and drawing equipment;

museums and institutions for the use and loan of material; and anonymous re-

viewers for helpful comments on the manuscript.

LITERATURE CITED

Burks, B. D. 1979. Chalcididae. pp. 860-874. Jn Krombein, K. V. et al. (eds.). Catalog of Hymenoptera

in America north of Mexico. Volume I. Smith. Instit. Press., Washington, D.C.

De Santis, L. 1979. Catalago de los himenopteros calcidoideos de America al sur de los Estados

Unidos. Comision de Investigaciones Cientificas de la Provincia de Buenos Aires, La Plata,

Argentina, 1-488.

Gibson, G. A. P, J. T. Huber & J. B. Woolley. 1997. Annotated Keys to the Genera of Nearctic

Chalcidoidea (Hymenoptera). NRC Research Press, Ottawa, Ontario, Canada.

Halstead, J. A. 1990. Revision of Hockeria Walker in the Nearctic region with descriptions of males

and five new species (Hymenoptera: Chalcididae). Proc. Entomol. Soc. of Wash., 92: 619-640.

Peck, O. C. 1963. A catalog of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Canad. Entomol.

Suppl., 30: 1092 p.

Received 30 Jul 1998; Accepted 15 May 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 55-57, (2000)

ENTOMOGNATHUS FROM CHINA WITH DESCRIPTION

OF A NEW SPECIES (HYMENOPTERA: SPHECIDAE)

QIANG LI! AND JUNHUA HE?

‘Department of Plant Protection, Shandong Agricultural University,

Taian, Shandong, 271018, P. R. China

*Department of Plant Protection, Zhejiang Agricultural University,

Hangzhou, Zhejiang, 310029, P. R. China

Abstract—A key to the species of the genus Entomognathus Dahlbom from China is provided,

and a new species, Entomognathus (Koxinga) aneurytibialis, is described.

Key Words.——Insecta, Hymenoptera, Sphecidae, Crabroninae, Entomognathus, China.

The genus Entomognathus Dahlbom has been represented by 61 species of

small to medium size predatory solitary wasps, of which 12 occur in the Pa-

laearctic, 10 in the Oriental, 25 in the Ethiopian and 14 in the Nearctic and

Neotropical Regions. Bohart & Menke (1976) revised the genera of Sphecidae of

the world. They provided a key to the subgenera and listed 42 species of the

genus Entomognathus. Tsuneki (1947, 1967, 1968, 1972, 1976, 1977) studied the

species and provided a key for the identification of east Asian forms. Pulawski

(1978) keyed the species of the northwest Palaearctic Region. Wu and Zhou

(1996) revised the species from China.

Entomognathus includes 4 subgenera, of which the subgenus Koxinga only has

6 species. In the course of a study on the fauna of Crabroninae from China, we

recognize 4 species of Entomognathus, of which one belonging to the subgenus

Koxinga is new to science.

KEY TO THE SPECIES OF ENTOMOGNATHUS FROM CHINA

1. Mesopleuron with stemaulus and verticaulus ...................--.4-- 2

— Mesopleuron without sternaulus and verticaulus

2. Hind tibia very swollen; pronotal collar, prepectus, scutellum and metan-

otum with large yellow spots. Sichuan, Zhejiang, Taiwan ..........

shee eesti i CAE ESSE SO tae Shwe Be) cob ewes E. (Koxinga) siraiya Pate

— Hind tibia normal; pronotal collar, prepectus, scutellum and metanotum

black. Yunnan ........ E. (Koxinga) aneurytibialis Li et He, new species

3. Propodeal enclosure enclosed by a narrow and shallow furrow; outer side

of hind tibia with fine spines. Neimenggu .......................-.

AE re Ae RRO ROIIR a Bay he E. (Entomognathus) sahlbergi (A. Morawitz)

— Propodeal enclosure enclosed by a broad and deep furrow; outer side of

hind tibia with coarse spines. Heilongjiang, Jilin, Neimenggu, Xinjiang,

1D ols ol Uta eed en. Oe A E. (Entomognathus) brevis (Vander Linden)

ENOTOMOGNATHUS (KOXINGA) ANEURYTIBIALIS, NEW SPECIES

(Figs. 1-4)

Type.—Holotype, male, Menga, 1050-1080 m, Xishuangbanna, Yunnan Province, China, 13 Oct

1958, S. Wang; deposited: the Insect Collections of Institute of Zoology, Academia Sinica.

56 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Figures 1-4. Entomognathus (Koxinga) aneurytibialis. Figure 1. Head, frontal view. Figure 2.

Antenna. Figure 3. Hind tibia, lateral view. Figure 4. Pygidial plate, dorsal view (scale line for Figures

1 and 3: 0.54 mm; for Figures 2 and 4: 0.48 mm).

Body length 3.5 mm. Black; mandible except apex, anterior margin of clypeus medially, apical half

of antenna beneath, pronotal lobe, trochanters to tarsi of legs, fore coxa, mid and hind coxae at apex

and abdomen at apex reddish yellow; mandible at apex, antenna above and tegula dark brown; basal

half of antenna beneath yellow; wing veins brown or dark brown. Eyes and body covered by white

short erect hairs. Head shiny; anterior margin of clypeus (Fig. 1) slightly prominent medially; upper

portion of frons densely punctate, with a shallow median furrow; vertex sparsely and finely punctate,

without orbital foveae; occipital carina flanged and foveate, contiguous to hypostomal carina; head

length: head width: postocellar distance: ocellocular distance = 59:100:14:15. Mandibles apically

simple, acuminate, externoventral margin notched medially. Antennae (Fig. 2), relative length of scape

: pedicel : flagellomere I:II:TH:IV:V = 33:8:5:5.5:5.5:5.5:5.5. Thorax shiny; pronotal collar densely

and finely punctate; scutum, scutellum and metanotum sparsely punctate; mesopleuron and metapleu-

ron densely and finely punctate; propodeum densely punctate, base of propodeal enclosure with short,

longitudinal rugae; posterior side of propodeum with broad, shallow median furrow, without rugae

and carinae; lateral side of propodeum without rugae and carinae, with lateral propodeal carina. Fore-

wing with R, extending beyond apex of marginal cell. Hind tibia (Fig. 3) and tarsi normal. Abdomen

shiny, sparsely punctate; tergite I, length: width at posterior margin = 58:60; pygidial plate (Fig. 4)

densely and coarsely punctate (Figs. 1—4).

Diagnosis.—This new species is related to E. (K.) siraiya Pate. It can be dis-

tinguished from the latter by the characters of hind tibia not swollen, posterior

side and lateral side of propodeum without rugae and carinae, the shape of anterior

margin of clypeus (Fig. 1), no yellow spot on thorax except the pronotal lobe

which are reddish yellow, abdomen black except apex, coloration of legs as out-

lined in the text, and a rather smaller body.

Etymology.—The name is derived from one of its main characters: an- = not

or without (originated from Greek words); -eury- = broad (originated from Greek

words); -tibialis = tibial (originated from Greek words also). The hind tibia of

this species is normal, not swollen and broad.

Material Examined.—See Type.

ACKNOWLEDGMENT

We are grateful to Professor Yanru Wu (Institute of Zoology, Academia Sinica,

Beijing) for providing us with specimens deposited in the Insect Collections of

Institute of Zoology, Academia Sinica.

2000 LI & ME: CHINESE ENTOMOGNATHUS 57

LITERATURE CITED

Bohart, R. M. & A. S. Menke. 1976. Sphecid wasps of the world, a generic revision. Univ. of

California Press, Berkeley, Los Angeles, London, pp. 1-695.

Pulawski, V. V. 1978. The family Sphecidae. pp. 73-279. In Classification of the insects in the Eu-

ropean part of the USSR. Volume 3. Hymenoptera. Part I. Zoological Institute Press. Leningrad.

[In Russian. ]

Tsuneki, K. 1947. On the wasps of the genus Crabro s. 1. from Hokkaido, with descriptions of new

species and subspecies (Hymenoptera). J. Fac. Sci. Hokkaido Univ., 9: 397—435.

Tsuneki, K. 1967. Further studies on the fossorial Hymenoptera from Manchuria. Etizenia, 23: 1-17.

Tsuneki, K. 1968. Studies on the Formosan Sphecidae (V), the subfamily Crabroninae (Hymenoptera)

with a key to the species of Crabronini occurring in Formosa and Ryukyus. Etizenia, 30: 1-34.

Tsuneki, K. 1972. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei, 280.

Sphecidae (Hymenoptera). IV—V. Acta Zoologica Academiae Scientiarum Hungaricae, 18: 147-

232.

Tsuneki, K. 1976. A fourth contribution to the knowledge of Sphecidae (Hymenoptera) of Manchuria,

with remarks on some species of the adjacent regions. Knotyu, Tokyo, 44: 288-310.

Tsuneki, K. 1977. H. Sauter’s Sphecidae from Formosa in the Hungarian Natural History Museum

(Hymenoptera). Annales Historico-Naturales Musei Nationalis Hungarici, Tomus, 69: 261-296.

Wu, Y. & Q. Zhou. 1996. Economic insect fauna of China, Fasc. 52. Hymenoptera: Sphecidae. Science

Press, Beijing [In Chinese.], pp. 1-197.

Received 1 Dec 1997; Accepted 24 Aug 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 58-70, (2000)

HISTORICAL REVIEW OF THE GENERA ALEIODES AND

ROGAS IN MEXICO, WITH A REDESCRIPTION OF

ALEIODES CAMERONI (HYMENOPTERA: BRACONIDAE)

H. DELFIN G.! AND R. A. WHARTON?

‘Facultad de Medicina Veterinaria y Zootecnia, Universidad Aut6noma de

Yucatan, Apartado postal 4-116 Itzimna, Mérida, Yucatan, México

*Department of Entomology, Texas A&M University,

College Station, Texas 77843, U.S.A.

Abstract—A brief nomenclatural history is provided for the species of Aleiodes and Rogas

previously recorded from Mexico. Variation in Aleiodes cameronii (Dalla Torre) is detailed to

facilitate comparisons with other species, and its distribution in Mexico discussed. The presence

of dorsal abdominal pits in male Aleiodes is reviewed, with new records for the dispar Curtis

species group from Africa and Australia.

Key Words.—distribution, abdominal pits, Rogadinae, parasitoids.

Aleiodes Wesmael, 1838 is a cosmopolitan genus of parasitic wasps, all species

of which are endoparasitoids of Lepidoptera. Species of Aleiodes have frequently

been placed in Rogas Nees, 1834, though both generic names are now considered

valid (van Achterberg 1991, Shaw 1997). Described species and known hosts are

listed by Shenefelt (1975) and Fortier (1997). Shaw et al. (1997) recently provided

a preliminary key to species groups for the Nearctic region.

In general, members from the western Palearctic and Nearctic regions are well

known, but the Afrotropical, eastern Palearctic and Neotropical species of Aleio-

des are poorly known. For example, there are 75 described species from the New

World and close to 200 undescribed species (Shenefelt 1975, Marsh 1979, Shaw

et al. 1997), with most of these described species from the Nearctic region. Except

for the recent description of 7 species and a redescription of 12 species (Shaw

1993; van Achterberg & Penteado-Dias 1995; Shaw et al. 1997, 1998a, b; Marsh

& Shaw 1998), most of the Neotropical species of Aleiodes are known only from

limited original descriptions. The identity of these Neotropical species is further

complicated by previous nomenclatural confusion involving application of generic

names. Thus, prior to clarification of the status of Rogas and Aleiodes by van

Achterberg (1982, 1991), some (e.g., Shenefelt 1975) or all (e.g., Labougle 1980)

of the species described in Aleiodes were placed in Rogas. Similarly, following

van Achterberg’s (1982) earlier work, many species formerly placed in Rogas

were automatically transferred to Aleiodes by authors of regional lists and similar

publications, without sufficient evidence to validate such changes. Correct place-

ment of the described species will require a critical assessment of the applicability

of the names Rogas and Aleiodes to the New World fauna, as is currently being

done for Nearctic Aleiodes in the excellent treatments by Shaw et al. (1997, 1998a,

b) and Marsh and Shaw (1998).

Our collections from Mexico indicate that the Aleiodes fauna is highly diverse,

perhaps equivalent to the fauna of America north of Mexico. Yet, only a few

species have previously been recorded from Mexico, and specific localities have

rarely been mentioned. To provide a baseline for work on the biodiversity of the

2000 DELFIN & WHARTON: REVIEW OF ALEIJODES a0

Mexican fauna, and to clarify the status of certain names, the 22 species of Al-

eiodes and Rogas previously recorded from Mexico are listed below. We have

used a catalog format, with information on their current status, previously re-

corded distribution within Mexico, prior combinations, and recent catalog listings

(which should be consulted for additional localities outside Mexico). For localities

listed by Cameron (1887), we have followed Selander and Vaurie (1962), thus

correcting earlier records suggesting that the type locality of Aleiodes cameronii

(Dalla Torre) was either in Veracruz or Texas. New combinations are made for a

few of the species, when sufficiently diagnostic characters were mentioned in the

original descriptions, or authoritatively determined material was available.

Abbreviations for specimen depositories are as follows: BMNH, British Mu-

seum of Natural History, London; CAS, California Academy of Sciences, San

Francisco; CER—-UADY, Coleccién Entomolégica Regional de la Universidad

Auténoma de Yucatan, Mérida; IB—UNAM, Instituto de Biologia, Universidad

Aut6énoma de México, México City; ANSP, Philadelphia Academy of Natural

Sciences, Philadelphia; TAMU, Texas A&M University Insect Collection, College

Station; ZMPA, Polish Academy of Sciences, Warsaw.

Additionally, we redescribe Aleiodes cameronii (Dalla Torre), and use this op-

portunity to discuss distribution and characterization of the pulchripes Wesmael

and dispar Curtis species groups. Terminology for the description follows van

Achterberg (1991, 1993) and, for wing venation, Sharkey and Wharton (1997)

and Shaw et al. (1997). Maximum width of head is measured in dorsal view

across the eyes and across the temples immediately posteriorad the eyes. Quan-

titative values are based on a minimum of five specimens when no sexual di-

morphism was evident. Voucher specimens of A. cameronii are deposited in

TAMU.

SPECIES OF ALEIODES AND ROGAS PREVIOUSLY RECORDED FROM MEXICO

atriceps Cresson.

Aleiodes atriceps Cresson, 1869 Trans. Am. Ent. Soc. 2:380. Type locality: ‘““Mex-

ico.”’ Holotype deposited ANSP (#1662.1).

Rhogas atriceps, Fox 1895:3.

Rogas atriceps, Shenefelt 1975:1218.

Dimorphomastax peculiaris Shenefelt, 1979:133.

Distribution MEXICO. BAJA CALIFORNIA SUR: Margarita Is. (Fox 1895).

Remarks.—Shaw et al. (1998a) synonymized peculiaris with atriceps, uniting

males with an enlarged, tooth-like outgrowth from the base of the mandible with

more normal looking females. Shaw et al. (1998a) also transferred atriceps back

to Aleiodes. Current valid combination: Aleiodes atriceps Cresson.

aztecus Cameron.

Rhogas aztecus Cameron, 1905 Trans. Am. Ent. Soc. 31:385. Type locality:

‘““Mexico.”’ Holotype depository unknown.

Rogas aztecus, Shenefelt 1975:1219.

Distribution—MEXICO (Cameron 1905); no specific localities published to date.

Remarks.——Generic placement needs verification. Current valid combination:

Rogas aztecus Cameron.

60 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

burrus Cresson.

Aleiodes burrus Cresson, 1869 Trans. Am. Ent. Soc. 2:381. Type locality: ‘“‘Illi-

nois.””’ Holotype deposited ANSP (#1670.1).

Rhogas burrus, Cameron 1887:224.

Rogas burrus, Muesebeck and Walkley 1951:170, Marsh 1979:179, Shenefelt

1975:1220.

Distribution—MEXICO (Cresson 1869); no specific localities published to

date.

Remarks.—Transferred back to Aleiodes on basis of material examined during

this study. Current valid combination: Aleiodes burrus Cresson, NEW STATUS.

cameronii Dalla Torre.

Rhogas mexicanus Cameron, 1887 Biol. Centr.-Am., Hym 1:389. Type locality:

“Mexico, Presidio.’’ Holotype deposited BMNH (#3.c.235).

Rhogas cameronii Dalla Torre, 1898 Cat. Hym. 4:216.

Rogas cameronii, Shenefelt 1975:1220.

Aleiodes cameronii, Shaw et al. 1997:17.

Distribution—-MEXICO. SINALOA: Presidio (Cameron 1887). Shaw et al.

(1997) record this species as occurring from southern U.S. through Mexico to

Costa Rica, but do not give specific localities within Mexico. Additional localities

are given below under the redescription of this species.

Remarks.—The name cameronii was proposed by Dalla Torre (1898) as a re-

placement name for mexicanus Cameron, 1887 (not mexicanus Cresson, 1869).

Both nominal species are valid in Aleiodes. Current valid combination: Aleiodes

cameronii (Dalla Torre).

enderleini Shenefelt: see vaughani.

fascipennis Cresson.

Aleiodes fascipennis Cresson, 1869 Trans. Am. Ent. Soc. 2:378. Type locality:

‘““Mexico.”” Holotype deposited ANSP (#1665).

Rhogas fasciipennis, Dalla Torre 1898:218 (emendation).

Pelecystoma fasciipennis, Shenefelt 1975:1207.

Distribution MEXICO (Cresson 1869); no specific localities published to

date.

Remarks.—Current valid combination: Rogas fascipennis (Cresson), NEW

STATUS.

ferrugineus Enderlein.

Rhogas ferrugineus Enderlein, (1918) 1920 Arch. Naturgesch. 84 A (11):156.

Type locality: “‘Mexiko, Chiapas.’’ Holotype deposited ZMPA.

Rogas ferrugineus, Shenefelt 1975:1229.

Distribution MEXICO. CHIAPAS (Enderlein 1920). We have seen additional

material of this widespread species from the following Mexican localities: MEX-

ICO. AGUASCALIENTES: 12.8 km NE of Aguascalientes; Calvillo. CHTHUA-

HUA: Santa Clara Canyon, 5 mi W Parrita. COAHUILA: 39 km S of Agua Nueva,

1.6 km SE of Saltillo. COLIMA: 14.4 & 16 km NE of Comala. DISTRITO FED-

ERAL:; Primary and Tertiary Secc. Bosque de Chapultepec. GUANAJUATO: Ce-

2000 DELFIN & WHARTON: REVIEW OF ALEJODES 61

laya; El Copal; Inchamacuaro; Las Trancas; Purisima de Bustos; Roque; San Bar-

tolomé; Tarandacuaro; Tierra Blanca. GUERRERO: Iguala; 28.8 km S of Chil-

pancingo; 5 km W of Tixtla; 9.6 km NE of Tixtla. HIDALGO: Tulancingo. JAL-

ISCO: Guadalajara; 4.8 km SE of Plan de Barrancas; 10 km NE of Jalostotitlan;

Sierra de Manantlan Lab. Mat. Las Joyas; Arroyo Las Joyas; Rancho La Quinta;

Teocaltiche; Zapopan. MEXICO: Chapingo; Tenango del Aire; Texcoco; Tonatico.

MICHOACAN: Morelia; La Huerta; 4.8 km E of Carapan; 17.6 km W of Hidalgo;

19.2 km NW of Zitacuaro. MORELOS: Cuernavaca; 2.5 km N of Huautla Esta-

ci6n CEAMISH; Huejotengo; Tepoztlan; Yautepec. NAYARIT: Jess Maria. NUE-

VO LEON: Linares. OAXACA: 12.8 km NE of El Punto; Oaxaca; 25.6 km NW

of Totolapan; Puerto Escondido. QUERETARO: 11.2 km N of Querétero. SAN

LUIS POTOSI: 17 km NE of Ciudad del Maiz; 59.2 km S of San Luis Potosi;

13.9 km S of Santa Maria del Rio. SINALOA: Concordia. TAMAULIPAS: Hidalgo

Conrado Castillo; Rio Soto La Marina; Soto La Marina: VERACRUZ: 19 km NW

of Ciudad Mendoza; 12.8 km S of Jalapa; Cérdoba; Orizaba. ZACATECAS: Con-

cepcion del Oro; 6.4 km NE of Concepcién del Oro.

Remarks.—Transferred to Aleiodes on basis of material examined during this

study. Current valid combination: Aleiodes ferrugineus (Enderlein), NEW COM-

BINATION.

fumialis Shenefelt.

Rhogas fumipennis Cameron, 1887 Biol. Centr.-Am., Hym. 1:389. Type locality:

‘““Mexico.”’ Holotype deposited BMNH.

Rogas fumialis Shenefelt, 1975:1230.

Distribution — MEXICO (Cameron 1887); no specific localities published to

date.

Remarks.—The name fumialis was proposed by Shenefelt (1975) as a replace-

ment name for fumipennis Cameron, 1887 (not fumipennis Cresson, 1869). Both

nominal species belong in Aleiodes, though only fumialis is currently considered

valid. Current valid combination: Aleiodes fumialis (Shenefelt), NEW COMBI-

NATION.

fumipennis Cameron: see fumialis.

fumipennis Cresson: see texanus.

fusciceps Cresson.

Aleiodes fusciceps Cresson, 1869 Trans. Am. Ent. Soc. 2:382. Type locality:

‘“‘Mexico.”” Holotype deposited ANSP (#1673).

Rhogas fusciceps, Dalla Torre 1898:218.

Pelecystoma fusciceps, Shenefelt 1975:1207.

Distribution MEXICO (Cresson 1869); no specific localities published to

date.

Remarks.—Current valid combination: Rogas fusciceps (Cresson), NEW STA-

TUS.

laphygmae Viereck.

Rogas laphygmae Viereck, 1912 Proc. U.S. Natn. Mus. 43:581. Type locality:

Texas, Brownsville. Holotype deposited USNM (#15012). Muesebeck and

Walkley 1951:171; Shenefelt 1975:1236; Marsh 1979:180.

62 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Distribution MEXICO. NUEVO LEON: Marin. This species has been re-

corded from southern U.S. and Central America, but we know of no specific,

published records from Mexico. This is a widespread species, however, and we

list here a specific record for reliably determined specimens from Mexico in

TAMU.

Remarks. Current valid combination: Aleiodes laphygmae (Viereck), NEW

COMBINATION.

melanocephalus Cameron.

Rhogas melanoce phalus Cameron 1887 Biol. Centr.-Am., Hym. 1:391. Type lo-

cality: ““Mexico, Cordova.”’ Holotype deposited BMNH.

Macrostomion melanoce phalus, Szépligeti, 1904:82.

Pelecystoma melanoce phalum, Enderlein, 1920:148; Shenefelt, 1975:1208.

Distribution —MEXICO. VERACRUZ: Cérdoba (Cameron 1887).

Remarks.—Current valid combination: Rogas melanoce phalus Cameron. Ge-

neric placement needs verification.

mexicanus Cameron: see cameronii.

mexicanus Cresson.

Aleiodes mexicanus Cresson, 1869 Trans. Am. Ent. Soc. 2:378. Type locality:

‘“‘Mexico.”” Holotype deposited ANSP (#1658).

Rhogas mexicanus, Dalla Torre 1898:216, 220.

Rogas mexicanus, Shenefelt 1975:1238.

Distribution MEXICO. CHIAPAS, SINALOA, VERACRUZ (Shaw 1993). We

have seen material from the following Mexican localities in addition to those

listed by Shaw (1993): MEXICO. CHIAPAS: 6.4 km SW of Simojovel. MEXICO:

Xilitla. MORELOS: Yautepec. SAN LUIS POTOSI: 16 km NE of entronque Ray-

6n-Cardenas. TAMAULIPAS: 3.5 km W of Gomez Farias. TABASCO: Teapa.

VERACRUZ: Cordoba; Fortin de las Flores; Puente Nacional 7.2 km SE of Rin-

conada; 40 km S of Acayucan.

Remarks.—Shaw (1993) returned mexicanus to Aleiodes, including it in the

subgenus Eucystomastax along with three other Neotropical species. Current valid

combination: Aleiodes mexicanus Cresson.

molestus Cresson.

Rogas molestus Cresson 1872, Trans. Am. Ent. Soc. 4:188. Type locality: ‘““Mex-

ico.”” Holotype deposited USNM (#1625). Muesebeck and Walkley 1951:171,

Shenefelt 1975:1239; Marsh 1979:180.

Rhogas molestus, Dalla Torre 1898:221.

Aleiodes molestus, Shaw et al. 1998a:70.

Rogas rufocoxalis Gahan 1917:207.

Distribution MEXICO. PUEBLA-OAXACA (Labougle 1980). We have seen

additional material of this widespread species from the following Mexican local-

ities: MEXICO. AGUASCALIENTES: 12.8 km NE of Aguascalientes. CHIAPAS:

Las Rosas; Rancho Sanchez; San Cristobal de las Casas. CHIHUAHUA: Chihua-

hua. JALISCO: Rancho La Quinta; Teocaltiche. MEXICO: Santa Maria; Tonatico;

Valle de Bravo. MICHOACAN: 7.2 km N of Cheran; Jungapeo. NAYARIT: Jests

2000 DELFIN & WHARTON: REVIEW OF ALEIJODES 63

Maria. NUEVO LEON: 8 km S of Linares. TAMAULIPAS: Hidalgo, Conrado

Castillo. VERACRUZ: Veracrus.

Remarks.—Shaw et al. (1998a) synonymized ruficoxalis with molestus. Current

valid combination: Aleiodes molestus (Cresson).

nigriceps Enderlein: see vaughani.

nigripes Enderlein.

Pelecystoma nigripes Enderlein (1918) 1920 Arch. Naturgesch. 84 A 11:148.

Type locality: ““Mexiko, Chiapas.’ Holotype deposited ZMPA. Shenefelt 1975:

1208.

Aleiodes nigripes, van Achterberg 1991:61.

Distribution MEXICO. CHIAPAS (Enderlein 1920).

Remarks. Current valid combination: Aleiodes nigripes (Enderlein).

nigristemmaticum Enderlein.

Rhogas nigristemmaticum Enderlein (1918) 1920 Arch. Naturgesch. 84 A (11):

156. Type locality: ““Mexiko, Chiapas.”” Holotype deposited ZMPA.

Rogas nigristemmaticum, Wolcott 1948:759, Shenefelt 1975:1240.

Aleiodes nigristemmaticum, Marsh and Shaw (1998).

Distribution -MEXICO. CHIAPAS (Enderlein 1920). We have seen one ad-

ditional Mexican specimen from the following locality: MEXICO. OAXACA: 4.2

km NW of El Cameron.

Remarks——Current valid combination: Aleiodes nigristemmaticum (Enderlein).

ornatus Cresson.

Aleiodes ornatus Cresson, 1869 Trans. Am. Ent. Soc. 2:380. Type locality: “‘Mex-

ico.”’ Holotype deposited ANSP (#1666).

Rhogas ornatus, Dalla Torre 1898:221.

Pelecystoma ornatus, Shenefelt 1975:1208.

Distribution MEXICO (Cresson 1869); no specific localities published to

date.

Remarks. Current valid combination: Rogas ornatus (Cresson), but the ge-

neric name Triraphis Ruthe, 1855 may be more appropriate for this species.

peculiaris Shenefelt: see atrice ps.

pedalis Cresson.

Aleiodes pedalis Cresson, 1869 Trans. Am. Ent. Soc. 2:379. Type locality: ‘““Mex-

ico.”’ Holotype deposited ANSP (#1664).

Rhogas pedalis, Dalla Torre 1898:221.

Rogas pedalis, Shenefelt 1975:1242.

Distribution MEXICO (Cresson 1869); no specific localities published to

date.

Remarks.—Shaw et al. (1997) transferred pedalis back to Aleiodes. Current

valid combination: Aleiodes pedalis Cresson.

rossi Marsh and Shaw.

Aleiodes rossi Marsh and Shaw, 1997 J. Hym. Res. 6:32. Type locality: Texas,

Brownsville. Holotype deposited CAS.

64 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Distribution MEXICO. SAN LUIS POTOSI: E| Salto (Shaw et al. 1997).

Remarks.—Current valid combination: Aleiodes rossi Marsh and Shaw.

rufocoxalis Gahan: see molestus.

scriptipennis Enderlein.

Heterogamus scriptipennis Enderlein, (1918) 1920 Arch. Naturgesch. 84 A (11):

152. Type locality: ‘““Mexiko, Chiapas.’’ Holotype deposited ZMPA, Shenefelt

1975:1202.

Distribution—MEXICO. CHIAPAS (Enderlein 1920).

Remarks.—Current valid combination: Aleiodes scriptipennis (Enderlein),

NEW COMBINATION. The new combination is based on the current treatment

of Heterogamus as a synonym of Aleiodes (van Achterberg 1991:24), but verifi-

cation is needed.

sonorensis Cameron.

Rhogas sonorensis Cameron, 1887 Biol. Centr.-Am., Hym. 1:390. Type locality:

‘“‘Mexico, Northern Sonora.” Holotype deposited BMNH (#3.c.236).

Rogas sonorensis, Shenefelt 1975:1251.

Distribution —MEXICO: northern SONORA (Cameron 1887).

Remarks.—Current valid combination: Aleiodes sonorensis (Cameron), NEW

COMBINATION. The combination proposed here is based on the original de-

scription only, and needs verification.

texanus Cresson.

Aleiodes texanus Cresson, 1869 Trans. Am. Ent. Soc. 2:378. Type locality: ‘“Tex-

as.”” Holotype deposited ANSP (#1655.1).

Rhogas texanus, Cresson 1887:224

Rogas texanus, Cresson 1872:188; Shenefelt 1975:1254.

Heterogamus texanus, Ashmead 1889:632.

Aleiodes fumipennis Cresson, 1869:378.

Distribution —northem MEXICO (Shaw et al. 1998b); no specific localities

published to date.

Remarks.—Shaw et al. (1998b) synonymized fumipennis with texanus, and

transferred texanus back to Aleiodes. Current valid combination: Aleiodes texanus

Cresson.

vaughani Muesebeck.

Rogas vaughani Muesebeck, 1960 Ent. News 71:257. Type locality: Nicaragua,

Managua Holotype deposited USNM (#65047).

Rhogas nigriceps Enderlein, 1920:155.

Rogas enderleini Shenefelt, 1975:1227.

Aleiodes vaughani, Shaw et al. 1997:33.

Distribution MEXICO (Shaw et al. 1997); no specific localities published to

date, but we have one Mexican specimen from the following locality: MEXICO.

VERACRUZ: Estacion de Biologia Tropical Los Tuxtlas.

Remarks.—Shenefelt (1975) renamed nigriceps Enderlein. The replacement

name was unnecessary because he transferred nigriceps Wesmael back to Aleiodes

while retaining nigriceps Enderlein in Rogas. Both nominal species are now in

2000 DELFIN & WHARTON: REVIEW OF ALEIODES 65

Aleiodes, but following the synonymy of vaughani with nigriceps (Shaw et al.

1997), an older name is now available. Current valid combination: Aleiodes

vaughani (Muesebeck).

vestitor Say.

Bracon vestitor Say, 1832 Boston J. Nat. Hist. 1:254. Type locality: ‘““Mexico.”’

Type: lost.

Rogas vestitor, Muesebeck 1925:82.

Distribution MEXICO (Say 1832); no specific localities published to date.

Remarks.——The generic assignment of this species is questionable, since the

type has been lost for about 150 years and the description is vague. Current valid

combination: Rogas vestitor (Say).

Aleiodes cameronii (Dalla Torre)

Redescription.—T otal length: female, 6.5—9.2 mm; male, 5.5—7.0 mm; specimens from Yucatan at

large end of spectrum. Head: Antennae with 63-69 segments, with no apparent sexual dimorphism

in length or number of flagellomeres; second antennal segment square, as long as wide; apical fla-

gellomere with prominent spine; first flagellomere short, 0.62—0.67 X outer (shortest) length of scape;

ocellar field very large, width of head at temples 2.48—2.73 X width of ocellar field, ocelli nearly

touching eye, ocello-ocular line 0.08—0.13 X length of lateral ocellus, eye very large, with deep

emargination, in dorsal view width of head at eyes 1.14—-1.21 X width at temples; vertex finely

granular, with oblique striae near ocelli; frons smooth; face occasionally with a weakly protruding,

median triangle extending from clypeus to just dorsad middle of face, where it narrows to the median

ridge that extends between bases of antennae (the interantennal carina), face transversely to obliquely

strigose laterally, weakly to distinctly longitudinally strigose within triangle (or in an equivalent region

of the face when an elevated triangular area is not apparent); malar space short, closely strigose, 0.10—

0.13 X eye height, 0.56—-0.66 X basal width of mandible, malar suture absent; clypeus irregularly

rugulose-punctate, 2.18—2.37 X wider than high (slightly but distinctly taller in specimens from Yu-

catan: 2.00 X wider than high), not protruding; cyclostome opening 1.22 X wider than high; occipital

carina usually complete above, though often weak and occasionally (10%) absent mid-dorsally, well-

developed laterally, becoming indistinct ventrally, where it either terminates just short of hypostomal

carina, or gives off weak, irregular striae, which sometimes (<< 10%) reach hypostomal carina distinctly

removed from base of mandible, occipital carina thus does not extend to hypostomal carina as a well-

developed ridge; mandible short, length along dorsal surface about 0.80 X width at base; gena not

protruding in frontal view; maxillary palp 2.2 X longer than labial palp, third and sixth maxillary

palpomeres subequal in length, fourth slightly longer, third maxillary palpomere broadest, gradually

widening from base to apex, second labial palpomere not obviously dilated, but distinctly broader than

the slender third and fourth palpomeres. Wings: Fore wing: 2.2—2.48 X longer than mesosoma; ve-

nation as follow: M+CU sinuate, 1.83-1.91 X longer than 1M+1RS; 1M evenly curved; 1RS 0.42-—

57 X length of parastigma; (RS+M)a straight; (RS+M)b tubular, depigmented, short, 0.44—0.67 X

length of 2RS; 2RS tubular, weakly to sharply bent at posterior fourth, 0.62—0.78 X length of 3RSa;

3RSa 1.58-1.92 X longer than r (r quite variable in length, even within populations); r-m straight to

weakly bowed, tubular, depigmented; 2M straight, 1.23-1.37 x longer than 3RSa; m-cu slightly

curved; Icu-a distad 1M by 0.32-0.60 X length of Icu-a; 1CUa 0.12-0.22 X length of 1CUb; 2CUa

short, 0.54—0.72 X length of m-cu; 3-1A present. Hind wing: Vein M+CU straight, 0.93-1.0 X length

of 1M; 1M slightly arched; m-cu nearly always (96%) present, short to very short, never extending

more than half way to wing margin, postfurcal to r-m; 2-1A present; RS and 2M complete but not

tubular, RS diverging from anterior margin of wing at its basal 0.3, distal 0.7 of marginal cell thus

distinctly widening. Mesosoma: Pronotum mid-dorsally twice as long as shortest distance between

occipital carina and lateral ocellus; dorsally with thin, low carina along anterior margin, this bordered

posteriorly by shallow, narrow, crenulate groove, dorsal surface otherwise slightly uneven; pronotum

shagreened to weakly granular dorsally and antero-dorsally in lateral view, laterally with striae radi-

ating dorsally, posteriorly, and ventrally from small, polished, smooth to weakly sculptured spot; angle

between pronotum and anterior declivity of mesoscutum slightly more than 135°. Notaulus very weakly

66 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

impressed anteriorly, sometimes barely discernible ending in weak, broad, median depression poste-

riorly; mesoscutum granular/shagreened, except median depression rugose to weakly rugulose; scu-

tellar sulcus irregularly strigose, often without clearly defined central carina, short, 0.36-0.43 X median

length of scutellum; scutellum nearly smooth, polished medially, with only a trace of shagreened

sculpture, weakly strigose-punctate laterally; posterior margin of mesonotum forming an unsculptured,

polished band. Median longitudinal carina of propodeum distinctly elevated and visible in lateral view

only on anterior 0.25, otherwise variable: complete in nearly all specimens examined from U.S., but

weak to absent over posterior half in most specimens from Yucatan and Oaxaca; propodeum varying

from uniformly granular or nearly so (rugulose only along midline posteriorly) to granular antero-

laterally grading to densely but finely granular-rugose medially and posteriorly. Mesopleuron with

small smooth, polished area medially extending to posterior margin above speculum then dorsally

along posterior margin to wing base, otherwise striate to strigose dorsally, sometimes very strongly

so and punctate to weakly striate over ventral half; precoxal sulcus absent. Metapleuron weakly punc-

tate medially, rugose dorsally and along ventral margin. Inner spur of hind tibia short, 0.48-0.52 x

length of basitarsus. Posterior tarsal claw pectinate throughout, teeth large, those in middle nearly as

tall as apical claw. Metasoma: First tergite with basal triangle well developed, extending onto dorsal

surface from anterior declivity. T1, T2, and basal 0.75—0.95 of T3 aciculate, remaining terga largely

smooth and without median carina; median carina usually extending from basal triangle of T1 to

middle of T3, gradually merging with surrounding sculpture over apical half of T3, carina usually

well developed throughout, rarely (15%) absent on T3 and/or weakly developed posteriorly on T2,

equally rarely with carina complete to posterior end of T3. Lateral lobe of dorsope distinctively

expanded, carinate, but short, shorter than distance from end of carina to spiracle. Lateral margin of

T3 sharp, lateral margin of T4—T6 rounded. Median triangle at base of T2 small, largely hidden by

well-developed median distal lobe of T1. T2/T3 suture crenulate, distinctly impressed. Males with

distinct pits medially on T4—6 (never present on T7 in our material, though T7 and sometimes T8

more densely setose medially); pit on T4 absent in one-third of specimens, when present, always

smaller than pits on TS and T6, T4 pit oval to heart-shaped, divided at extreme base and extending

internally at antero-lateral corners; T5 and T6 with larger but variably sized pits, width of pit on TS

0.09-0.29 X width of tergum; all pits densely setose. Tl 1.27—1.40 X longer than T2, T2 1.24—1.38

X longer than T3,T2 + T3 1.27-1.40 X longer than T1. Females with hypopygium truncate; ovipositor

about 1.2 X longer than hypopygium, straight, with well-developed node subapically, strongly nar-

rowed medially, and strongly dilated basally, setae on ovipositor sheath longer, denser on dorsal half,

equal in length to depth of sheath. Color: Orange to pale yellow; flagellum and stemmaticum black;

scape, pedicel, and dorsal side of telotarsus dark brown; wings hyaline, stigma yellow; ovipositor

sheath brown apically, yellow basally.

Biology.—Unknown.

Material examined —MEXICO. OAXACA: 17 km N of Miltepec, 11 July 1973, Mastro & Schaffner,

1 female. YUCATAN: Reserva Especial de la Biosfera de Ria Lagartos, El Cuyo, 4-5 May 1994, H.

Delfin, 3 females, 5 males; 11 km N of Mérida, 27 May 1996, R. Wharton, 1 male; Xmatkuil, 18-—

28 June 1996, H. Delfin & FE Leon, 1 male. USA. ARIZONA. MARICOPA Co.: 12 km NE Apache

Jct., 17 July 1998, J. Oswald, 1 female. PIMA Co.: 17 km NW Arivaca, 18 July 1998, J. Oswald, 2

males. NEW MEXICO. LEA Co.: 32°24.8'N, 103°41.5'W, 1 August 1979, J. Delorme & C. McHugh,

1 female. TEXAS. BRAZOS Co.: Bryan, 25 May 1974, J. Schaffner, 1 male; College Station, 12-18

April 1978, J. Jackman, 1 female. BREWSTER Co.: Big Bend National Park, N Rosillos Mts, Buttrill

Springs, 22 March—8 April 1991, Wharton & Whitefield, 3 females, 3 males; same except 10 March

1991 and 23-25 April 1991, Wharton, Woolley & Zolnerowich, 2 females, 1 male. LASALLE Co.:

Chaparral Wdlf Mgmt Area, 29-30 September 1989, J. Schaffner, 1 female. RANDALL Co.: Palo Duro

Cyn, 14 June 1960, R. Fischer, 1 female. VAL VERDE Co.: 9 April 1960, 1 female.

Discussion.—Shaw et al. (1997) included cameronii in the pulchripes species

group, and gave a detailed diagnosis. Our material from Yucatan differs from this

diagnosis in certain details (notably development of propodeal carina and place-

ment of pits on the male terga), and we initially concluded that this was an

undescribed species. Subsequent examination of numerous specimens of came-

ronii, many of them determined by Shaw, has enabled us to characterize this

species more completely, and revise our initial assessment. For most characters,

2000 DELFIN & WHARTON: REVIEW OF ALEIJODES 67

Figure 1. Dorsal abdominal pits of Aleiodes cameronii from the same Pima Co., Arizona locality,

showing differences in size and placement of pits on terga 4-6. A. Pits on terga 5 and 6 only. Arrow

= terga 4 without pit. B. Pits on terga 4—6.

specimens from southern Mexico do not differ significantly from those collected

in the U.S., and variation within popuations is equivalent to that among popula-

tions from widely scattered localities. For example, pit size and placement on

male terga is variable, and the largest pits in our material occur on one specimen

from Yucatan and one from Arizona. In each case, other males from the same

locality have distinctly smaller pits (roughly half the size, as in Fig. 1). Shaw et

al. (1997) stated that males of cameronii have large dorsal median circular pits

on metasomal terga 5—7, but all of our specimens have the pits on terga 4—6 (Fig.

1B) or only terga 5 and 6 (Fig. 1A). We found few regional differences: specimens

from southern Mexico tend to have a weaker median carina on the propodeum,

and those from Yucatan have a slightly taller clypeus and are pale yellow. Our

material matches Cameron’s (1887) original description (including placement of

pits) with one exception. Cameron (1887) noted far fewer antennal segments in

his original description, but the number he gives (47) must have represented an

antenna that was broken.

Shaw et al. (1997) included 17 species in the pulchripes group, and although

they stated that the group is restricted to the New World, they undoubtedly meant

only those species with dorsal abdominal pits. Five of the species included in the

pulchripes group by Shaw et al. (1997) are known only from the Palaearctic. Six

of the 12 previously known New World species have males with setose pits (sim-

ilar to those of cameronii) on terga 4—6, 4—7 or 5—7. While the presence of these

pits is a potential synapomorphy uniting these species within the pulchripes spe-

cies group, the distribution of this character state is not congruent with other

characters that could be used to subdivide this species group, such as the arrange-

ment of teeth on the claws.

Aleiodes cameronii differs from all other members of the pulchripes species

68 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

group by the placement of fore wing cross-vein lcu-a, which arises near 1 M (an

unusual feature in Aleiodes). The curvature of hind wing RS is also distinctive.

The combination of tergal pits, closely spaced pattern of the enlarged teeth on

the tarsal claws, and uniformly colored body and flagellum further separates ca-

meronii from all but A. rossi Marsh and Shaw. In addition to the placement of

fore wing lcu-a, rossi differs in lacking hind wing m-cu and by having a shorter

hind wing r-m and more rugose propodeum. Both rossi and A. earinos Shaw have

large dorsal pits equivalent to those of cameronii. Although the venation in ear-

inos is more similar to that of cameronii than is rossi’s, the tarsal claws of earinos

are incompletely pectinate.

Dorsal abdominal pits have a restricted distribution within Aleiodes, occurring

in 14 described species. They are presently known only for males from the pul-

chripes and dispar species groups, but not all species within these groups have

males with abdominal pits (van Achterberg 1985, van Achterberg & Penteado-

Dias 1995, Shaw et al. 1997). These two species groups are readily separated

from each other, with major differences in the size of the pronotum, curvature of

the propodeum, size of eye and ocelli, relative length of the hind trochantellus,

and development of the precoxal sulcus (sternaulus) (van Achterberg & Penteado-

Dias 1995, Shaw et al. 1997). The pits themselves also differ slightly, being

confined to terga 2 and 3 in the dispar group and terga 4—7 in the pulchripes

group. Though work on the relationships among the species of Aleiodes is still

on-going (e.g., Fortier 1997), it seems unlikely that these two groups are sister

taxa. Specific features associated with the occipital carina, head shape, and body

sculpture suggest that members of the pulchripes group are more closely related

to species within other groups than to the species within the dispar group.

Though species with abdominal pits from the pulchripes group are apparently

confined to the New World, those from the dispar group are more widespread

(previously recorded from the Palaearctic, Neotropical, and Oriental Regions).

With the exception of A. excavatus (Telenga), however, males of the dispar group

with abdominal pits are rare in collections (van Achterberg & Penteado-Dias,

1995), and the disjunct distribution pattern of this group is likely an artefact.

Specimens in the TAMU collection indicate a more nearly cosmopolitan distri-

bution for these species, with representatives from Namibia and three localities

in Australia (two in Queensland and one in South Australia): the first records for

these two continents. The African and Australian individuals are typical members

of the dispar group, as defined by van Achterberg and Penteado-Dias (1995). As

there are only four specimens representing three species, and no accompanying

females, they are not described here.

ACKNOWLEDGMENT

We are particularly grateful to J. Fortier for checking our material, to S. Shaw

and P. Marsh for determining or verifying the identity of hundreds of specimens

from the TAMU collection, to C. van Achterberg for providing additional spec-

imens for comparison and reprints of his work related to the dispar group, to J.

Schaffner (Texas A&M University) for providing the Selander and Vaurie refer-

ence, and P Marsh and S. Shaw for comments about their work on Aleiodes. S.

Lewis provided information on some of the Cameron types and D. Azuma pro-

vided similar information on the Cresson types, for which we are most apprecia-

2000 DELFIN & WHARTON: REVIEW OF ALEIODES 69

tive. The work has been supported in part by the Texas Agricultural Experiment

Station, and in part by the National Science Foundation under grant No. 9712543.

LITERATURE CITED

Achterberg, C. van. 1982. Notes on some type-species described by Fabricius of the subfamilies

Braconinae, Rogadinae, Microgastrinae and Agathidinae (Hymenoptera, Braconidae). Ent. Ber.,

42: 133-139.

Achterberg, C. van. 1985. IV. The Aleiodes dispar-group of the Palaearctic region (Hymenoptera:

Braconidae: Rogadinae). Zool. Med. Leiden, 59: 178-187.

Achterberg, C. van. 1991. Revision of the genera of the Afrotropical and W. Palaearctic Rogadinae

Foerster (Hymenoptera: Braconidae). Zool. Verh. Leiden, 273: 1-102.

Achterberg, C. van. 1993. Illustrated key to the subfamilies of the Braconidae (Hymenoptera: Ichneu-

monoidea). Zool. Verh. Leiden, 283: 1-189.

Achterberg, C. van. & A. M. Penteado-Dias. 1995. Six new species of the Aleiodes dispar group

(Hymenoptera: Braconidae: Rogadinae). Zool. Med. Leiden, 69: 1-18.

Ashmead, W. H. (1888) 1889. Descriptions of new Braconidae in the collections of the U.S. National

Museum. Proc. U.S. Natl. Mus., 11: 611-671.

Cameron, P. 1887. Fam. Braconidae. Biol. Centr.-Am., 1: 312—419.

Cameron, P. 1905. Descriptions of new species of neotropical Hymenoptera. Trans. Am. Ent. Soc.,

31: 373-389.

Cresson, E. T. 1869. List of the North American species of the genus Aleiodes Wesmael. Trans. Am.

Ent. Soc., 2: 377-382.

Cresson, E. T. 1872. Hymenoptera Texana. Trans. Am. Ent. Soc., 4: 153-292.

Cresson, E. T. 1887. Synopsis of the families and genera of the Hymenoptera of America, North of

Mexico together with a catalogue of the described species, and bibliography. Trans. Amer. Ent.

Soc., (Suppl.): 1-350.

Dalla Torre, C. G. 1898. Catalogus Hymenopterorum. 4. Braconidae. G. Englemann, Leipzig.

Enderlein, G. (1918) 1920. Zur kenntnis auBereuropaeischer Braconiden. Arch. Naturges., 84A: 51—

224.

Fortier, J. C. 1997. Cladistics of the Aleiodes lineage of the subfamily Rogadinae (Hymenoptera:

Braconidae). Ph. D. Thesis, University of Wyoming Laramie, U.S.A.

Fox, W. J. (1894) 1895. Report on some Mexican Hymenoptera, principally from lower California.

Proc. Cal. Acad. Sci., 4: 1-25.

Gahan, A. B. 1917. Descriptions of some new parasitic Hymenoptera. Proc. U.S. Nat. Mus., 53: 195—

217.

Labougle, R. J. M. 1980. Analisis sobre la sistematica de la familia Braconidae (Ins. Hym.) y su

situacion actual en Mexico. Bachelor’s degree Thesis, UNAM—Facultad de Ciencias. Mexico.

Marsh, P. 1979. Family Braconidae. pp. 144-195. In Krombein, K. V., P. D. Hurd, D. R. Smith, and

B. D. Burks (eds.). Catalog of Hymenoptera in America North of Mexico. 1.

Marsh, P. M. & S. R. Shaw. 1998. Revision of the North American Aleiodes Wesmael. Part 3: the

seriatus (Herrich-Schaeffer) species-group (Hymenoptera: Braconidae: Rogadinae). Proc. Ent.

Soc. Wash., 100: 395-408.

Muesebeck, C. E W. 1925. A revision of the parasitic wasps of the genus Microbracon occurring in

America north of Mexico. Proc. U.S. Nat. Mus., 67: 1-85.

Muesebeck, C. EK W. 1960. New reared Neotropical species of Rogas Nees (Hymenoptera: Braconidae).

Ent. News, 71: 257-261.

Muesebeck, C. EF W. & L. Walkley. 1951. Braconidae. pp. 90-184. Jn Muesebeck, C. E W., K. V.

Krombein, and H. K. Townes. Hymenoptera of America north of Mexico. Synoptic Catalogue.

Ag. Monogr. 2.

Nees von Esenbeck, C. G. 1834. Hymenopterorum Ichneumonibus affinium monographiae, genera

Europaea et species illustrantes. Cotta, Stuttgart & Tubingen, 1: 1-320 and 2: 1-448.

Ruthe, J. EK 1855. Beitraege zur Kenntnis der Braconiden. (Exothecus, Ascogaster). Stett. Ent. Ztg.,

16: 291-294.

Say, T. 1832. Descriptions of new species of North American Hymenoptera, and observations on some

already described. Boston J. Nat. Hist., 1: 209-305.

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Selander, R. B. & P. Vaurie. 1962. A gazetteer to accompany the “‘Insecta” volumes of the ‘‘Biologia

Centrali-Americana.’’ Am. Mus. Novitates, 2099: 1—70.

Sharkey, M. J. & R. A. Wharton. 1997. Morphology and Terminology. pp. 19-37. In Wharton, R. A.,

Marsh, P. M. and Sharkey, M. J. (eds.). Manual of the New World Genera of the Family

Braconidae Hymenoptera. International Society of Hymenopterists. Special publication 1.

Shaw, S.R. 1993. Systematic status of Eucystomastax Brues and characterization of the Neotropical

species (Hymenoptera: Braconidae: Rogadinae). J. Hym. Res., 2: 1-11.

Shaw, S. R. 1997. Rogadinae s.s. pp. 403-412. In Wharton, R. A., PR. M. Marsh, and M. J. Sharkey

(eds.). Manual of the New World Genera of the Family Braconidae Hymenoptera. International

Society of Hymenopterists. Special publication 1.

Shaw, S. R., P M. Marsh & J.C. Fortier. 1997. Revision of North American Aleiodes Wesmael (Part

1): the pulchripes Wesmael species-group in the New World (Hymenoptera: Braconidae, Ro-

gadinae). J. Hym. Res., 6: 10-35.

Shaw, S. R., PR M. Marsh & J. C. Fortier. 1998a. Revision of North American Aleiodes Wesmael. Part

2: the apicalis (Brullé) species-group in the New World (Hymenoptera: Braconidae, Rogadinae).

J. Hym. Res., 7: 62-73.

Shaw, S. R., PR M. Marsh & J. C. Fortier. 1998b. Revision of North American Aleiodes Wesmael (Part

4): the albitibia Herrich-Schaeffer and praetor Reinhard species-groups (Hymenoptera: Bra-

conidae, Rogadinae) in the New World. Proc. Ent. Soc. Wash., 100: 553-565.

Shenefelt, R. D. 1975. Braconidae 8: Exothecinae, Rogadinae. Jn J. van der Vecht and R. D. Shenefelt

(eds.). Hymenopterorum Catalogus (nov. ed.), 12: 1115-1262.

Shenefelt, R. D. 1979. Some unusual Braconidae (Hymenoptera). Proc. Ent. Soc. Wash., 81: 125-134.

Szépligeti, G. 1904. Hymenoptera, Braconidae. Genera Insectorum, 22: 1-253.

Viereck, H. L. 1912. Descriptions of one new family, eight new genera, and thirty-three new species

of ichneumon-flies. Proc. U.S. Nat. Mus., 43: 575-593.

Wesmael, C. 1838. Monographie des Braconides de Belgique, 4. Nouv. Mem. Acad. Sci. R. Bruxelles,

11: 1-166.

Wolcott, G. N. 1948. The insects of Puerto Rico—Hymenoptera. J. Agric. Univ. Puerto Rico, 32: 751-

763.

Received 9 Jan 1999; Accepted 21 Apr 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 71-73, (2000)

Scientific Note

A RARE FIND: THE CAPTURE OF A PRIMARY QUEEN

OF THE WESTERN SUBTERRANEAN TERMITE

Field discoveries of true, macropterous-derived, physogastric, primary queens

of subterranean termites rarely occur (Snyder, T. E. 1935. Our enemy the termite.

Comstock Publishing Co., Ithaca, New York; Potter, M. EK 1997. Termites. Chapter

6. pp. 233-333. In Mallis, A. 1997. Handbook of pest control. 8th.ed.GIE Pub-

lishers Inc. Ohio; Thorne, B. L. 1998. Part 1. pp. 1-30. Biology of subterranean

termites of the genus Reticulitermes. NPCA research report on subterranean ter-

mites. NPCA, Dunn Loring, Virginia).

This is not at all surprising in that subterranean termites, particularly those of

the genus Reticulitermes, are cryptobiotic in nature. The critical life processes of

most subterranean termite colonies take place in the soil and/or in logs, stumps,

poles, posts, tree roots, etc. which are in the ground. Also, Reticulitermes nests

are not clearly defined, and the reproductive forms are known to migrate from

soil to wood and vice versa as well as within wood in response to changes in

temperature and moisture (Snyder 1935). All of these factors combine to make

the capture of true primary queens in the field rare events.

The first record of queens being found in a Nearctic subterranean termite colony

was reported in 1893 (Joutel, L. H. 1893. J. N. Y. Ent. Soc., 1: 89-90). The fact

that 23 queens were reported from one colony of Reticulitermes flavipes (Kollar)

confirms that these were secondary reproductives and not primary queens. The

first record of a true queen of Reticulitermes having been taken occurred in 1901.

The specimen in question was actually captured in June 1898. A notation in this

article stated that: “‘This is the first true Termite queen which has been found in

North America’? (Anonymous 1901. Proc. Ent. Soc. Wash., 4: 347). The capture

of a true queen of R. flavipes was recorded in 1902 (Schaeffer, C. 1902. J. N. Y.

Ent. Soc., 10: 251). A second record of the taking of a primary queen of R.

flavipes occurred in 1912 (Schaeffer, C. 1912. Bull. Brooklyn Ent. Soc., 8: 30).

Another report of the discovery of a true queen of R. flavipes was documented

in 1912 (Snyder, T. E. 1912. Proc. Ent. Soc. Wash., 14: 107-108). Apparently,

unaware of Schaeffer’s (1902) find, the preceding article stated that: “‘It is believed

that is the first fertilized true queen ever found of this species.”

This paper reports the capture of a primary queen of the western subterranean

termite, Reticulitermes hesperus Banks in Hemet, Riverside County, California

on 4 April 1995. Information is presented on the circumstances surrounding her

capture and the condition of this queen.

A few days before the discovery of this queen, the Terminix Riverside, Cali-

fornia office received a phone call from one of its customers reporting termites

swarming within one of their buildings. A Terminix representative was dispatched

to conduct an inspection and investigate the situation. Upon arrival at the property,

he was ushered into a concrete slab building which housed a shuffle board court/

auditorium and was told that termites had swarmed by the bleachers a few days

ago. Underneath one set of bleachers, a piece of partially delaminated, water-

72 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

Figure 1. A primary queen of the western subterranean termite, Reticulitermes hesperus Banks.

damaged plywood measuring approximately 70 cm X 70 cm X 1.4 cm covering

a recessed area in the ground was discovered. The sides of this sunken area in

the soil were fortified with form boards. On removing the plywood cover and

turning it over, extensive subterranean termite shelter tubes were found on the

form boards and on the bottom of the plywood cover. When the shelter tubes on

the underside of the cover were opened, a primary queen of R. hesperus was

found within the workings on the surface of the plywood (Fig. 1). Alates and

soldiers were also retrieved from this colony. They were keyed to species using

keys found in Weesner, E M. 1965. The termites of the United States—a hand-

book. National Pest Control Association, Dunn Loring, VA. It is important to note

that this imaginal queen was found in shelter tubes on wood essentially above

soil level. This queen may have been induced to move up to this area because of

the swarming which took place about two days before she was discovered. It is

known that swarming in Reticulitermes colonies is a frantic event which generates

enormous excitement, great activity, and much frenzy among members of a colony

(Snyder 1935).

2000 SCIENTIFIC NOTE 73

Observations on this queen in situ confirm what has been previously reported

in the literature in that these first form physogastric queens are very ambulatory

and are quite capable of traveling about on their own within a colony (Snyder, T.

E. 1920. Proc. Ent. Soc. Wash., 22: 109-150; Snyder 1935).

This primary queen of R. hesperus measured 12 mm long and 4 mm wide at

the greatest dorsal width of its abdomen. Reticulitermes queens have been reported

to reach the largest dimension of 14.5 mm in length and 4 mm wide (Snyder

1920; Snyder, T. E. 1934. Chapter 16. pp. 187-195. In Kofoid, C. A. (ed.) 1934.

Termites and termite control. Univ. Calif. Press, Berkeley). This first form queen

may not have achieved its maximum size in that it was an immaculate specimen.

Old queens of Reticulitermes have been described as having lost portions of their

antennae, legs, and margins of their thoraxes probably due to activities such as

constant social grooming (Snyder 1935).

The fact royal imaginal queens are rarely found in field colonies of Reticuli-

termes should not be interpreted to mean that these colonies are rarely headed by

true primary queens. The rarity of macropterous queens is an anthromorphic phe-

nomenon which is directly related to the inability to find these primary repro-

ductives in a cryptobiotic social insect.

Acknowled gment.—I would like to thank Stoy Hedges, Lonnie Anderson, and

Tony Borski for reviewing the manuscript and offering suggestions for its im-

provement.

Hanif Gulmahamad, 3547 Centurion Way, Ontario, California 91761.

Received 1 Mar 1999; Accepted 27 Oct 1999.

PAN-PACIFIC ENTOMOLOGIST

76(1): 74-76, (2000)

Scientific Note

NEW DISTRIBUTION RECORDS FOR THE ELDERBERRY

LONGHORN BEETLE DESMOCERUS CALIFORNICUS

HORN (COLEOPTERA: CERAMBYCIDAE)

Desmocerus californicus Horn is a large (12.3 to 27.2 mm length), black and red

cerambycid beetle which occurs only in California. The beetle’s obligate host and food

plant is elderberry (Sambucus spp., Caprifoliaceae) bushes which in California are

widely distributed and range from near sea-level to 3048 m in elevation. The beetle

larvae bore the bushes’ living stem pith and adults consume the leaves and flowers.

Adults emerge from living stems during spring or summer, leaving diagnostic oval-

shaped holes in the bark (Linsley, E. G. & J. A. Chemsak. 1972. Univ. Calif. Publ.

Entomol. 69; U.S. Fish and Wildlife Service (USFWS). 1984. Portland, Oregon;

USFWS. 1991. Sacramento, California).

Two subspecies of D. californicus are described: D. c. californicus Horn (California

Elderberry Longhorn Beetle [CELB]) and D. c. dimorphus Fisher (Valley Elderberry

Longhorn Beetle [VELB]). Prior to 1972, the VELB was considered a different and

valid species. The subspecies are differentiated by male characters of elytra color,

body length, and antennal hair color. Females of the two subspecies are alike (Linsley

& Chemsak 1972).

In 1980, the VELB was listed as a threatened subspecies (i.e., likely to become

endangered and possibly extinct in the future) by the USFWS (1980. Federal Register

45: 52803-52807). Within the VELB’s range, surveys for beetles and elderberry bush-

es are required for development projects. Also, the destruction of any elderberry bush

mandates extensive mitigation to avoid U.S. Endangered Species Act (1973) violations

and possible law enforcement and judicial actions.

When listed as threatened, the VELB was known from the Sacramento Valley (i.e.,

Sacramento and Davis) and the upper San Joaquin Valley (i.e., one site on the Merced

River). The nonthreatened CELB was known from numerous sites in coastal Califomia

from Mendocino County south to Los Angeles and Riverside counties (Linsley & Chem-

sak 1972). During the 1980s and 1990s, the collection of specimens and records of emer-

gence hole sightings improved our knowledge of the beetle’s distribution (USFWS 1984,

1991); however, that data and new information is unpublished and not readily available.

To clarify the range of D. californicus, 46 new locality records (29 from adult beetles

and 17 from emergence holes) obtained from field, literature, and museum surveys

are presented (Fig. 1). These data show that the species is widely distributed in Cal-

ifornia, and is significant in extending the known range of D. californicus northward

into Trinity County; into the southern San Joaquin Valley, the Sierra Nevada foothills,

and Mojave Desert mountain ranges; and southward to San Diego. These are the first

published records of D. californicus in Fresno, Kern, Madera, Mariposa, San Diego,

San Joaquin, Sutter, Tehama, Trinity, and Tulare counties. Additional locality records

are presented for Merced, Riverside, San Bernardino, and Santa Clara counties.

Subspecific names are not assigned to the new locales because the subspecies tax-

onomy is problematic. The differentiating characters for the subspecies intergrade and

overlap (USFWS 1984, 1991; Authors, personal observations). The new record males

from the Merced River (only one of two) and one from the Mokelumne River resemble

Linsley & Chemsak’s (1972) description of VELB. The other new record males (in

the San Joaquin Valley, Coast Range, Sierra Nevada foothills, and remainder of state)

resemble the description of CELB, the nonthreatened subspecies. Regardless, the

USFWS currently considers the Central Valley and surrounding foothills (below 914

2000 SCIENTIFIC NOTE 75

PIT RIVER

TRINITY RIVER

EEL ae

RUSSIAN RIVER

SACRAMENTO RIVER

FEATHER RIVER

AMERICAN RIVER

COSUMNES RIVER

MOKELUMNE RIVER

CALAVERAS RIVER

STANISLAUS RIVER

TUOLUMNE RIVER

MERCED RIVER

SAN JOAQUIN RIVER

KINGS RIVER

SALINAS RIVER KAWEAH RIVER

TULE RIVER

KERN RIVER

SANTA YNEZ RIVER

SANTA CLARA RIVER

RECORDS BASED UPON: SAN DIEGO RIVER

mi — EMERGENCE HOLE(S)

@ — ADULT SPECIMEN(S)

0 50 100

SCALE IN MILES

Figure 1. Updated distribution of the elderberry longhorn beetle Desmocerus californicus Horn

(Coleoptera: Cerambycidae). A symbol may represent several nearby locales. Locales of U.S. Fish

and Wildlife Service (1991) are mapped but not presented as new records in the text. The clear area

of California denotes its Central Valley.

m elevation) from Redding south through Kern County as the threatened subspecies’

range (USFWS. 1996. Sacramento, California).

New distributional records based upon adult beetles include: USA. CALIFORNIA. FRESNO Co.:

Kings Riv near Hwy 180, 119 m (390 ft), 13 Apr 1989, J. A. Halstead & J. A. Oldham, 1 male, 2

females (CAS—Calif. Acad. of Sciences, San Francisco); W Fork of Byrd Slough near Hwy 180, 117

m (385 ft), 12 Apr 1989, J. A. Halstead & J. A. Oldham, 1 female (CAS); Kings Riv near Annadale

Ave, 107 m (350 ft), 18 Apr 1989, J. A. Oldham, J. R. Single & J. A. Halstead, 2 females examined

and released; Herndon, 7 May 1970, L. H. Walker, 1 male (Calif. State Univ., Fresno); Table Mtn

about 3.1 mi N of Marshall Station, 366 m (1200 ft), 28 Apr 1995, D. York (personal communication).

KERN Co.: No locale, H. K. Morrison, 1 male, 1 female (NMNH—Natl. Mus., Nat. Hist., Wash.,

D.C.). MADERA Co.: Coarsegold, 671 m (2200 ft), 4 Jun 1989, R. M. Sadeghi & J. A. Halstead, 1

male (CAS); Hwy 41 about 8 mi S of Coarsegold, 375 m (1230 ft), 9 Jun 1991, J. A. Oldham & J.

A. Halstead, 1 female (CAS); San Joaquin Riv at Hwy 41, 84 m (275 ft), 1986, D. Mitchell (personal

communication), 1 female examined and released. MARIPOSA Co.: Mariposa, Jun 1974, 1 male (Acad.

Nat. Sciences of Phil., Pennsylvania). MERCED Co.: Merced Riv at Rd J7, 37 m (120 ft), 4 & 13

Apr 1990, J. A. Halstead & J. A. Oldham, 2 males (CAS); Los Banos Crk, Los Banos Val, 6 mi SE

of San Luis Reservoir, 122 m (400 ft), 11 Apr, 5 May 1987, D. Giuliani, 1 male, 1 female (CDFA—

Calif. Dept. of Food & Agric., Sacramento). RIVERSIDE Co.: 9 mi E of Temecula, 366 m (1200 ft),

76 THE PAN-PACIFIC ENTOMOLOGIST Vol. 76(1)

27 May 1969, R.R. Snelling, 2 males (LACM—Los Angeles Co. Mus., California); Menifee Val, 31

May 1976, S. I. & S. L. Frommer (Univ. of Calif., Riverside); SW of Anza, 25 Jun 1949, Simonds,

2 males (CDFA). SAN BERNARDINO Co.: Cedar Cyn, 1554 m (5100 ft), 1 Apr 1981, T. Griswold,

1 female (USU—Utah State Univ., Logan); Mitchells Cavern State Park, 24 Mar 1980, T. Griswold,

1 male (USU). SAN DIEGO Co.: Pala, May 1962, M. E. P, 1 male (LACM); Mission Val, 15 May

1931, 1 male, 1 female (SDNHM-San Diego Nat. Hist. Mus., California). SAN JOAQUIN Co.: Mo-

kelumne Riv, Clements Glen View Cem, 41 m (135 ft), 15 May 1991, C. Barr, 1 male, 1 female

(LACM). SANTA CLARA Co.: Coyote Crk, 10 Aug 1974, M. Robey, 1 male (TCACO—Tulare County

Agric. Comm. Office, Visalia, California); Uvas Mdws, 25 May 1969, W. F & B. C. Tyson, 1 female

(W. Tyson, personal collection, Oakhurst, California); Mount Madonna, 7 Apr 1972, H. J. Denk, 1

male (SDNHM); Silver Creek, 16 Apr 1940, G. S. Mansfield (CAS). TRINITY Co.: No locale, 14

May 1925, E. R. Leech, 1 male (NMNH). TULARE Co.: Kaweah, 13 Jun 1937, FE T. Scott, 1 male

(TCACO); Kaweah Power Sta #3, Sequoia Natl Park, Ash Mtn Park Hdarts, 671 m (2200 ft), 22 May

1982, 19 Jun 1983, 9 May 1986, 1 male, 2 females (TCACO); Campbell-Moreland Ditch (trib of Tule

Riv) at A.T. & S.E Railroad between Worth Ave & Hwy 190, 149 m (490 ft), 21 Apr 1991, C. Barr,

1 female (LACM); Lane Slough (trib of Kaweah Riv), Rd 196 & Hwy J27, 123 m (405 ft), 30 Apr

1991, C Barr, 1 male (LACM).

New distributional records based upon emergence hole sightings include: U.S.A. CALIFORNIA.

FRESNO Co.: Reedley, Kings Riv Col, 104 m (341 ft), 1993, W..M. Rhodehamel (personal communi-

cation); Channel Rd about 3 mi SE of Sanger, 104 m (340 ft), 1990, J. A. Oldham; 1 mi W of Piedra,

Hughs Crk, 168 m (550 ft), 1990, J. A. Oldham & J. A. Halstead; Hwy 180 at Alta Main Canal, 122

m (400 ft), 1990, J. A. Oldham & J. A. Halstead; San Joaquin Riv at Riverside Golf Course, 91 m (300

ft), 1989, M. Boland, J. A. Oldham & J. A. Halstead; Hwy 168 about 5 mi NE of Prather, Sierra Natl

For, T10S, R23E, Sec 15, 732 m (2400 ft), 1994; D. York (personal communication); Trimmer Sprgs

Rd near Secata Crk, N of Pine Flat Reservoir, Sierra Natl For, 640 m (2100 ft), 1995, J. A. & B.S.

Halstead. MADERA Co.: Oakhurst, 884 m (2900 ft), 1991, J. A. Oldham & J. A. Halstead; Fresno Riv

about 3 mi W of Oakhurst, 732 m (2400 ft), 1991, J. A. Oldham & J. A. Halstead; Quartz Mtn Rd

about 5 mi SE of Coarsegold, 683 m (2240 ft), 1993, J. A. & P. S. Halstead; Chowchilla Riv at Rd 19,

82 m (270 ft), 1993, J. A. Halstead & J. C. Stebbins. MERCED Co.: Livingston, Merced Riv at Hwy

99 (Calif. Nat. Diversity Data Base. 1992. Calif. Dept. of Fish & Game, Sacramento (CNDDB)). SUTTER

Co.: Feather Riv near Live Oak (CNDDB 1992). TEHAMA Co.: Jellys Ferry Rd about 15 mi N of Red

Bluff, near Sacramento Riv, 122 m (400 ft), 1989, J. A. Oldham & J. A. Halstead. TULARE Co.: Tule

Riv Indian Reserv, 305 m (1000 ft), 1993, J. A. Halstead; Sequoia Natl Park, Potwisha Cmpgd, 549 m

(1800 ft), 1993, R. D. Haines; 1994, PR S. & J. A. Halstead; 1.5 mi N of Lemoncove, Kaweah Riv at

Hwy 216, 152 m (500 ft), 1997, Alice Karl, P S. & J. A. Halstead.

The types of both subspecies and 428 specimens of D. californicus were examined from the following

locations (excluding the new record locales): U.S.A. CALIFORNIA. ALAMEDA Co.: Leona Heights

(Park); Mission San Jose; Oakland. BUTTE Co.: Ordbend. COLUSA Co.: Grimes; Colusa. CONTRA

COSTA Co.: Martinez. GLENN Co.: Butte City. LAKE Co.: Adams Sprg (Adams); Gravelly Val, Hull-

ville. LOS ANGELES Co.: Chatsworth; Eagle Rock; Los Angeles; Pasadena; Santa Monica; La Brea.

MARIN Co.: Lagunitas; Marin City; Novata; no locale. MENDOCINO Co.: Yorkville; no locale. NAPA

Co.: Chiles Val. ORANGE Co.: Atwood; Fullerton; Huntington Beach; Laguna Beach; Newport Bay;

Newport Beach; Newport, Back Bay; Long Beach. RIVERSIDE Co.: Riverside; Sonorian Region; no

locale. SACRAMENTO Co.: Sacramento; Cosumnes Riv near Sloughhouse. SAN BENITO Co.: SW of

Idria (New Idria). SAN BERNARDINO Co.: Lytle Crk; San Bernardino, Upland. SAN LUIS OBISPO

Co.: no locale. SANTA BARBARA Co.: San Roque Crk, Coastal Slope; Montecito; Santa Barbara. SANTA

CLARA Co.: San Jose; Stanford Univ. SOLANO Co.: Cold Crk near Monticello Dam; G. L. Stebbins

Cold Cyn Reserv. SANTA CRUZ Co.: Ben Lomond. SONOMA Co.: Eldridge; Healdsburg; Petaluma;

Sonoma. TEHAMA Co.: Red Bluff. VENTURA Co.: Ojai. YOLO Co.: Davis; Knights Landing.

Acknowled gment.—We thank the Kings River Conservation District for conducting

the studies; other biologists, researchers, and agencies for providing assistance and

information; museums and private collectors for loaning specimens and colleagues and

reviewers for their advice and helpful editorial comments.

Jeffrey A. Halstead and Jonathan A. Oldham!, Environmental Division, Kings River

Conservation District, 4886 E. Jensen Avenue, Fresno, California 93725. !Current

address: Coast Branch Project, California Department of Water Resources, 3220 S.

Higuera, #304, San Luis Obispo, California 93401.

Received 20 Jan 1998; Accepted 24 Aug 1999.

PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

See volume 74: 248-255, October 1997, for detailed general format information and the issues thereafter for examples; see below for

discussion of this journal’s specific formats for taxonomic manuscripts and locality data for specimens. Manuscripts must be in English,

but foreign language summaries are permitted. Manuscripts not meeting the format guidelines may be retumed. Please maintain a copy

of the article on a word-processor because revisions are usually necessary before acceptance, pending review and copy-editing.

Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 X 11 in,

nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS OF

ANY FIGURES TO THE 8.5 < 11 INPAGE. Number pages as: title page (page 1), abstract and key words page (page 2), text pages

(pages 3+), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last. List

the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right comer. The title

must include the taxon’s designation, where appropriate, as: (Order: Family). The ABSTRACT must not exceed 250 words; use five

to seven words or concise phrases as KEY WORDS. Number FOOTNOTES sequentially and list on a separate page.

Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases

underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet

1988, Dorly et al. 1989), (Burton in press) and (R. F. Tray, personal communication). For multiple papers by the same author use:

(Weber 1932, 1936, 1941; Sebb 1950, 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith

1987: table 3).

Taxonomy. — Systematics manuscripts have special requirements outlined in volume 69(2): 194-198; if you do not have access to that

volume, request a copy of the taxonomy/data format from the editor before submitting manuscripts for which these formats are

applicable. These requirements include SEPARATE PARAGRAPHS FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED

(INCLUDING A SPECIFIC FORMAT), and a specific order for paragraphs in descriptions. List the unabbreviated taxonomic author

of each species after its first mention.

Data Formats. — All specimen data must be cited in the journal’s locality data format. See volume 69(2), pages 196-198 for these

format requirements; if you do not have access to that volume, request a copy of the taxonomy/data format from the editor before

submitting manuscripts for which these formats are applicable.

Literature Cited. — Format examples are:

Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.

Blackman, R. L., P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometrics provide

some answers? pp. 233-238. Jn Holman, J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics and

taxonomy of aphids and Thysanoptera. Proc. intemational symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985. SPB

Academic Publishing, The Hague, The Netherlands.

Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.

Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.

Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 X 181 mm while maintaining label

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Mount all illustrations. Label illustrations on the back noting: (1) figure number, (2) direction of top, (3) author’s name, (4) title of

the manuscript, and (5) journal. FFGURE CAPTIONS must be on a separate, numbered page; do not attach captions to the figures.

Tables. — Keep tables to a minimum and do not reduce them. Table must be DOUBLE-SPACED THROUGHOUT and continued on

additional sheets of paper as necessary. Designate footnotes within tables by alphabetic letter.

Scientific Notes. — Notes use an abbreviated format and lack: an abstract, key words, footnotes, section headings and a Literature Cited

section. Minimal references are listed in the text in the format: (Bohart, R. M. 1989. Pan-Pacific. Entomol., 65: 156—-161.). A short

acknowledgment is permitted as a minor headed paragraph. Authors and affiliations are listed in the last, left indented paragraph of

the note with the affiliation underscored.

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THE PAN-PACIFIC ENTOMOLOGIST

Volume 76 January 2000 Number 1

Contents

RUNGROJWANICH, K. & G. H. WALTER—tThe Australian fruit fly parasitoid Diachasmi-

morpha kraussii (Fullaway): life history, ovipositional patterns, distribution and hosts

(Hymenoptera: Braconidae: Opiinae)

RUNGROJWANICH, K. & G. H. WALTER—tThe Australian fruit fly parasitoid Diachasmi-

mor pha kraussii (Fullaway): mating behavior, modes of sexual communication and

crossing tests with D. longicaudata (Ashmead) (Hymenoptera: Braconidae: Opiinae) ___.

CHAO, R.-F. & C.-S. CHEN—Formosozoros newi, a new genus and species of Zoraptera

(Insecta) from Taiwan

COVILLE, R. E., C. GRISWOLD & P. L. COVILLE—Observations on the nesting biology and

behavior of Trypoxylon (Trypargilum) vagulum (Hymenoptera: Sphecidae) in Costa Rica _

GOMEZ, J. & O. GARCIA—A new species of Encarsia (Hymenoptera: Aphelinidae), a para-

sitoid of whitefly Aleurodicus sp. (Homoptera: Aleyrodidae) in Mexico

HALSTEAD, J. A—A new species of Hockeria Walker from Mexico (Hymenoptera: Chalcididae) _.

LI, Q. & J. HE—Entomognathus from China with description of a new species (Hymenoptera:

Sphecidae)

DELFIN G., H. & R. A. WHARTON—Historical review of the genera Aleiodes and Rogas in

Mexico, with a redescription of Aleiodes cameronii (Hymenoptera: Braconidae)

SCIENTIFIC NOTES

GULMAHAMAD, H.—A rare find: the capture of a primary queen of the western subterranean

termite

HALSTEAD, J. A. & J. A. OL. DHAM—New distribution records for the elderberry longhom

beetle Desmocerus californicus Horn (Coleoptera: Cerambycidae)

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 76 April 2000 Number 2

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)