Nesting Behavior of Prionyx "thomae" (Fabricius) (Hymenoptera: Sphecidae)

Author(s): E. E. Grissell
Source: Journal of the Kansas Entomological Society, Vol. 54, No. 1 (Jan., 1981), pp. 16-21
Published by: Allen Press on behalf of Kansas (Central States) Entomological Society
Stable URL: http://www.jstor.org/stable/25084126 .
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 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
54(1), 1981, pp. 16-21

Nesting Behavior of Prionyx "thomae" (Fabricius)

(Hymenoptera: Sphecidae)
E. E. Grissell

Systematic Entomology Laboratory, IIBIII, Agricultural Research,

Science & Education Administration, USDA1

abstract: In Florida, Prionyx "thomae" (Fabricius) nested in

firm sand along the edge of a salt marsh. Six females were observed
to restructure burrows of the fiddler crab, Uca rapax (Smith). Be
havior of one female, observed during a complete nest cycle, dif
fered from previous records for Nearctic species by construction
of a nest prior to prey hunting and by carriage of prey in flight.
Possible explanations for such different behavior include adapta
tion to a littoral habitat and the possibility that this represents a
cryptic species which has been confused with P. thomae.

While studying nesting biology of the sphecid Pluto littoralis (Malloch),

several females of Prionyx "thornae" (Fabricius) (see discussion section
below for citation of specific name in quotations) were observed and their
behavior recorded. My observations differ significantly from other records
for the species in 3 respects: 1) females used preexisting holes of the fiddler
crab, Uca rapax (Smith), which were restructured to accommodate prey;
2) nests were constructed prior to prey hunting, a habit generally not as
sociated with Nearctic Prionyx ; and 3) prey was carried in flight for at least
some distance before being placed near the nest and then dragged into it.
Known biology o? Prionyx has been reviewed by Evans (1958) and Menke
(1976). All species provision with Acrididae (see Table 9 in Menke, 1976).
Nearctic species collect prey and place it in a protected place before digging
a nest. Evans (1958:182) noted a possible exception to this behavior with
the comment that a wasp he observed may have reversed the "usual prey
?? nest sequence." After digging a unicellular nest, Nearctic Prionyx drag
the prey over the ground by its antennae, place it in the nest, head-in,
dorsum-up, and lay an egg with one end at the base of the hindcoxa (Evans,
1958). Several Neotropical species dig the nest prior to hunting but are
otherwise similar in habit to Nearctic species (Menke, 1976). Only one
species, P. spinolae, is known to carry its prey in flight and to make a
multicellular nest each provisioned with 5 to 10 grasshoppers.

1
Mailing address: % U.S. National Museum of Natural History, Washington, D.C. 20560.
Received for publication 15 April 1979.

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VOLUME 54, NUMBER 1 17

Observations
In a previous article I described and illustrated the study site which was
located at Cedar Key, Levy County, Florida (Grissell, 1979). Briefly, it was
a narrow strip of firm sand at the edge of a Spartina -dominant salt marsh.
Although above high tide level, the subsurface sand was constantly moist.
Burrows of Uca rapax were abundant on this sand strip (Fig. 1). Obser
vations were made on 1, 7, and 16 September 1975. One female was ob
served during a complete nesting cycle, and eight additional females were
seen investigating crab holes, of which 5 actively restructured the burrows.
At no time did a female begin excavation on bare ground.
The following account is of a single female which initiated and completed
nest construction in about 45 minutes (2:15-2:58 p.m., EDST). Other fe
males will be discussed below. At 2:15 a female went headfirst into a vertical
crab hole (Fig. 2). She immediately turned around and began gouging sand
at a right angle to the shaft about 1 cm below the surface of the entrance.
The sand was scraped with the mandibles and fell down the shaft of the
crab hole. The female did nothing with this sand until later. After about 5
minutes of digging, the female abruptly turned to a point 180? from the
burrow she had been digging and began excavating at a point opposite the
first one. When the main shaft of the crab hole became backfilled to a point
just below the entrance of the burrow under construction, the female took
soil from the excavation and deposited it just outside the crab hole entrance.
At 2:30 (after 15 minutes of excavation), the female left the nest (head

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 18 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

first) and walked ca. 10-15 cm in one direction, pivoted around several
times, then walked back to the nest entrance. This process was repeated in
several different directions at which time the female flew up to a Baccharis
bush, paused, then flew down the sand road for at least 10 m before flying
out of sight. At 2:34 the female returned, in flight, carrying a grasshopper
beneath her body. She landed on the ground ca. 1 m from the nest and
placed the grasshopper in a small clump of grass. Then she flew directly to
the nest, entered, spent almost 60 seconds inside, and walked directly back
to the prey. The prey was straddled, dorsal side up, and the bases of both
antennae were taken in her mandibles. The forelegs were placed behind the
prey's head, and the mid and hindlegs were used to walk forward to the
nest. Here the female placed the grasshopper headfirst at the nest entrance,
went into the nest, turned around, and pulled the prey in by its antennae.
Within a minute the female returned to the main shaft and began tearing
down the inside perimeter of the burrow with her mandibles, kicking soil
behind her into the tunnel leading to the prey chamber. Occasionally she
would turn around, and pack the soil into the side tunnel with her head.
This was accompanied by much wing buzzing. As the side tunnel became
filled she began scraping sand from the backfill area, thus partially reexca
vating the original crab hole. By 2:43, or 7 minutes after depositing the prey
in the nest, the female completely emerged from the main burrow, and began
breaking down the rim of the original entrance hole and kicking sand from
around the entrance into the nest. Her head was used from time to time to
tamp the sand firmly into place. After the nest and crab hole were com
pletely filled and packed to surface level, the female began kicking sand into
a pile over the entrance. She spent several minutes covering the area, and
then flew off at 2:48. At 2:50 she returned, did some more kicking for 3
minutes, and then left again. A minute later she returned and kicked more
sand around the area for 4 minutes. Then she left and did not return. I
excavated the nest (Fig. 3) and found a weakly paralyzed, mature grass
hopper (Clinocephalus ele g ans), in a head-in, dorsum-up position with an
egg at the base of the left hindcoxa.
Portions of the behavior noted for this female were observed in other
females but not during a complete nest cycle. Five females were seen re
structuring old Uca burrows. While the actual nest construction provision
cycle seemed relatively short (ca. 40 minutes for the female just discussed),
there was apparently much tentative exploring before a nest was completely
dug. One female worked for an hour, digging and refilling 5 crab holes within
a 200 cm2 area. Several times she backfilled one burrow with the sand from
a nest in close proximity, only to turn around and reexcavate the first bur
row. After 60 minutes she abandoned the nest site without provisioning any
cells. Evans (1958:182) reported a female which started and abandoned 6
nests in a 27 minute period.

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VOLUME 54, NUMBER 1 19

Fig. 3. Nest diagram of Prionyx "thomae." Stippling indicates sand; diagonal lines indicate
filled burrow; A) abandoned crab burrow, B) partially excavated wasp nest (abandoned), C)
complete cell with grasshopper provision. (Approximately life size.)

Five prey were

collected, 3 from nests, and 2 from females in the act of
provisioning. They were all mature Clinocephalus elegans, measuring from
27-31 mm in length. The body position and egg placement of excavated
prey were the same as described above. Of the 3 buried prey, 2 were missing
one or other of the hindlegs. In all 3 instances the egg was at the base of
the left hindcoxa.

Discussion
Evans (1958) summarized the nesting habits of Prionyx thomae as re
ported from Texas, Kansas, and Missouri, and added further notes for the
former 2 states. He also gave a list of ethological characters shared in com
mon by all Prionyx, as well as characters of perhaps "greater importance,"
but for which exceptions were known. The species I observed differs from
all other known Prionyx by the use of preexisting burrows for nest con
struction. It differs from published biologies of other species, except the
Neotropical P. spinolae, by carriage of the prey in flight at least for part of
the return trip. And it differs from Nearctic species (except for one obser
vation by Evans, 1958:182) by capturing prey after the nest is finished.
Because the biology of females I observed varied so much from previous
reports for thomae, it might be conjectured that what I observed was a
cryptic species easily confused with thomae. According to Bohart and
Menke (1963:160) thomae ranges widely throughout western North America
but is "uncommonly" found in the southeastern United States. They also
pointed out that male specimens taken from coastal South Carolina differed
morphologically from western specimens. A reexamination of Eastern pop
ulations of "thomae" might reveal the presence of a second species.2

-
Regrettably, voucher specimens of female P. "thomae" used in this study have been either
lost or misplaced.
in transit The identification of this species was based upon several female
specimens examined by Bohart, Menke, and myself. Males were not collected. Attempts at
further observation and collection at the nest site were met by a refractory landowner.

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 20 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Since reported biological observations have been from inland sites, I may
have worked with a population of thomae which has adapted to a littoral
habitat. Firstly, the carriage of prey in flight might be explained by the off
shore capture of grasshoppers on salt marsh grass {Spartina alterniflora
Loiseleur-Deslongchamps). It would generally not be possible for a female
to walk back with prey, even at low tide, because of standing water. Sec
ondly, the use of crab holes as starting points for nests may have been a
factor of their abundance (Fig. 1) but did not appear to be due to hardness
of the soil {Pluto littoralis easily began nests in the same soil; Grissell,
1979). However, using restructured, preformed burrows does not seem in
herently more advantageous than initiating new ones, given the softness of
the sand and the shallowness of the nest. Finally, there would seem to be
no inherent adaptiveness to a littoral habitat (above high tide) which would
cause females to provision in any particular sequence either before or after
nest construction. Further comparison of western and eastern populations
of Prionyx thomae seems warranted.
Barber and Matthews (1979) provided another recent example of possible
habitat adaptation in their studies of the sphecid Trypargilum politum (Say)
which normally constructs mud nests which resemble organ pipes. At the
Okefenokee Swamp, Georgia, however, this species was found to nest in
preexisting cavities (bamboo trap nests) in 2 cypress habitats. Reasons for
this atypical behavior were summarized as possible occasional aberation
without real adaptive benefit, adaptation to a habitat with little mud, ad
aptation to interspecific competition due to other mud-wasps, and lack of
suitable natural substrates upon which to build a mud nest.
Observations such as reported by Barber and Matthews (1979), and for
Prionyx in this paper, demonstrate the potential relationship of geography
and habitat adaptation to the behavior of fossorial Hymenoptera. While
numerous studies have been made for individual species of nesting Hyme
noptera, relatively few comparative studies of populations have been made.
This would seem to provide abundant opportunity for evolutionary studies.

Acknowledgments
I wish to thank Drs. A.
S. Menke (Systematic Entomology Laboratory,
USDA, Washington, D.C.) and R. M. Bohart (Department of Entomology,
University of California, Davis) for identifying P. "thomae", Dr. R. E.
Woodruff (Florida Department of Agriculture and Consumer Services,
Gainesville) for identifying the grasshopper prey, and Mr. R. Zranz (Zoology
Department, University of Florida, Gainesville) for identifying the species
of crab. Drs. H. E. Evans (Department of Entomology, Colorado State
University, Fort Collins) and R. F. Denno (Department of Entomology,
University of Maryland, College Park) reviewed the manuscript, and I wish
to thank them for many helpful suggestions.

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VOLUME 54, NUMBER 1 21

Literature Cited

Barber, M. C, and R. W. Matthews. 1979. Utilization of trap nests by the pipe-organ mud
dauber, Trypargilum polit urn. Ann. Entomol. Soc. Amer. 72:260-262.
Bohart, R. M., and A. S. Menke. 1963. A reclassification of the Sphecinae with a revision of
the Nearctic species of the tribes of Sceliphronini and Sphecini. Univ. California Pub.
Ent. 30:91-181.
Evans, H. E. 1958.Studies of the nesting behavior of digger wasps of the tribe Sphecini. Part
I: Genus Prionyx Dahlbom. Ann. Ent. Soc. Amer. 51:177-186.

Grissell, E. E. 1979. Nesting biology of Pluto littoralis. J. Kans. Ent. Soc. 51:269-275.
Menke, A. S. 1976. Sphecinae. Pages 79-154 in R. M. Bohart and A. S. Menke, eds. Sphecid
wasps of the world, a generic revision. Univ. Calif. Press, Berkeley and Los Angeles.

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