T
conid subfamilies and genera (Marsh et al. 1987,
Shaw and Huddleston 1991, Sharkey 1993, Shaw
1995, Wharton et al. 1997).
Most braconids are primary larval–larval parasitoids of other insects, ovipositing into and emerging from the host larva (Shaw and Huddleston
1991). A lesser number are larval–pupal parasitoids, egg–larval parasitoids, or parasitoids of adult
insects. Phytophagy is known to occur, although it
is rare (Shaw 1995). Many braconids kill major
insect pests and therefore are used extensively in
biological control (Shaw 1995); they are among
the most frequently and successfully used beneficial organisms in classical biological control programs (Greathead 1986).
For such an economically significant group of organisms, it is surprising that the family Braconidae
has no common name, other
than “braconid wasps.” More
often than not, they are called
“parasitic wasps” along
with numerous unrelated
families (Quicke 1997).
Because they are parasitoids that nearly always kill their hosts,
Shaw (1994) proposed the common
name
“death
wasps.” Sharkey
(1994) has called
them “silk wasps” or
“cocoon wasps” for
their ability, as larvae,
to produce silk from
An observation of the characteristics of the Meteorus cocoon,
labial silk glands. None
of these suggested comand the variety of suspended-cocoon forms in the genus
mon names has been
Meteorus, with focus on this highly interesting but
widely adopted.
underemphasized aspect of braconid biology.
Silk production, cocoon
morphology, and behavior
associated with cocoon formation and adult emergence from
the cocoon are aspects of braconid
biology that have received relatively
little attention. Quicke (1997) provides
the most comprehensive literature review
of parasitic wasp silk production and related
topics; Shaw and Huddleston (1991) and Shaw
(1995) provide general reviews of braconid cocoon forms. The objective of this paper is threeand Carlson 1979, Achterberg 1984). However, fold: to review the variety of suspended-cocoon
these estimates may be low because many tropical forms in the genus Meteorus, present speculative
ideas about the adaptiveness of these cocoon forms
species remain undescribed (Shaw 1997).
Although braconid species richness encom- and, therefore, focus attention on this highly inpasses a great diversity of adult morphological teresting but underemphasized aspect of braconid
forms, all braconid wasps are readily distinguish- biology.
Meteorus is a large genus with a cosmopolitan
able from ichneumonids by the fusion of the second and third metasomal tergites. The fusion of distribution (more than 200 described species). All
these tergites is usually rigid, but occasionally it is species of Meteorus are koinobionts: the host larva
flexible; and a sculptured or smooth suture often is not paralyzed during oviposition and the host
indicates the junction of the two (Shaw 1995). Sev- continues to grow while parasitized (Quicke 1997).
eral sources provide keys and diagnoses of bra- They are also endoparasitic, living and feeding inhe genus Meteorus Haliday is a member of
the family Braconidae, a large and important group of parasitoid wasps. Within the
hyperdiverse order Hymenoptera, the Braconidae
and its sister group the Ichneumonidae are the two
largest families (Wahl and Sharkey 1993, Gauld
and Shaw 1995, Grissell 1999). The Braconidae
comprises more than 14,890 described species
worldwide (Wharton 1997), but it has been estimated to contain 40,000 to 50,000 species (Marsh
From Meteors
to Death Stars:
Variations on a Silk Thread
(Hymenoptera:Braconidae:
Meteorinae)
Nina M. Zitani and Scott R. Shaw
228
AMERICAN ENTOMOLOGIST • Winter 2002
side their hosts. Most species of Meteorus attack
young caterpillars, but some attack beetle larvae.
»
Building a Suspended Cocoon
The life cycle of a typical Meteorus species begins with the adult female inserting an egg into the
body of an exposed feeding host caterpillar. The
egg hatches and the larva feeds on the blood of the
host (Madel 1963), while the host remains active
and responsive to stimuli (Askari et al. 1977). Just
before pupation of the host, the nearly or fully
developed Meteorus larva chews a hole in the abdomen of the host and exits the body (Simmonds
1947, Madel 1963, Askari et al. 1977). The host
caterpillar dies within a few hours of emergence of
the parasitoid (Fuester et al. 1993), although it
may remain alive for up to 36 h after emergence
(Askari et al. 1977).
The Meteorus larva then moves to a nearby leaf
or twig and forms a silk pad on the substrate. Next
the larva suspends itself from the silk pad by a silk
thread, usually ≈3.0 cm long (Fig. 1), but occasionally up to 18.0 cm (Fig. 2), and forms its cocoon.
Based on the resemblance of this suspended cocoon to a meteor plummeting from the sky,
Alexander H. Haliday named the genus in 1835.
Within the family Braconidae, this suspended cocoon-forming behavior is unique.
The wasp pupates facing downward within the
cocoon (i.e., with its head at the end of the cocoon
opposite the suspending thread). It is possible to
observe this directly in living specimens because
freshly spun Meteorus cocoons often are translucent. The end of the cocoon opposite the suspending thread is therefore referred to as the anterior
end, and the end attached to the thread is referred
to as the posterior end. When the wasp has completed its development and is ready to emerge, it
cuts a neat, circular cap from the anterior end of
the cocoon. The cap is symmetrical about the long
axis, and the emergence hole and cap edge are
smooth (Fig. 3).
Another characteristic of the Meteorus cocoon
is that the silk at the anterior apex is very thick
compared with the silk of the rest of the cocoon.
This thickened area of silk is most apparent if observed from the interior of the cap. This silk also
appears to be of a different texture than the surrounding silk (Fig. 3). As observed by Askari et al.
(1977), the Meteorus larva, “. . . [pays] particular
attention to the anterior apex where a very thick
button of white silk is spun.” Because of this thickened area of silk, the cocoon appears asymmetrical
when whole: the anterior end is more elongated
and tapered, whereas the posterior end is more
rounded (Askari et al. 1977).
The function of this thickened area of silk at
the anterior apex of the cocoon is unknown; however, a few observations are worth noting. After
emergence, adult Meteorus rest on the cocoon before taking flight (NMZ, personal observation).
The adult Meteorus escapes from a cocoon that is
suspended in midair. Wasps emerging from the
anterior end, as they do in nature, have a readyAMERICAN ENTOMOLOGIST • Volume 48 Number 4
Another
characteristic of the
Meteorus cocoon is
that the silk at the
anterior apex
is very thick
compared with
the silk of
the rest of
the cocoon.
Figs. 1–3. (1) Typical Meteorus suspended cocoon,
»3.0 cm in length, that has been removed from the
substrate to which it was attached. Note the silk pad
marking the origin of the thread (arrow). Scale = 1.0 cm.
(2) Suspended cocoon of Meteorus with unusually long
suspending thread, »18.0 cm long. Scale = 1.0 cm. (3)
Close-up of a typical Meteorus cocoon from which the
adult has emerged. Note the circular emergence hole,
the smooth edge of the emergence hole and cap, and
the interior texture of the cap (arrow). Photographs by
N. Zitani.
229
Figs. 4–5.(4) M. papiliovorus cocoon removed from the substrate, showing short
(»2.0 mm) suspending thread. Photograph by S. R. Shaw. (5) Intact M. papiliovorus
cocoons showing pronounced nipplelike anterior ends (arrow). Photograph by G.
Thorn, Pitilla Biological Station, Area de Conservacion Guanacaste, Costa Rica.
The strategy of
suspending oneself
is not novel in
nature. Within the
animal kingdom,
numerous
unrelated species
suspend themselves
or form suspended
structures
made platform (the remainder of the cocoon,
which is still attached to the substrate by the
thread) on which to emerge and remain, until they
are ready to fly. If wasps were to emerge from the
posterior end, they would cut the cocoon from
the suspending thread and, while still inside the
remainder of the cocoon, fall to the ground. These
wasps would be more likely to die, so natural selection would favor exiting from the anterior end
of the cocoon.
Once the cocoon is complete and the Meteorus
larva is ready to pupate, how does it determine
which end of the cocoon is the anterior end? During the process of cocoon spinning, the Meteorus
larva reverses its position numerous times, spinning layers of silk at one end and then turning on
end to spin layers at the other end (Askari et al.
1977). As cocoon construction continues, the larva
progressively isolates itself from its surroundings.
Perhaps the parasitoid determines its orientation
within the cocoon simply by gravity, or perhaps
the thickened area of silk at the anterior end serves
as a guide mark. The construction of guide marks
for proper orientation within cocoons was documented in an ichneumonid by Salt (1977). The
Meteorus larva may be able to detect the thickened
area of silk at the anterior apex by touch and then
orient itself properly before pupating.
Why Form a Suspended Cocoon?
Several years ago, Meteorus papiliovorus Zitani,
a species with a short (≈2.0 mm) cocoon-suspending thread was described (Zitani et al. 1997) (Figs.
230
4–5). The asymmetry of the cocoon is pronounced
in this species; the anterior end forms a nipple shape
(Fig. 5). If the question prior to this 1997 publication had been, “Why does Meteorus form a suspended cocoon?” it now became, “Why bother
forming a suspending thread that is only 2.0 mm
long?”
The first question is an interesting one. It has
been suggested that the suspended cocoon makes
the pupating wasp inaccessible to some potential
enemies (Shaw and Huddleston 1991, Quicke
1997). This seems plausible given that an ant, or
other predator, crawling over the surface of a leaf
in search of food is less likely to discover a hanging
cocoon than one that is attached directly to the
leaf. For a suspended organism, the area of contact
between the substrate, or searching area of a crawling predator, is reduced to a single point.
The strategy of suspending oneself is not novel
in nature. Within the animal kingdom, numerous
unrelated species suspend themselves or form suspended structures, presenting excellent examples
of convergent evolution (e.g., the cocoon of some
campoplegine ichneumonids, some social wasp
nests, butterfly chrysalids, the nests of some birds
such as orioles and oropendolas). Insects use a
variety of materials to suspend themselves: plant
fibers surrounded by an oral secretion, silk, larval
exuvium, cremaster (a series of hooks on a butterfly pupa imbedded in a silk pad on the substrate)
(Figs. 6–9).
Many insects suspend themselves while molting, so that emergence from the old cuticle is aided
by the force of gravity (Chapman 1982). This may
be an explanation for the crysalid (a suspended
butterfly pupa), but the evolution of suspended
structures has undoubtedly been influenced by a
variety of forces. It has been argued that predation
by ants is a major selective force in the evolution of
nest form in tropical social wasps. Some tropical
wasps (e.g., Polistes spp., Mischocyttarus spp.)
build a nest that is suspended from the substrate
by a narrow petiole. They also smear the petiole
with a glandular secretion that has ant-repellant
properties. It appears that the narrow nest petiole
and the ant-repellent secretion are an effective system of defense against ant predation (Jeanne 1970,
1975).
Even if predation has been a major force in the
evolution of the suspended cocoon in Meteorus
spp., why do we see so much interspecific variation
in the length of the cocoon-suspending thread? In
Costa Rica, a small Citrus tree was discovered that
nearly had been defoliated by leaf-cutter ants, except for one leaf with a cluster of M. papiliovorus
cocoons (Fig. 10). This suggests that M.
papiliovorus has evolved an additional defense
mechanism to deter crawling predators, especially
ants (Zitani et al.1997). The Citrus-feeding
papilionid host caterpillars secrete defensive chemicals from their osmeteria (fleshy eversible processes
located dorsally just behind the head). Perhaps M.
papiliovorus takes advantage of the host
caterpillar’s own defense by causing it to lay down
AMERICAN ENTOMOLOGIST • Winter 2002
Figs. 6–9. (6) Vespid
wasp on nest suspended
by petiole. Photograph by
E. S. Ross, Vila Amazona,
Brazil, June 1964. (7)
Campoplegine
ichneumonid cocoon
suspended by silk strand.
Photograph by E. S.
Ross, Lake Berryessa,
CA. (8) Endomychid
beetle pupae suspended
by cast larval exuviae.
Photograph by E. S.
Ross, Tingo Maria, Peru.
(9) Chrysalis of zebra
butterfly, Heliconius
charitonius (L.).
Photograph by S. R.
Shaw, lab colony.
repellant chemicals from its osmeterium before it is
killed. Osmeterial secretions have been shown to
be an effective defense against ants (Eisner and
Meinwald 1965, Honda 1983).
In addition to attacking a Citrus-feeding
papilionid, M. papiliovorus attacks an Aristolochiafeeding papilionid (Zitani et al. 1997). Related
Aristolochia-feeding species sequester toxic chemicals from their host plants (Euw et al. 1968, Nishida
and Fukami 1989). Barbosa et al. (1982) found
the presence of nicotine in the cocoons of a
microgastrine braconid that attacks the tobacco
hornworm, Manduca sexta (L.) (Sphingidae), suggesting that the parasitoid takes in nicotine from
its host and incorporates it into the cocoons. Perhaps M. papiliovorus is incorporating the toxic
chemicals from its host into its silk cocoons, providing an additional defense against ants.
If the chemical defense is adequate, then why
suspend the cocoon at all? The presence of a sus-
Figs. 10–11. (10) M. papiliovorus cocoons on nearly defoliated Citrus tree. Photograph by S. R. Shaw, La Selva Biological Station, Costa
Rica. (11) M. congregatus cocoons adjacent to sphingid host cadaver. Photograph by D. H. Janzen, Area de Conservacion Guanacaste,
Costa Rica.
AMERICAN ENTOMOLOGIST • Volume 48 Number 4
231
pending thread indicates the relatedness of M.
papiliovorus to other Meteorus spp. that spin longthreaded cocoons. Maybe the short threads are
simply a result of natural selection: If the chemical
defense deters crawling predators, then a longer
suspending thread no longer provides an advantage. Individuals with a chemical defense and a short
suspending thread may be just as protected from
crawling predators as individuals that make a
longer thread.
The Gregarious Meteorus
Gregarious parasitism is another interesting aspect of Meteorus biology. It is well known that
some Meteorus spp. are gregarious (Huddleston
1980; Maetô 1989, 1990; Shaw and Huddleston
1991; Shaw 1995). Gregarious parasitism results
when two or more parasitoid larvae from the same
mother develop in a single host. Gregariousness
may be the result of multiple eggs laid in the host,
Figs. 12–16. (12) Gerardo Vega looking at suspended communal cocoon
mass of M. townsendi (arrows). Notice exceptionally long suspending thread
(»3.0 m). (13) Manduca sp. host with suspending threads of M. townsendi
cocoon mass (arrow). (14) Close-up of M. townsendi communal cocoon
mass. (15) Close-up of M. townsendi larvae forming the communal cocoon
mass. (16) Second close-up of M. townsendi larvae forming the communal
cocoon mass. Photographs by G. Gentry, La Selva Biological Station, Costa
Rica.
232
or it may result from the repeated division of a
single egg (polyembryony) (Gauld and Bolton
1988); however, Meteorus is not known to be polyembryonic. Within the Braconidae, in general,
brood size corresponds to the number of eggs deposited, with the exception of the Macrocentrinae
(Shaw 1995).
In terms of numbers of individuals per host,
gregariousness reaches its peak in Meteorus
congregatus Muesebeck. Up to 250 individuals have
been reported from a single Manduca sexta caterpillar, feeding on Solanum torvum Sw. (Solanaceae),
in Costa Rica. (Zitani et al. 1998) (Fig. 11). M.
congregatus cocoons are attached directly to the
substrate; they have no suspending thread. Phylogenetic analyses suggest M. congregatus is derived
from species that make suspended cocoons (NMZ,
unpublished data).
Meteorus townsendi Muesebeck is a Neotropical gregarious species that probably surpasses all
other Meteorus spp. in volume of silk production
per larva. It also is interesting because it produces a
suspended communal cocoon mass, suggesting
some amount of cooperation among the larvae.
When the type specimens of M. townsendi were
examined, we found that the suspending thread of
the cocoon mass was the longest (45.0 cm) of any
Meteorus sp. Unfortunately, it had been cut, so the
original length was impossible to determine. This
was not the first time that such a problem had been
encountered. Frequently Meteorus cocoons are not
preserved or are not preserved intact (either the
suspending thread has been cut or the cocoon cap
is missing, or both).
In summer 1999, NMZ met Grant Gentry at
the La Selva Biological Station in Costa Rica. Gentry had come upon M. townsendi in the field and
had taken a superb series of photographs. The
first of these photographs shows the exceptionally
long suspending thread of the M. townsendi communal cocoon mass and documents the longest
thread in the genus (Fig. 12); it was ≈3 m long (G.
Gentry, personal communication)! Several closeup shots show the Manduca sp. host (Fig. 13), and
the M. townsendi larvae in the process of constructing their cocoons, which will eventually become
part of the larger cocoon mass (Figs. 13–16). To
the best of our knowledge, this is the first photographic record of M. townsendi larvae in the field.
Most of us are familiar with caterpillars and
spiders that drop from foliage on silk threads, and
we consider spider silk to be especially strong because of its ability to hold a violently wriggling
prey item captive. The exceptionally long suspending thread of M. townsendi suggests that Meteorus
silk also is quite strong and durable. The suspending threads of various species in museum collections are extremely durable and remain flexible for
decades. However, Meteorus silk never has been
tested experimentally for strength or any other
quality.
The architecture of a suspended communal cocoon mass reaches its zenith in the highly organized, radially symmetrical cocoon mass of the AfAMERICAN ENTOMOLOGIST • Winter 2002
Figs. 17–20. (17)
Suspended spherical
cocoon mass of M.
komensis removed from
substrate. Scale = 0.5
cm. (18) Close-up of
cocoon mass of M.
komensis showing intact
cocoons. (19) Close-up
of cocoon mass of M.
komensis showing adult
emergence holes and
hyperparasitized cocoon
(arrow). (20) Second
close-up of cocoon mass
of M. komensis showing
hyperparasitized cocoon
(arrow). Photographs by
N. Zitani.
rican Meteorus komensis Wilkinson. The larvae
(≈100 from a single host) form a very regular,
spherical cocoon mass with the nipple-shaped anterior ends of all the cocoons facing outward (Figs.
17–18). We affectionately call this communal cocoon mass the “death star”, in reference to Shaw’s
(1994) suggested common name for the
Braconidae, and its starlike appearance. After M.
komensis adults emerge, the silk sphere appears to
have holes cut in its surface (Fig. 19). We know of
only two specimens, from the National Museum
of Natural History, Smithsonian Institution (Figs
17–20).
A hyperparasitoid is a parasitoid that develops
on another parasitoid (Gauld and Bolton 1988),
and Meteorus solitary cocoons often are recorded
as being hyperparasitized (e.g., Lyle 1914,
Simmonds 1947). Meteorus gregarious cocoon
masses also are attacked by hyperparasitoids. A
M. komensis cocoon that contained a
hyperparasitoid is apparent because of the emergence hole cut in the side of the cocoon (Figs. 19–
20). Because Meteorus always emerges by cutting
an anterior cap, it is clear that what emerged from
this cocoon was not Meteorus.
At least eight cocoons of the M. komensis coAMERICAN ENTOMOLOGIST • Volume 48 Number 4
coon mass contained hyperparasitoids (possibly
more, because not all of the adults emerged), but
most of the cocoons were not attacked (Fig. 19).
Given that hyperparasitoids fail to attack all the
potential hosts in a cocoon mass, it has been proposed that the gregarious cocoon mass offers more
protection against hyperparasitoids than solitary
cocoons (Gauld 1991, Hanson and Gauld 1995).
The specific architecture of the M. komensis “death
star” may provide even more protection from
hyperparasitoids: Only the outer tip, maybe 10%
of the cocoon surface area, is exposed to searching
hyperparasitoids. It may be a result of the same
basic strategy as the suspended cocoon (i.e., that of
reducing the surface area exposed to predators)
but, in this case, flying hyperparasitoids.
Meteorus communal cocoon masses, particularly those of the M. komensis “death star,” are
strikingly beautiful and raise interesting questions
about the behavior of the larvae. If not for the
protection from hyperparasitism, why do the larvae of M. komensis build such an organized structure, how do they build it, and what keeps them
from wandering off and building their own solitary cocoons? Perhaps we see in Meteorus a series
of strategies, solitary and communal, for the de-
From meteors
to death stars,
Meteorus spp.
construct some
of the most
remarkable
cocoon forms
in the insect
world.
233
fense of the pupating larvae. From meteors to death
stars, Meteorus spp. construct some of the most
remarkable cocoon forms in the insect world.
Acknowledgments
We gratefully acknowledge Grant Gentry (Mesa
State College, Grand Junction, CO), Daniel H.
Janzen (University of Pennsylvania, Philadelphia),
Edward S. Ross (California Academy of Sciences,
San Francisco), and Greg Thorn (University of
Western Ontario, London) for the use of their
outstanding photographs; the curators of the
National Museum of Natural History,
Smithsonian Institution, Washington, DC; Canadian National Collection, Ottawa; and the Natural History Museum, London, UK, for the loan
of specimens; Ian Craig (University of Western
Ontario) and Gary Fetter (University of Wyoming)
for assistance in creating digital images from color
slides; Greg Thorn, and two anonymous reviewers for providing helpful comments on the manuscript. We express our appreciation to J. E.
McPherson for his support, patience, and critical
review of the manuscript.
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AMERICAN ENTOMOLOGIST • Volume 48 Number 4
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Zitani, N. M., S. R. Shaw, and D. H. Janzen. 1997.
Description and biology of a new species of
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Meteorinae) from Costa Rica, parasitizing larvae
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Systematics of Costa Rican Meteorus (Hymenoptera: Braconidae: Meteorinae) species lacking a dorsope. J. Hymenoptera Res. 7: 182–208.
Nina M. Zitani is a Ph.D. candidate in the Department of Renewable Resources, University of Wyoming (Laramie, WY 82071-3354). She currently
resides in London, Ontario (email: [email protected]).
Scott R. Shaw is professor of entomology and curator of the U.W. Insect Museum, University of
Wyoming (Laramie, WY 82071-3354; email:
[email protected]).
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