Geographic Range
Hagenius brevistylus
, commonly known as the dragonhunter, has a wide distribution across 34 states in
the eastern United States and in the adjacent 5 provinces of southeastern Canada.
Dragonhunters are endemic, or native and unique, to North America east of the Great
Plains.
Habitat
Dragonhunters are found in the temperate regions of North America, where adults prefer the banks of forested streams and rivers that have moderate to fast flow. Dragonhunter adults can be found foraging in the woods near streams.
Larvae live in or along the edges of woodland streams, amid leaf litter and detritus,
which match their own dark coloration. Older dragonhunter larvae prefer sheltered
shorelines with silt-sand sediment bottoms. In the water, larvae are found at depths
of 0.5 to 10 m in streams that contain beds of muddy sand and organic debris.
- Habitat Regions
- temperate
- terrestrial
- freshwater
- Terrestrial Biomes
- forest
- Aquatic Biomes
- rivers and streams
- Other Habitat Features
- riparian
Physical Description
Dragonhunters are large dragonflies that possess long and powerful legs and wings. Adults range in size from 73 to 90 mm long, the span of their hindwings measures 47 to 58 mm, and they weigh an average of 1.2 g.
Adults have green eyes and a small, black head with a face striped in yellow. The body is characterized by black and yellow markings, with two thick, yellow stripes on the sides of the thorax. The dorsal surface of abdominal segments 9 and 10 are completely black, while the rest of the abdomen has yellow spots. The legs are entirely black, with short, heavy spines on the lower sections to assist in taking large prey. The posterior end of the abdomen is enlarged, forming a club. The club of a dragonhunter is relatively small compared to other clubtail dragonflies. The wings are coarse and slightly tinted, with blackish veins running across them.
Males and females are similar in color and markings, but the club on a male is bent
downward in a sideways J-shape when in flight or perched. The anal loop (a pattern
on the hindwing) is more broad in the female than in the male.
The larvae, or nymphs, of dragonhunters have stilt-like legs and possess a broadly
flattened, nearly circular abdomen with serrated edges. These characteristics provide
camouflage, as they create an overall resemblance to the dead leaves in which the
nymphs hide. The mouth of a dragonhunter larva consists of a pair of heavy ridges
to support a stout labium, which is an extendable lower lip that is used to catch
prey. At the end of the labium is a pair of palpal lobes lined with hooks, spines,
and teeth.
- Other Physical Features
- ectothermic
- heterothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes shaped differently
Development
Hagenius brevistylus
follows the general pattern of
dragonfly
development, undergoing incomplete metamorphosis, which does not include a pupal
stage. Eggs that are laid late in the summer may undergo diapause, or hibernation,
and wait until the spring to hatch. The larval stage of dragonhunters can be exceptionally
long, ranging from 4 to 7 years. Larvae appear to be freeze tolerant, which enables
them to survive during the winter. The rate at which the nymphs molt and grow depends
on ambient temperatures.
- Development - Life Cycle
- metamorphosis
- diapause
Reproduction
Little is known of the
Hagenius brevistylus
mating system. Like most
dragonflies
, dragonhunters probably are polygynandrous, such that males and females mate with
multiple partners. Little information is available about courtship behavior and mate
attraction in dragonhunters. The papillae at the base of the antennae may play a role
in courtship and reproduction, but in general,
dragonflies
depend on vision, and males exhibit territorial behavior to announce availability
and suitable egg-laying sites.
Reproduction in dragonflies usually involves two formations between a mating pair: the tandem position and the copulation wheel.
First is the tandem position. Males in the family Gomphidae pursue and quickly grasp females from behind without any apparent behavioral or visual display. The male grasps the female’s head or thorax and curves his abdomen under his body to grip the back of her head with his anal appendages, or claspers. The pair then flies off with the male in the lead, towing the female behind him. This process usually takes less than a second, and at this stage the couple is in the tandem position. The pair stays in this position for seconds to minutes and may fly around in tandem or find a suitable place to settle.
Before mating occurs, the
dragonflies
in the tandem pair adjust the positions of their bodies to form a copulation wheel.
The female curls her abdomen underneath her thorax while in tandem, moving the abdomen
forward to contact the genitalia of the male. The couple forms a heart-shaped wheel
as the male and female mate. Male dragonhunters use their genital hamuli (small forked
appendages) to support the erected penis during mating.
Dragonhunter males are notorious for inflicting severe damage on females while mating.
The spines of the male’s anal appendages grasp the female in a vise-like grip, gouging
the female’s eyes and usually piercing her head. Copulation is brief, and the couple
seldom flies while in tandem, because physical damage can occur. Tandem pairs probably
perch in treetops for the majority of the mating interval to prevent linkage strain
caused by flying in tandem.
- Mating System
- polygynandrous (promiscuous)
The flight and breeding period of Hagenius brevistylus is approximately 3 months in the summer, during which male and female dragonhunters mate as many times as possible. Like most dragonflies , sperm is injected into the female during copulation, but the eggs are not fertilized until they are laid by the female. Males compete for their sperm to be used in fertilization by removing any sperm that previously was deposited in the female before injecting their own.
Dragonhunter females oviposit (lay their eggs) while flying in a zigzag course, circling
over a comparatively small area of open water. They rise and descend repeatedly, dipping
the tip of their abdomen into the water with each descent to deposit a little cluster
of eggs. Information is scarce regarding the number of eggs laid by female dragonhunters,
but in most
dragonflies
, the number varies from a few hundred to a few thousand.
When the eggs of dragonhunters enter the water, they scatter and drift apart as they
fall to the bottom. The eggs are evenly covered by a jelly except at the anterior
pole, which is left uncovered so that hatching is unhindered. As the eggs fall through
the water, silt sticks to their surface, and they become hidden at once. Neither parent
protects the eggs after oviposition.
Little information is available about the gestation period of dragonhunters, but most dragonfly eggs hatch within 8 to 30 days, depending on the environmental conditions. Dragonhunter eggs typically hatch during the summer and develop and grow in the water; the hatched larvae shift closer toward the shoreline as they mature. Eggs that are laid late in the summer may undergo diapause, or hibernation, and hatch the following spring.
Parental care has not been observed in this species, and like most dragonflies , H. brevistylus larvae appear to be independent the moment they hatch. The duration of the larval stage in dragonhunters ranges from 4 to 7 years, and the larvae appear to be freeze tolerant, which enables them to survive during the winter. The rate at which the nymphs molt and grow largely depends on ambient temperatures.
Adults emerge after metamorphosis in late June and early July, often on the shoreline,
but occasionally as much as a meter high on trees in the forest. Dragonhunters exhibit
“mass emergence”, also known as “explosive emergence”, in which most of the annual
population appears at once within a short period of time. There is little information
on the time of sexual maturity for dragonhunters, but most
dragonflies
become sexually mature about 2 to 3 weeks after reaching adulthood.
- Key Reproductive Features
- semelparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- oviparous
- sperm-storing
- delayed fertilization
As in most
dragonflies
, yolk from the mother nourishes the hatching larvae. No further parental investment
or care is provided by male or female dragonhunters.
- Parental Investment
- no parental involvement
-
pre-fertilization
- provisioning
-
pre-hatching/birth
-
provisioning
- female
-
provisioning
Lifespan/Longevity
The lifespan of dragonhunters is about 4 to 7 years. The rate at which the nymphs
molt and grow largely depends on ambient temperatures. Adults live no longer than
3 months.
Behavior
Hagenius brevistylus is a relatively solitary species. It is known for its fearlessness, as adults have been observed to relentlessly pursue songbirds that are much larger than the dragonflies themselves.
Dragonhunters display territorial behavior and defend an area vigorously from other individuals of the same species. Little information is available about the size of the guarded territories.
Adults change their orientation with respect to the sun to control their body temperature.
For example, at high temperatures, dragonhunters perch with their long abdomens hanging
directly downward, parallel to the sun’s rays, thus minimizing their shadow and reducing
direct exposure to the sun. Dragonhunters are active when exposed to sunlight.
When hunting,
H. brevistylus
adults can reach speeds of up to about 40 km per hour. They often attack from above,
sometimes knocking prey into water before going in for the kill. Dragonhunters use
their long legs to grasp prey and their large mandibles to deliver a powerful and
often fatal bite. Adult dragonhunters have a habit of flying along a regular path
with definite perches while foraging, and they usually perch to consume large prey.
They most often choose to perch on a thorny branch to eat
Ebony Jewelwings
(a species of broad-winged damselfly) and take approximately 25 minutes to chew and
ingest them. When eating
monarch butterflies
, dragonhunters use their forelegs to rotate the monarch and press the wings of the
monarch together for easier handling. Because monarchs sequester unpalatable chemicals
in their bodies, dragonhunters avoid eating the more toxic portions of the prey. For
instance, dragonhunters almost never eat monarch wings, as the wings contain high
levels of toxins. Dragonhunters appear to prefer feeding on monarchs, especially when
the butterflies are abundant.
Dragonhunter larvae are slow-moving sprawlers and hiders, as opposed to burrowers;
they rest on substrate using their long, extended legs for support. Nymphs occasionally
appear near the surface of water to molt. A diverse assortment of larval instars usually
can be found together.
- Key Behaviors
- flies
- diurnal
- motile
- sedentary
- hibernation
- solitary
- territorial
Home Range
Information is scarce about the home range of dragonhunters. Most evidence suggests
that dragonhunter larvae stay within a relatively small area as they mature.
Communication and Perception
Information is scarce regarding the methods of communication and perception specific to Hagenius brevistylus . The papillae at the base of its antennae may be used to communicate with mates, involving touch and chemical signals. Adult dragonflies perceive the environment visually, and they can see UV light. Male dragonhunters are territorial, displaying their size and speed.
Larvae use mechanical, visual, and chemical cues to sense their prey.
- Perception Channels
- visual
- ultraviolet
- tactile
- vibrations
- chemical
Food Habits
Hagenius brevistylus
is an insectivore, with adults primarily eating butterflies and other insects, especially
other
dragonflies
. Dragonhunters use their long legs to grasp prey and their large mandibles to deliver
a powerful and often fatal bite. Dragonhunters are known to eat
Lake Darners
,
Ebony Jewelwings
,
Cyrano Darners
,
swallowtail butterflies
, and
monarch butterflies
. Dragonhunters seem to prefer feeding on monarchs, especially when the butterflies
are abundant. Dragonhunters appear able to overcome the toxic defenses of
monarchs
,
bees
and
wasps
.
One experiment concluded that an adult dragonhunter would have to consume 60% of its
body weight each day to meet its daily water requirements at 25°C, which would be
enough food to power its flight for 4.6 hours. However, due to constraints in the
experiment, the calculations may have overestimated the daily requirements.
Larvae are highly predatory and eat anything of suitable size, including other
odonate
larvae,
fish
, and
amphibians
.
- Primary Diet
- carnivore
- Animal Foods
- amphibians
- fish
- insects
Predation
Scarce information is available about the specific predators of dragonhunters. In general, crustaceans , true bugs , beetles , turtles , and waterfowl are predators of dragonfly larvae. Hagenius brevistylus larvae avoid predation mostly by blending in with leaves and detritus in their aquatic habitat.
When transitioning from an aquatic to a terrestrial habitat, and as adults,
dragonflies
are attacked by
ants
,
spiders
,
frogs and toads
,
shrews
,
raccoons
, and various
birds
.
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Dragonhunter larvae are aquatic carnivores and insectivores; dragonhunter adults are
aerial insectivores. As major predators of
monarch butterflies
,
Hagenius brevistylus
adults can influence monarch behavior. For instance, if dragonhunters are active
where monarchs are present, the monarchs avoid predation by feeding in the shade despite
their usual preference for sunlight.
Dragonhunters have been affected by the introduction of
zebra mussels
to the Great Lakes. Because dragonhunter larvae possess a wide abdomen and exhibit
sprawling behavior, they are more susceptible to
zebra mussel
colonization than the slimmer, burrowing larvae of other
dragonfly
species. The presence of mussels on
H. brevistylus
larvae has a negative effect on the survivorship of the larvae. Indeed, colonized
larvae show decreased mobility and increased susceptibility to predation. However,
zebra mussel
colonization does not appear to hamper the feeding rate of dragonhunter larvae.
Economic Importance for Humans: Positive
As insectivorous predators, dragonhunters may be effective in regulating insect populations (some of which are pests to humans), such as mayflies , stoneflies , caddisflies , and possibly mosquitoes . Dragonhunters also serve as indicators of clean water in stream ecosystems.
Some research includes
Hagenius brevistylus
as the study species because it is relatively easy to observe and because of its
unique predatory behavior.
- Positive Impacts
- research and education
- controls pest population
Economic Importance for Humans: Negative
There are no known negative impacts of Hagenius brevistylus on humans.
Conservation Status
Dragonhunters are common (though rarely abundant) throughout their geographic range,
and their populations are stable. They are present in many federal, state, local,
and private reserves and do not require conservation measures at this time.
Additional Links
Contributors
Allison Liao (author), University of Michigan-Ann Arbor, Elizabeth Wason (author, editor), Animal Diversity Web Staff.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- terrestrial
-
Living on the ground.
- freshwater
-
mainly lives in water that is not salty.
- forest
-
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- ectothermic
-
animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature
- heterothermic
-
having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
- metamorphosis
-
A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.
- diapause
-
a period of time when growth or development is suspended in insects and other invertebrates, it can usually only be ended the appropriate environmental stimulus.
- polygynandrous
-
the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.
- semelparous
-
offspring are all produced in a single group (litter, clutch, etc.), after which the parent usually dies. Semelparous organisms often only live through a single season/year (or other periodic change in conditions) but may live for many seasons. In both cases reproduction occurs as a single investment of energy in offspring, with no future chance for investment in reproduction.
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- internal fertilization
-
fertilization takes place within the female's body
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- sperm-storing
-
mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.
- delayed fertilization
-
a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.
- diurnal
-
- active during the day, 2. lasting for one day.
- motile
-
having the capacity to move from one place to another.
- sedentary
-
remains in the same area
- hibernation
-
the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.
- solitary
-
lives alone
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- chemical
-
uses smells or other chemicals to communicate
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- vibrations
-
movements of a hard surface that are produced by animals as signals to others
- chemical
-
uses smells or other chemicals to communicate
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
References
Ackerman, J. 2006. Dragonflies Strange Love. National Geographic Magazine , 209/4: 104. Accessed March 23, 2012 at http://ngm.nationalgeographic.com/2006/04/dragonfly-mating/ackerman-text .
Berger, C. 2004. Dragonflies . Mechanicsburg, PA: Stackpole Books. Accessed June 17, 2013 at http://www.kilibro.com/book/preview/78915_wg-dragonflies .
Bienek, R., S. Hickner. 2007. "The effects of colonization by Zebra mussels, Dreissena polymorpha , on fitness in two Anisopteran species: Hagenius brevistylus and Didymops transversa " (On-line pdf). Deep Blue at the University of Michigan. Accessed January 28, 2012 at http://deepblue.lib.umich.edu/bitstream/handle/2027.42/57456/Bienek_Hickner_2007.pdf?sequence=1 .
Brooks, S. 2003. Dragonflies . Washington, D.C.: Smithsonian Books.
Brunelle, P. 2010. Dragonflies and damselflies ( Odonata ) of the Atlantic Maritime Ecozone. Pp. 333-369 in Assessment of Species Diversity in the Atlantic Maritime Ecozone . Ottawa: NRC Research Press. Accessed March 22, 2012 at http://books.google.com/books?id=c2n7FPi3Lu0C&pg=PA338&lpg=PA338&dq=Hagenius+brevistylus+development&source=bl&ots=HispK5fDAc&sig=y9rRxjJvHlXR0YU8vYcpycL4SXs&hl=en&sa=X&ei=5-dtT_f0C8v6ggeN7NFr&ved=0CEcQ6AEwBg#v=onepage&q=Hagenius%20brevistylus%20development&f=false .
Burcher, C., L. Smock. 2002. Habitat distribution, dietary composition and life history characteristics of odonate nymphs in a Blackwater coastal plain stream. The American Midland Naturalist , 148/1: 75-89.
Catling, P. 2004. Why are Hagenius brevistylus nymphs so distinctive?. Ontario Odonata , 5: 27.
Corbet, P. 1962. A Biology of Dragonflies . Chicago: Quadrangle Books.
Corbet, P. 2004. Dragonflies : Behaviour and Ecology of Odonata . Colchester: Harley Books.
Cresswell, S. 2009. " Hagenius brevistylus : Dragonhunter" (On-line). American Insects. Accessed January 28, 2012 at http://www.americaninsects.net/d/hagenius-brevistylus.html .
Fincke, O., D. Santiago, S. Hickner, R. Bienek. 2009. Susceptibility of larval dragonflies to zebra mussel colonization and its effect on larval movement and survivorship. Hydrobiologia , 624/1: 71-79. Accessed June 17, 2013 at http://faculty-staff.ou.edu/F/Ola.M.Fincke-1/Finckeetal09.pdf .
Harris, W., D. Forman, R. Battell, M. Battell, A. Nelson, P. Brain. 2011. Odonata colour: more than meets the eye?. International Journal of Odonatology , 14/3: 281-289.
Johnson, C. 1972. Tandem Linkage, Sperm Translocation, and Copulation in the Dragonfly , Hagenius brevistylus ( Odonata : Gomphidae ). American Midland Naturalist , 88/1: 131-149.
McCormac, J. 2005. Fierce Dragonflies Display Killer Skills When They Hunt. The Columbus Dispatch , Tuesday, August 16: 7D.
Mead, K. 2003. Dragonflies of the North Woods . Duluth: Kollath-Stensaas.
Mead, K. 2008. " Dragonflies in Winter" (On-line). Minnesota Odonata Survey Project. Accessed March 21, 2012 at http://www.mndragonfly.org/journal.html .
Medland, V. 2008. " Dragonhunter " (On-line). Eco-notes: The Cofrin Center for Biodiversity. Accessed March 21, 2012 at http://www.uwgb.edu/biodiversity/econotes/2007/dragonhunter.htm .
Needham, J., M. Westfall, M. May. 2000. Dragonflies of North America . Gainesville, FL: Scientific Publishers.
Tracy, B., C. Tracy, D. Dobkin. 1979. Desiccation in the black dragon, Hagenius brevistylus Selys. Experientia , 35/6: 751-752.
Tracy, C., B. Tracy, D. Dobkin. 1979. The Role of Posturing in Behavioral Thermoregulation by Black Dragons ( Hagenius brevistylus Selys; Odonata ). Physiological Zoology , 52/4: 565-571.
Westfall, M., K. Tennessen. 1996. Odonata . Pp. 164-211 in An Introduction to the Aquatic Insects of North America . Dubuque, IA: Kendall/Hunt. Accessed March 22, 2012 at http://books.google.com/books?id=UUtEoRejk0AC&pg=PA164&lpg=PA164&dq=Hagenius+brevistylus,+mating+system&source=bl&ots=f_VFxYEY0v&sig=s37YhYcggeVUfi4Lp_KvNgKLuZI&hl=en&sa=X&ei=dzBuT4fPJYWMgweY5dxr&ved=0CEYQ6AEwCA#v=onepage&q=Hagenius%20brevistylus%2C%20mating%20system&f=false .
White, D., O. Sexton. 1989. The monarch butterfly ( Lepidoptera : Danaidae ) as prey for the dragonfly Hagenius brevistylus ( Odonata : Gomphidae ). Entomological News , 100/3: 129-132. Accessed January 28, 2012 at http://www.biodiversitylibrary.org/item/21676#page/142/mode/1up .
2010. " Hagenius brevistylus : Dragonhunter" (On-line). Encyclopedia of Life. Accessed January 28, 2012 at http://eol.org/pages/1034811/overview .