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Insect Order Ephemeroptera (Mayflies)

Pictures Below
Mayflies may be the most important insects for trout anglers to understand. They are an ancient order of insects, famous outside the fly-fishing world for their fragile beauty and short adult lifespan, often a single day to mate and die. The mayfly's poignant drama attracts poets and anglers alike, but anglers make the most of it.

Mayflies live more than 99% of their lives as nymphs on the river or lake bottom, filling many crucial roles in freshwater ecosystems as they feed and grow. They eventually emerge from the water as winged sub-adults called "subimagos" by scientists and "duns" by anglers. Duns evolved to be good at escaping the water, with a hydrophobic surface and hardy build, but they are clumsy fliers. Within a day or two they molt one last time into "imagos" or "spinners," the mature adults, a transformation captured in this photo series of a dun molting into a spinner. They have longer legs and tails, and sleeker, more lightweight bodies, giving them the airborne speed, agility, and long grasp they need for their midair mating rituals. They are usually darker than the duns and have shinier, more transparent wings. They die within minutes or hours after mating.

Hatching Behavior


The importance of mayflies comes largely from their emergence and mating behavior. While many organisms assure the survival of their species using individualistic tools like stealth, speed, venom, or parental care, mayflies are famous instead for their "strength in numbers" approach. They coordinate their emergence and mating times (both time of year and time of day) so that they leave their safe habitats and emerge together in large numbers in a very short period of time. This can trigger feeding frenzies in every nearby insect-eater, from trout to birds to dragonflies, but there are simply so many mayflies at once that many luck out and survive to reproduce. These trout feeding frenzies are the stuff of legend among fly anglers, and they also pose one of our greatest challenges, because trout feeding feverishly on a thick hatch are often unwilling to strike any fly that doesn't properly imitate the mayfly of the hour.

The duns of each species typically emerge during an hour or two each day for a couple of weeks in the spring or summer, though there are some important fall hatching species. Some species follow more sporadic emergence strategies, and many of these combine to create a sort of "background noise" of miscellaneous mayfly activity on many trout streams throughout much of the summer. This mixed bag of mayflies provides good opportunities for anglers to catch rising trout that aren't too picky.

Mayfly nymphs emerge into duns in several different ways. Most often, the nymph swims to the water's surface and splits open its exoskeleton above the thorax (Thorax: The thorax is the middle part of an insect's body, in between the abdomen and the head, and to which the legs and wings are attached.). The dun wriggles out onto the surface, where its wings fill with fluid hydraulically and allow it to take flight. (In contrast to the common myth of mayflies needing time to "dry their wings," this process is more like inflating a raft.) Different species may be quick or slow at each stage of this process. Some take a long time to escape their nymphal shuck (
Here's an underwater view of the pupal shucks of several already-emerged Brachycentrus numerosus caddisflies.
Here's an underwater view of the pupal shucks of several already-emerged Brachycentrus numerosus caddisflies.
Shuck: The shed exoskeleton left over when an insect molts into its next stage or instar. Most often it describes the last nymphal or pupal skin exited during emergence into a winged adult.
)
, making flies that imitate these "emergers" especially effective. Some species very quickly take flight when they hit the surface, while others ride the surface for some minutes like little sailboats, a prime target for hungry trout and a welcome sight for the dry fly angler. Cool or windy weather may prolong these struggles and increase the availability of mayflies to trout.

Many important species follow completely different emergence patterns. In some, full winged duns emerge on the bottom of the stream and float to the surface. Others swim to shore and crawl out on land before emerging. Learning to identify mayflies and associate them with the right behaviors gives an angler an advantage: the ability to make a good guess about which style and stage of emergence to imitate, simply from seeing and recognizing some duns or mature nymphs.

Spinner Behavior


Once mayflies have molted into spinners (imagos), they usually gather in swarms to mate, usually over riffles. When they're done they fall dead, or spent (Spent: The wing position of many aquatic insects when they fall on the water after mating. The wings of both sides lay flat on the water. The word may be used to describe insects with their wings in that position, as well as the position itself.), on the water in an event anglers call a spinner fall. Spinner falls are usually better coordinated than emergences, because spinners gather in swarms for mating. This means some species with sporadic, unnoticed dun emergences become far more concentrated and important to anglers as spinners.

Spinner falls are also usually more predictable than emergences, because so many of them (although not all) take place at dusk, and they are preceded by visible aerial spinner swarms, which may start hours earlier at treetop level and descend gradually toward the water as night falls. Dusk spinner falls often mark the angler's best chance to see a good rise of trout each day. However, like most things in nature, mayfly spinners aren't as predictable as we'd like. Sometimes clouds of thousands of spinners will gather over a riffle in the evening and fly back into the woods as quickly as they came, never falling spent (Spent: The wing position of many aquatic insects when they fall on the water after mating. The wings of both sides lay flat on the water. The word may be used to describe insects with their wings in that position, as well as the position itself.). When that happens, anglers must swallow their disappointment and look for them to finish the job in the morning.

Mayfly females face the extra duty of laying their eggs after mating. Many species release their eggs as they fall spent (Spent: The wing position of many aquatic insects when they fall on the water after mating. The wings of both sides lay flat on the water. The word may be used to describe insects with their wings in that position, as well as the position itself.) on the water near the males after mating. Some land on the water, release a few eggs and take off again. Others fly low over the water and dip the tips of their abdomens below the surface for just a moment to release eggs. Still others drop their eggs from high in the air. In one very common genus, Baetis, the females land near shore and crawl underwater to lay their eggs in neat little rows on rocks and logs.

Nymph Biology


Mayfly nymphs or "naiads (Naiad: Naiad is the technical term for nymph used by modern entomologists.)" grow underwater for a period ranging from 3 months to 2 years, depending on the species. Like the stoneflies of Plecoptera, their development follows "incomplete" metamorphosis, meaning they do not undergo a dramatic transformation to adulthood via a pupal stage like butterflies and caddisflies (Trichoptera) do. Their changes are more gradual, at least internally. As the nymphs grow they proceed through numerous slightly different developmental stages called instars (Instar: Many invertebrates molt through dozens of progressively larger and better-developed stages as they grow. Each of these stages is known as an instar. Hard-bodied nymphs typically molt through more instars than soft-bodied larvae.), between which they molt out of their old exoskeletons and expose new ones.

Anglers recognize four categories of mayfly nymphs: swimming, burrowing, clinging, and crawling:

  • Some streamlined swimmers move like little bullets, faster then fish of the same size, and they swim upstream against strong current without a problem. Others inhabit slow water and use their speed to dart between leaves in the weed beds.

  • Clingers of the family Heptageniidae are typically flat nymphs with strong legs and claws for holding on to rocks. Some have evolved further adaptations for clinging in fast water; for example, the genus Rhithrogena has gills resembling suction cups. There is great variation among the clingers and some species have adapted to slow water.

  • Crawlers come in the most varied forms; they are a catch-all group for "average" families that usually excel at neither swimming nor clinging. The Hendricksons and Sulphurs of the Ephemerella genus are typical crawlers. There are tiny crawlers like Tricorythodes, and there are oddballs like Baetisca. The crawlers in Leptophlebiidae are quite good at swimming, and those in Drunella (especially Drunella doddsii) are quite good at clinging.

  • The distinctive burrowers of Ephemeridae (and the less important Polymitarcyidae) are pale nocturnal creatures which use tusks to carve U-shaped burrows into the river bottom, where they live most of the time. Their long yellow bodies and feathery gray gills make them unmistakable. Their hatches are some of the angler's favorites, especially the giant Hexagenia limbata flies of the Midwest and West or the Eastern Green Drakes, Ephemera guttulata.



Entomologists have a similar system, but even their line between categories is a blurry one. Some burrowers swim well, some crawlers cling well, and some families like Leptophlebiidae and Potamanthidae straddle the boundary between categories.

If you fish a fertile stream, watch the bottom ahead of you as you walk. Sometimes, especially in April and May, you'll see lots of mayfly nymphs in front of you swimming out of your way or scurrying to the undersides of rocks. You don't need to be down on all fours with a magnifying glass to see mayfly life underwater.

Pictures of 678 Mayfly Specimens:

Specimen Page:1234...69
Hexagenia limbata (Hex) Mayfly NymphHexagenia limbata (Hex) Mayfly Nymph View 9 Pictures
Collected June 8, 2005 from the Namekagon River in Wisconsin
Added to Troutnut.com by Troutnut on May 26, 2006
Specimen Page:1234...69

41 Streamside Pictures of Mayflies:

Streamside Photo Page:12345
This Ephemerella invaria sulphur dun got stuck in its shuck trying to emerge.  This isn't exactly a "natural" pose for a photograph, but it kind of shows what an emerger pattern could look like.  In this picture: Mayfly Species Ephemerella invaria (Sulphur Dun). From the Neversink River in New York.
This Ephemerella invaria sulphur dun got stuck in its shuck (
Here's an underwater view of the pupal shucks of several already-emerged Brachycentrus numerosus caddisflies.
Here's an underwater view of the pupal shucks of several already-emerged Brachycentrus numerosus caddisflies.
Shuck: The shed exoskeleton left over when an insect molts into its next stage or instar. Most often it describes the last nymphal or pupal skin exited during emergence into a winged adult.
)
trying to emerge. This isn't exactly a "natural" pose for a photograph, but it kind of shows what an emerger pattern could look like.

In this picture: Mayfly Species Ephemerella invaria (Sulphur Dun).
Date TakenMay 20, 2007
Date AddedJun 5, 2007
AuthorTroutnut
CameraPENTAX Optio WPi
Often mayflies can be found on houses near the river.  This one molted from a dun into a spinner on the outside of our kitchen window.

Any lit dwelling near the river can attract a lot of mayflies at night.  A good way to determine what's hatching is to visit a gas station (or anything else with bright lights) close to the river early in the morning.  In this picture: Mayfly Family Baetidae (Blue-Winged Olives). From the West Fork of the Chippewa River in Wisconsin.
Often mayflies can be found on houses near the river. This one molted from a dun into a spinner on the outside of our kitchen window.

Any lit dwelling near the river can attract a lot of mayflies at night. A good way to determine what's hatching is to visit a gas station (or anything else with bright lights) close to the river early in the morning.

In this picture: Mayfly Family Baetidae (Blue-Winged Olives).
Date TakenMay 26, 2005
Date AddedFeb 8, 2006
AuthorTroutnut
Shed exoskeleton from what was very likely an Ephemerella aurivillii nymph that emerged on this rock.  In this picture: Mayfly Species Ephemerella aurivillii. From Mystery Creek # 170 in Alaska.
Shed exoskeleton from what was very likely an Ephemerella aurivillii nymph that emerged on this rock.

In this picture: Mayfly Species Ephemerella aurivillii.
StateAlaska
Date TakenJul 11, 2012
Date AddedJul 14, 2012
AuthorTroutnut
CameraCanon PowerShot D10
Streamside Photo Page:12345

67 Underwater Pictures of Mayflies:

Underwater Photo Page:1234...8
Several Baetidae nymphs line up on a rock.  In this picture: Mayfly Family Baetidae (Blue-Winged Olives). From Mongaup Creek in New York.
Several Baetidae nymphs line up on a rock.

In this picture: Mayfly Family Baetidae (Blue-Winged Olives).
Date TakenApr 19, 2006
Date AddedApr 22, 2006
AuthorTroutnut
CameraPENTAX Optio WPi
A crayfish chews on a Hexagenia limbata nymph shortly after a small Hex emergence.  I didn't catch any fish, but playing around with my flashlight and camera in the rocks proved productive.  In this picture: Arthropod Order Decapoda (Crayfish) and Mayfly Species Hexagenia limbata (Hex). From the Namekagon River in Wisconsin.
A crayfish chews on a Hexagenia limbata nymph shortly after a small Hex emergence. I didn't catch any fish, but playing around with my flashlight and camera in the rocks proved productive.

In this picture: Arthropod Order Decapoda (Crayfish) and Mayfly Species Hexagenia limbata (Hex).
Date TakenJun 14, 2006
Date AddedJun 30, 2006
AuthorTroutnut
CameraPENTAX Optio WPi
This picture from below shows a stillborn Ephemerella subvaria (Hendrickson) dun drifting on the surface amidst a number of shed pupal skins from Brachycentrus caddisflies which were heavily hatching that day.  In this picture: Mayfly Species Ephemerella subvaria (Hendrickson) and Caddisfly Species Brachycentrus appalachia (Apple Caddis). From the East Branch of the Delaware River in New York.
This picture from below shows a stillborn (
This stillborn Ephemerella subvaria dun is trapped in its shuck.
This stillborn Ephemerella subvaria dun is trapped in its shuck.
Stillborn: In fly fishing, a stillborn insect is one which got stuck in its nymphal or pupal shuck during emergence and floats helplessly on the surface instead of flying away. It is a specific class of cripple, although it is sometimes used interchangeably with that term.
)
Ephemerella subvaria (Hendrickson) dun drifting on the surface amidst a number of shed pupal skins from Brachycentrus caddisflies which were heavily hatching that day.

In this picture: Mayfly Species Ephemerella subvaria (Hendrickson) and Caddisfly Species Brachycentrus appalachia (Apple Caddis).
Date TakenApr 19, 2006
Date AddedApr 22, 2006
AuthorTroutnut
CameraPENTAX Optio WPi
Underwater Photo Page:1234...8

Recent Discussions of Ephemeroptera

rotunda 8 Replies »
Posted by CharlieBugs on Jan 26, 2015 in the species Ephemerella invaria
Last reply on Feb 1, 2015 by Gutcutter
Your post on Ephemerella subvaria brought back some memories that might be of interest to some readers.
I got my master’s degree under Ed Cooper at Penn State in 1966. I studied the impact of low oxygen from Penn State’s sewage plant on the mayflies of Spring Creek. The plant mostly removed BOD (organic matter which causes low oxygen) by oxidizing it in a bacteria rich environment. But at that time the plant did not remove phosphorus (and nitrogen) which fertilized the macrophytic algae and other plant growth. There were far more macrophytes (large plants) in Spring Creek below the sewage plant entrance than above, and essentially no mayflies. What was there in the effluent that killed the mayflies? Mayflies put directly in the effluent did not die over a 16 hour day. But oxygen samples taken over 24 hours in summer showed a much greater variation below the effluent (from 16 ppm (or mg/l), 160% of saturation in late afternoon) to 3 ppm (30 percent saturation just before dawn) (vs 14 to 10 ppm, +- 20 percent saturation above the effluent.
I built a Rube Goldberg machine in the lab that would control the oxygen levels and temperature of control and experimental cages that each held 25 mayflies. The control would keep oxygen near saturation and the experimental one would lower the oxygen over 8 hours (the length of night in summer). Mortality was dependent upon both oxygen level and temperature. Virtually no mayflies would die if the oxygen was above 2 PPM (or mg/l) at 8 C, or at 4.5 at 20 C. Seventy five percent of larvae would die at 1 ppm at 8 degrees and 2.5 ppm at 20 degrees. So the mortality was much more if the temperatures were high. Hence most of the mortality would presumably occur in August when the high temperatures (20C) would increase the larvae metabolism and hence need for oxygen, but the water would hold less oxygen even before the night-time respiration of the macrophytes would reduce it much further (to only 3 ppm). Thus the low nighttime oxygen caused by excessive plant growth was a sufficient cause or the near total absence of mayflies below the sewage plant. Recognizing the aquatic impact Penn State started to use land disposal of its effluent which as far as I knew alleviated, and even stopped, the negative impacts on the mayflies. Can anyone verify this ? The otherwise well done “The Fishery of Spring Creek; A Watershed Under Siege “
By Robert F. Carline, Rebecca L. Dunlap Jason E. Detar, Bruce A. Hollender Has nothing on dissolved oxygen or aquatic insects. In my opinion we need much more of an ecosystems approach for streams (Which we are doing for Little Sandy Creek in N.Y.).


Now back to Ephemeralla subvaria. The title of the paper I published on this project (my first of nearly 300 publications) was:
1. Hall, C.A.S. 1969. Mortality of the mayfly nymph, Ephemerella rotunda, at low dissolved oxygen concentrations. J. Elisha Mitchell Sci. Soc. 85(1): 34-39 (M.S. Thesis, Pennsylvania State University, 1966).

Whoa! Ephemeralla rotunda? This was by far the most abundant mayfly in Spring Creek where I sampled! But I could not even find the name in your list. Also I had my samples verified by Burke, author of the authoritative "Mayflies of Illinois", and he said Well that’s what it keyed out to in my book”. (Talk about scientific ass covering! ) Well to make matters worse (as of 24 hours ago) my next girlfriend, Molly, at the University of North Carolina, loved the name of “Ephemerella rotunda”, the rotund one, which comes to think of it described her as well. I liked it too. But n’exist plus: where had the most abundant mayfly gone? Fortunately upon reading the rest of the post I found “Ephemerella invaria is one of the two species frequently known as Sulphurs (the other is Ephemerella dorothea). There used to be a third, Ephemerella rotunda, but entomologists recently discovered that invaria and rotunda are a single species with an incredible range of individual variation." Ahh neither rotunda nor Molly stood the test of time. So I assume what I called rotunda is still alive and well in Spring Creek as invaria. Again, can anyone verify that?

If anyone wants to follow up on the distribution and abundance of mayfly (or any other species) may I recommend: Hall, C.A.S., J.A. Stanford and R. Hauer. 1992. The distribution and abundance of organisms as a consequence of energy balances along multiple environmental gradients. Oikos 65: 377-390. I can send it if you cannot get it from google, which I think you can. (chall@esf.edu)
ReplySo is Ep Infrequens now known as Ep Dorothea? 20 Replies »
Posted by Wbranch on Feb 17, 2008 in the species Ephemerella dorothea infrequens
Last reply on Jul 1, 2014 by Crepuscular
These mayflies look more like the Sulfurs I see on the Delaware system than the PMD's I see in Montana. The Montana mayfly has a distinct yellow leading edge to an overall light dun gray wing and the abdomen and thorax have a more light greenish/yellow cloration so how is it that Infrequens is now known as Ep Dorothea Dorothea?
ReplyPredicting a hatch? 2 Replies »
Posted by U2ill on Apr 2, 2008
Last reply on Jun 24, 2014 by TNEAL
All,

Is it possible to predict a hatch (whether mayfly, caddis, stonefly, etc) based off of water, air, and weather data? So if I know streamflow, water temp, air temp, wind, cloudiness, etc...and I know what insects are on a stream...is that enough data to make a prediction like "There would be a good likelihood of a caddis hatch this afternoon"?
ReplyHex hatch water temperature range? 3 Replies »
Posted by NEMatt on May 23, 2014 in the species Hexagenia limbata
Last reply on May 25, 2014 by NEMatt
Hi,

New to the site - love it. I was wondering if there was a suggested range of water temperature at which the Hex likes to hatch.

Thanks
Matt
ReplyGreen Drake Hatch Temp? 1 Reply »
Posted by NEMatt on May 23, 2014 in the species Ephemera guttulata
Last reply on May 24, 2014 by Entoman
Hi

I was looking for a water temperature range for the Green Drake hatch. Anyone know?
Reply
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