Blobs of caddisfly eggs (Nemotaulius sp.) laid on a willow branch overhanging the water along a wetland. |
An adult Baetis mayfly spinner penetrating the surface film as it makes its way under water to lay eggs. |
Blobs of caddisfly eggs (Nemotaulius sp.) laid on a willow branch overhanging the water along a wetland. |
An adult Baetis mayfly spinner penetrating the surface film as it makes its way under water to lay eggs. |
A scene that normally plays out underwater, I flipped this log to find an adult caddisfly laying its gelatinous mass of eggs on the underside.
If you are not familiar with the life-cycle of the Caddisfly, this photo series should give you a few fly tying ideas to go on.
A typical caddisfly life-cycle. |
Anabolia sp. (Limnephilidae) larva - one of the larger caddisfly larvae found in Alberta lakes. |
A close-up of Anabolia with its cool color pattern. As the larva grows it simply adds on to the front end of the case to make it bigger. |
A cut-away showing the pupa developing within the case. If oxygen levels drop too low, the pupa will undulate (similar to cased chironomid larvae) to help circulate water through the case. |
This pupa has almost completed its development. Note the well-developed legs, antennae, and wing-case all bunched together on the front half of the pupa. |
Once at the surface, the cuticle of the pupa splits along the back and the adult caddisfly crawls out onto the surface. |
If you see lots of these on the surface you know that caddisflies have been or are emerging. |
A typical adult caddisfly with its characteristic drab color and long antennae. |
Caddisfly hatches are more common late in the evening but you will sometimes see an emergence in the middle of the afternoon. |
Mating takes place on shore-line vegetation with egg laying taking place a short time later. |
Most of the larger species of caddisflies found in Alberta Lakes are in the families Limnephlidae and Phryganeidae. Both groups have some interesting case building behaviors, with each case designed to help the larva blend into its environment and avoid predators. Natural materials are scavenged and arranged in ways the enhance the camouflage provided by the case. Note that these photos were purposely taken with each caddisfly larva in an unnatural setting to make them more visible in the photos.
Caddisflies are closely related to the Lepidoptera (moths and butterflies). And like caterpillars, caddisfly larvae can produce silk from glands near their mouth, which they use to line their case and hold it all together.
If you have the patience to fish a cased caddisfly pattern in lakes, it can produce some nice fish. The trick is to keep your fly on or near the bottom, and move it ever so slowly. Fly patterns are usually quite simple - I have seen woolly worms with the palmered hackle trimmed short work well. One of my favorite patterns imitates the larva of Phryganea cinerea (Phryganeidae) with its case of spirally arranged short pieces of grass.
When it comes to fly design it helps to have a good idea of what the actual bugs looks like. Being able to take your time to really assess the shape and color of an aquatic insect in its natural environment gives the fly tier a big advantage. But most bugs rarely sit still for very long, and aquatic larvae removed from the water tend to look darker and more clumpy than they actually are. A bug in the hand may beat two in the bush, but a good photo trumps them both.
A Dragonfly darner nymph (Aeshna sp.). You can tell that this one has more growing to do by the small wing-pads. |
A Dragonfly sprawler nymph (Cordulia shurtleffii) This one has fully developed wing-pads and is ready to emerge. |
This is what Cordulia shurtleffii looks like as an adult. Its metallic-green eyes earned this species the common name of American Emerald. |
An adult Variable Darner (Aeshna interupta) laying eggs in a rotten log. |
I have been working on some woven-body dragonfly nymphs this week. Reg Denny taught me how to do this many years ago. It makes for a very realistic looking abdomen on any dragonfly pattern - including the longer and darker darner type patterns. The first one imitates the stubby sprawler type dragonflies during the spring pre-hatch migration.
A short, stubby woven dragonfly sprawler imitation. |
I'm one of those tiers who doesn't always follow a recipe - most of the time I just wing it by memory. This dragonfly patterns start with two lengths of lead free wire laid along either side of the hook shank - this gives the body a somewhat wide and flat appearance like the naturals. Leave enough head space to attach the eyes. The hook for sprawlers is usually a 2XL #8 to keep the fly short and stout. I do up a bunch of the hooks with wire ahead of time and make sure the wire is tied down well and add a little head cement to keep it all in place.
The weave is an alternating over & under using two strands of yarn in slightly different colors - one color for the dorsal side and one for the ventral. If I haven't tied them for a while it seems I need to re-learn the technique and practice a bit before I start on the actual flies. There are a few YouTube videos out there that cover the woven body technique better than I can explain it. My preference is for soft legs using tufts of dyed mallard flank - this allows the legs to sweep back when using a darting retrieve. The thorax is not woven - just a simple wrap spaced to keep the legs in position. I used bead-chain for the eyes on these ones but I also use glass beads on heavy monofilament with the ends melted sometimes. The wing case is traditional turkey quill. Colors for the sprawls range from light olive to olive-brown.
A longer-bodied woven dragonfly darner imitation. |
The darner pattern is tied on a 4XL #8 hook using two strands of brown yarn in slightly different shades. Fine "nymph" rubber legs have a good action on the larger fly. Make sure to make the abdomen longer relative to the thorax for this one.
A dragonfly nymph (Epitheca spinigera) fresh out of the water and making its way up shoreline vegetation in search of an emergence site. |
It is not easy wiggling out of the nymphal exoskeleton - gravity is used to aid in the process. |
Once free, it can take up to an hour for the body and wings to fully extend. |
Wings and body starting to take shape. |
Almost ready to fly - the wings just need to harden a little more and extend out to the sides. |
A close-up of the large compound eyes. |
As soon as the adults can fly, they make their way to nearby trees or bushes to rest and allow for any final hardening to take place. (Spiny Baskettail - Epitheca spinigera) |
In my last post I talked about how chironomid larva that build tubes or cases use body undulations to maximize oxygen intake. This behavior is used by not just chironomid larva living in the hypoxic zone on the bottom, but also by chironomid and caddisfly larvae that build portable cases. They do this when oxygen levels are low or they need to refresh the stagnant water in their case.
This video shows how a chironomid (midge) larva undulates in its tube to circulate water to aid in respiration. This particular genus (Cricotopus) builds a portable tube, while other chironomid groups construct more extensive stationary tubes in the mud. The video also shows how chironomid larvae that build portable tubes move by reaching out and pulling themselves forward. In the final scene, the midge larva gets caught on a strand of vegetation and uses a small pebble to leverage itself up and over.
Note: This post may not make you a better fly
angler, but it will make you a more knowledgeable one.
One of the pitfalls of living in the mud at the
bottom of a wetland or lake is that there can be very little oxygen present. Oxygen
diffuses much slower through water than it does in air, and even slower through
the mud barrier where the decay of organic debris further depletes what little
oxygen is present. Under these low oxygen (hypoxic) conditions, life is limited
to organisms that are adapted to survive in these extreme environments.
Some chironomid (midge) larvae have adapted to
exploit these hypoxic habitats through the use of hemoglobin. Hemoglobin is the
protein component of red blood cells that circulates through our body and is
responsible for delivering oxygen to tissues. It also gives blood cells their red
color. Hemoglobin has a high affinity for binding to oxygen molecules making it
the perfect oxygen delivery system where constant circulation is present.
There are several groups of chironomids that
utilize hemoglobin to aid in survival under hypoxic conditions, but the two
most common ones are in the genera Chironomus
and Glyptotendipes (commonly known as blood-worms). The high concentration of hemoglobin in their body fluids
(hemolymph) gives them their distinctive red color. While there can be some passive
movement of the hemolymph through their body, it does not circulate like our
blood does. Instead, it simply surrounds the organs and tissues.
Note
that more evolved and larger insects may have an open circulatory system, but
even here, the hemolymph does not circulate very fast and oxygen uptake and
transport is primarily via the air-filled tracheal system.
A Chironomus larva with its typical bright red coloration. |
So how exactly does the hemoglobin help blood-worms
survive in hypoxic and sometimes anoxic (no oxygen) habitats? If there is
little to no oxygen to bind to the hemoglobin to begin with, what advantage can
there be?
Most blood-worms build extensive tube networks in and on the mud by tunneling and reinforcing the tube walls with mucus. These tubes provide protection from predators but they also allow the midge larva to circulate water through the tube by undulating its body within the tube (caddisflies also do this). This action pulls water with a slightly higher oxygen content through the tube. In some cases, the midge larva will also elevate the tube opening, even by just a few millimeters, to place the tube intake above the low oxygen boundary layer at the water-mud interface. Of course under low oxygen conditions they can't spend all of their time undulating to gain oxygen, they need to feed too. That is where the hemoglobin comes in - as the larva undulates and circulates water through the tube, the hemoglobin picks up oxygen (much like recharging a battery) - then when the larva stops undulating to feed; oxygen bound to the hemoglobin diffuses into the tissue to sustain metabolic activity. Once the hemolymph oxygen level falls to a critical point, the midge larva will once again take up its ventilation position in the tube to recharge its oxygen supply.
A Glyptotendipes larva out of its tube and foraging for food. |
The most productive part of a lake is the littoral zone where the penetration of sunlight allows for the rapid growth of aquatic plants and periphyton. Here is where you will find the highest level of bug biodiversity, including a hundred or so species of chironomid (midge) larvae. But this doesn’t mean that the deep water areas of a lake are devoid of life. In fact, there are some genera of chironomids (Chironomus and Glyptotendipes) that specialize in hiding in the low-oxygen layer of mud during the day, and coming out at night to forage for whatever tiny organic particles of food they can find.
While the diversity of midge larvae living in the
mud 15 to over 60 feet down is low, the biomass (number of organisms) can be
very high. This is because, with very little oxygen to support life in the hypoxic mud layer, there is little competition for food. The midge species that
can live in this layer of mud are able to monopolize on the resources, much
like the mono-culture of a wheat field.
One would think that there is not much to eat way
down there in the depths, but as organisms living in the upper layers of the limnetic
zone die (algae, zooplankton and cyanobacteria), or organic bits get blown onto
the lake’s surface (pollen and organic dust), they settle to the bottom. This
translates into a continuous shower of food for most of the year, even in the
deeper parts of a lake.
This video shows midge larvae foraging for food on
the lake bottom and their swimming behavior when they decide to move longer
distances to find better foraging sites. Their swimming behavior is unique, and
somewhat frantic, and not easily imitated by the fly angler.
Every fly tier who frequently fishes scud patterns has an opinion on this; should scud patterns be tied on straight shank or curved shank hooks?
Scuds at rest tend to have a very pronounced curve.
This observation has resulted in a large number of scud fly patterns being tied
on radically curved hooks – what used to be called scud or shrimp hooks.
A Scud (Gammarus lacustris) in a typical resting position. |
Here is an article I wrote for the 2016 fall issue of Fly Fusion magazine. Fly Fusion is probably the most informative and artistic fly fishing magazine in print. I have not written for them lately but I do still have a subscription - something I would suggest to anyone with an interest in fly fishing. This article was written with Halloween in mind. I hope you enjoy it.
It was clear from the get-go that something strange was afoot. Was this really happening? Could I be dreaming? This just didn’t make any sense...
Strange indeed; knowing what was normal, what
was expected, I was having a difficult time reconciling what I was bearing
witness to. The funny thing was that I was not the least bit afraid. The fact that
it was already mid-morning may have had something to do with my mindset leaning
more to simple curiosity than to morbid misgivings, but with the seemingly
sinister events unfolding before me, I simply had to know more.
I cautiously waded a little closer to a
small patch of weed growth in shallow water. Three brook trout circled above it
like sharks around a wounded mackerel, their dark olive backs and white fin
edges standing out in stark relief against the verdant background. Every so often
a pale colored scud could be seen spiraling lazily towards the surface, only
to be quickly picked off by the nearest fish. Suicide, I thought. What would possess these scuds to leave the safety
of the weeds like that, in broad daylight no less? I knew they didn’t hatch
into a winged terrestrial form like aquatic insects do. They really had no
reason to head for the surface. It would be several more years before I would
learn that those scuds were in fact possessed – infected by an insidious
parasite and no longer in control of their own bodies. Yes my friend, zombies
really do exist.
Scuds
normally lead peaceful and unassuming lives. Much of their time is spent simply
scavenging among aquatic vegetation and along the bottom for bits of dead
plant and animal material to eat. As scuds grow they periodically shed their plate-like
exoskeleton through a process called molting. Once maturity is reached they
will form mating pairs, swimming in tandem as they propagate the next
generation. The eggs develop in a pouch on the female called a marsupium and
the young are born looking just like their parents, only smaller. There is no
larval stage or metamorphosis. The average lifespan for a scud is about one
year and depending on conditions they may produce from one to several
generations in a summer. That effectively sums up the serene cycle of life for
a scud.
Of course being a perfectly packaged piece
of protein means that scuds are a sought after food source. Many birds and fish
will go out of their way to chase down any scuds that have been careless enough
to get caught out in the open. Unfortunately for the scuds they have little to
offer in the way of escape strategies. Even with all of those legs scuds are
relatively poor swimmers – often swimming in slow spirals, sometimes on their
side, and sometimes even upside down. And with their Mr. Magoo-like eyesight they
are constantly bumping into things. With antics like that it’s no wonder that
scuds make up a significant part of a trout’s diet wherever the two coexist.
For the slow moving scud staying hidden is really
the only available defense. Cryptic coloration that matches their habitat is a
good start. Being found in the weeds, or prowling in the muck, scuds tend to be
shades of green, olive, or olive-brown. They can’t quite change their color with chameleon-like adeptness but scuds can adapt over time to changes in the
background color of their environment using subtle shifts in shading. Staying
out of sight is another strategy that keeps scuds safe. Scuds are normally
photo-phobic – meaning they tend to avoid the light, preferring instead to frequent
dark and shadowy places. During the day they can be found tucked deep in the
vegetation or hiding under logs and debris. It is during periods of low light that
scuds tend to become more active, sometimes venturing out into the open (even
cloudy days can be enough to coax them out of hiding). Regardless of the lighting
conditions, a scud’s instinct is to always dive for cover when threatened.
Knowing all of this it’s easy to understand
why I was so intrigued upon witnessing the exact opposite behavior on my
earlier fishing expedition. And here is where our story takes an unsettling twist, one that fly anglers
can use to their advantage.
Parasitism is not uncommon in the
invertebrate world. Small bugs often serve as intermediate hosts to even
smaller bugs looking for a free lunch. One of the most common scud parasites is
a tiny creature known as a spiny-headed worm (Acanthocephala). Scuds become
infected with spiny-headed worms when the scud ingests eggs that are mixed in
with the organic debris they are feeding on. The eggs hatch in the gut and the “larvae”
(acanthella) burrow through the scud’s intestinal wall and into the body cavity
where they continue their development. The acanthella develop into sub-adults
known as cystacanths – at this stage they are fully formed worms but lack
reproductive structures. These tiny worms migrate to a central region, usually
along the back of the scud, where they encyst and wait for a definitive host to
eat the scud. Depending on the parasite species the definitive host is usually
a fish or bird. Final development and reproduction takes place in the gut of
the definitive host and the parasite eggs are released when the host animal
defecates. Scuds feed on the feces and the cycle continues.
Once the first phase of development has been
completed within the intermediate host, the life of the spiny-headed worm (or
continuation thereof) hangs in the balance. If the scud is not eaten by a
definitive host the scud will eventually die of old age, and all will be lost
for the progeny of the spiny-headed worm. Not leaving things up to chance, the
ancient acanthocephalan has evolved a few clever ways to beat the odds. By manipulating the hormonal system of the
scud, acanthocephalans are able to control the scud’s behavior. Instead of
avoiding the light, infected scuds in the cystacanth stage become attracted to
light and are more active during the day. Infected scuds will also develop a
tendency to swim towards any disturbance in the water (such as feeding fish or
birds) rather than head for cover. Acanthocephalans also make infected scuds
stand out more by altering their coloration. Cystacanths are typically bright
orange or yellow – they can easily be seen in infected scuds as a small spot
along the back. But more importantly, the cryptic coloration of a normal scud often
becomes muted and takes on a pale, bluish-grey, ghost-like hue as the infection
progresses (freshly molted scuds will also exhibit the same bluish-grey color but they will lack the orange spot and will have an even greater tendency to
stay hidden). These two factors combine to make an infected scud stand out like
a neon sign. No guarantee that the scud will be consumed by the correct
definitive host, but quite likely that some sort of fish or fowl will gobble it
up.
A scud (Gammarus lacustris) in the early stages of infection, before the natural color has faded. |
In fact, it has been shown experimentally
that fish will preferentially feed on infected scuds even when both infected
and uninfected scuds are readily available in equal numbers. This should come
as no surprise. Most fish rely heavily on search images to direct their feeding
effort. If they are seeing and successfully capturing more pale scuds than
normal ones, the search image for the pale scuds becomes a stronger feeding
trigger.
A scud (Gammarus lacustris) with its natural color faded to a bluish-grey due to the infection. |
If infected scuds stand out more, and are
eaten more frequently than their uninfected counterparts, doesn’t it make sense
to use a fly pattern that specifically mimics an infected scud? Once I had all
of the facts, it didn’t take me long to answer that question for myself. Several
years ago I tied a bunch of simple baggie shrimp on #10-12 hooks, some
using pale blue dun rabbit dubbing and some with the traditional olive dubbing.
I added a small orange dot to the back of some using a marker pen but left a
few of each color unmarked. After testing these patterns on several local trout
lakes the olive scud with the orange spot turned out to be slightly more
effective than the plain olive pattern, while the blue dun scud out fished both
olive patterns by a wide margin. The presence or absence of the orange spot on
the blue dun scuds didn’t seem to matter. This experiment was all very
unscientific of course, but the results were enough to convince me to permanently
add the Blue Dun Scud to my repertoire.
In the intervening years I have come across
fly tying recipes for other scud patterns in blue, light grey, and pearl – all
good candidates for an infected scud imitation. I prefer the simplicity of the
old-school blue dun baggie shrimp but some of the newer synthetic fly tying
materials are certainly worth experimenting with. The goal is to get that translucent
bluish-grey look of a zombie scud. Of course I still carry olive scuds for
those times when olive is what the fish seem to prefer, but more often than not
it is the blue dun scud that is in tatters on the end of my line.