Habitat Heterogeneity Through Time and Space

Habitat heterogeneity is a fun jargon term within the fields of ecology and land stewardship.  It’s a good one to know if you’re at a party trying to impress an ecologist.  Although, to be honest, if you’re trying to impress an ecologist, going to a party is probably not the best way to start.  You’d be better off meeting them “accidentally” on a hiking trail and bending down to closely examine a set of tracks or a fly feeding on a flower.  Ecologists are impressed when people notice things in nature.

Regardless of whether you’re trying to impress an ecologist, habitat heterogeneity is an important concept.  Most commonly, it is used to describe the variety of habitat conditions available in a particular location.  It’s generally good to have numerous habitat types represented across a site that each provide something a little different in terms of food or shelter for animals or growing conditions for plants.  That kind of variety supports the habitat needs of a broad diversity of species.

This prairie has at least four different habitat types available, including variation in the height and density of vegetation, as well as what species of plants are blooming.

It’s not hard to imagine why lots of different habitat conditions would lead to a high number of species thriving in a prairie.  However, if you really want to impress an ecologist, you’ll also consider the time elements, not just the spatial elements of habitat heterogeneity.  In other words, it’s not just important to vary the kinds of available habitat across space, it’s also important for each of those habitat patches to change through time.

Good farmers and gardeners know the value of rotating crops.  Rotation can help soil productivity because each crop uses soil resources differently.  Shifting the location of crops from year to year also helps reduce impacts from diseases and insect herbivores that can build up over time when their favorite food source is available year after year. 

Many of the same concepts apply very well to prairie and other natural land stewardship work.  When habitat conditions remain static in a given location, bad things can happen to the inhabitants of that patch.  Disease organisms (pathogens), parasites, and herbivores can build up high populations.  Predators can learn to be very effective at finding and catching prey.  Within plant communities, the plants best adapted to the current conditions will thrive at the expense of others.  If those conditions don’t change, the plants less adapted to those will struggle to survive.

Here are two specific examples to illustrate why variation through time matters so much:

Years ago, a wildlife refuge manager told me about a situation he’d encountered early in his career.  He was charged with managing wetlands for ducks.  Ducks like nesting in dense vegetation, so he was told to make sure all the wetlands in his site had thick cover around them to maximize the number of duck nests.  After a few years of this, though, he realized that nesting success was actually very low, even though he was providing the habitat ducks supposedly wanted. 

What he thinks happened is this: The consistent dense vegetation allowed the vole population around wetlands to grow.  That higher population attracted predators like bull snakes to feed on the voles.  Unfortunately, those predators also enjoyed eating duck eggs, which worked out poorly for those ducks. 

Bull snakes are just one of many predators that like feeding on both voles and duck eggs.

Regardless of whether the manager’s hypothesis was right or not, when he started varying the habitat structure from year to year, nesting success went up.  He managed the site so that each year, some of his wetlands were surrounded by dense cover and others weren’t.  Every nesting season, he’d change which wetlands had thick vegetation around them.  Even though there was less overall nesting cover each year, ducks did better because nests suffered from lower rates of predation.

The second example comes from my own experience.  Canada milkvetch (Astragalus canadensis) can be frustrating to harvest seeds from because the pods are often empty by the time we get to them.  Those empty pods are easily identified because of the exit holes left by larvae of a small weevil (Tychius liljebladi) that feed on the seeds and then burrow out of the pod when they’re done.  After leaving, the larvae dig into the soil beneath the plant, where they overwinter.  The next summer, they emerge as adults and will lay eggs on the flowers in mid-summer to continue their life cycle. 

Holes left by weevil larvae usually indicate there’s no point picking seed from a particular patch of Canada milkvetch plants.

I’ve seen populations of Canda milkvetch go many years in a row without ever producing seeds because of the recurrence of those weevils.  That’s not good for the longevity of a population.  However, we’ve found that if we can break that annual pattern, we can get a year of strong seed production, which is good for both us and the plants.  The easiest way to break the cycle is to prevent the plants from flowering for a year.  If the weevils emerge from the soil and don’t see flowers to lay eggs on, they’ll fly off in search of flowers elsewhere.

The following year, when Canada milkvetch blooms again, a few weevils might travel from other places to breed on the flowers, but there aren’t high numbers of them popping out of the soil right beneath the plants.  Usually, that results in low rates of seed predation.  Even delaying flowering by a few weeks can help.  In years where milkvetch plants are mowed or grazed in a way that makes them bloom later in the season, they tend to produce lots of seed because the weevils were done breeding by the time the flowers appeared.

Following the crop rotation model, the best way to vary habitat conditions through time is to create a kind of ‘shifting mosaic’ of habitat patches across a site.  The ‘mosaic’ part of that phrase represents the variety of available habitats.  Those can include patches with tall, thatchy vegetation as well as others with short plants and exposed bare ground – and still other patches where that tall and short structure is intermixed.  It’s also helpful to have patches differ in the plant species that are blooming or otherwise thriving.  That might include some patches where annual plants are temporarily flourishing instead of the usually dominant perennials.  Having all those patches arrayed near each other allows animals to move between them to look for food, escape predators, seek out shade or sunlight, and anything else they need.

The ‘shifting’ aspect of the mosaic, though, is also crucial.  Each habitat type should appear in a different place each year, or on some similar time scale.  That can be created by simply mowing or burning different patches within a site at varying times of the year.  You might end up with a patch that was burned in the early spring, another that was mowed in mid-summer, and another that hasn’t been mowed or burned for a while.  That idled patch could then be burned the following year and the previous year’s burned patch could be mowed sometime during the growing season.  Or whatever. Basically, you want to shake things up each year so you’re applying management in different places over time and always providing a good mix of habitat options.

Grazing can be used the same way, except that there’s a lot more flexibility in how it’s applied.  Patch-burn grazing is a terrific way to combine fire and grazing to create heterogeneity across time and space.  I’ve written about the ‘open gate rotational grazing’ approach we’ve been experimenting with.  That’s another potential option.  In both those examples, each part of a prairie is grazed for a full season or more and then allowed to rest for a couple years.  The grazing and rest cycles are staggered across the site so there are always patches in each stage of that cycle available for the plants and animals that thrive best in those conditions.

Patch-burn grazing is just one of many ways to create a shifting mosaic of habitats. Here, cattle are grazing a recently burned patch while mostly ignoring an adjacent unburned patch. Not shown in the photo are other nearby areas with more patchy habitat structure. Each of those provides different cover, foraging opportunities, and growing conditions for animals and/or plants.

Those grazing approaches, though, are only a couple examples among an infinite array of choices.  You can vary the stocking rate, timing, and duration of grazing to achieve lots of different results.  You can combine grazing with fire and/or mowing to create even more variation.  There’s lots of room for creativity, but more importantly, to observe, learn, and adapt over time.

Regardless of the stewardship options available to you, the key is to look for ways to provide a good variety of habitat options for animals and growing conditions for plants.  More to the point of this post, it’s important to make sure those habitat options shift around in location to help prevent anyone from winning too consistently at the cost of others. 

Or, if you’re still trying to impress that ecologist, you might say that you’re trying to foster both spatial and temporal heterogeneity.  If that doesn’t work, you might try talking about how fascinating parasitoid-host relationships are.  That always gets me going.

Flies That Can Eat You (if you’re a tiny invertebrate)

Flies are an astoundingly diverse and important group of organisms. I’ve written about them quite a bit, including broad overviews and more specific pieces about behavior, etc. I’m going to keep writing about them and there’s nothing you can do about it. I mean, you don’t have to read what I write, I guess. And if enough of you stop reading, I’d probably stop writing. But apart from that, just try and stop me!

Today, I want to talk about predatory flies. Because of their incredible diversity (estimated 61,000 species in North America), flies fill lots of roles in ecosystems. As adults and larvae, they are pollinators, scavengers, herbivores, decomposers, parasites, and more. A few of them even come into our dwellings, buzz around our windows and lights, and taste our food. If flies disappeared from the earth, it would be a huge problem, notwithstanding those food-tasting buzzers.

A robber fly (left) feeding on a tiger beetle in the Nebraska Sandhills.

If you know anything about flies as predators, it’s likely you’re familiar with robber flies. That’s a good start, though there are others we’ll get to presently. Robber flies include more than 7,500 species worldwide, according to bugguide.net. They are impressive aerial predators. The ones I know tend to hunt from perches, zipping into the air as another insect flies by. They streak toward their prey and intercept them violently – sometimes knocking them out of the air altogether.

Once they have their target, robber flies insert their sharp mouthpart and inject digestive chemicals into their prey’s body. Once their prey is sufficiently liquefied, robber flies suck out the good stuff and discard the husk. This is the basic strategy used by many of our coolest invertebrate predators, by the way, including spiders, assassin bugs, and others.

To clarify, it’s the liquefying process that’s used by many others. Spiders and assassin bugs don’t launch themselves through the air like missiles at airborne prey. That would be amazing, though, wouldn’t it? Can you imagine how people would feel about spiders if that was true? Its not that much more crazy than what people already think spiders can do, I guess. That’s another topic for another day, though.

A robber fly laying eggs, showing off its distinctive long, slender abdomen.
A robber fly feeding on a small bee.
This robber fly knocked a huge cicada out of the air. It then had to search for a chink in the armor so it could insert its mouthpart to dispatch and feed on its prey.
Here’s a bumblebee mimic robber fly. You can tell it’s not a real bumble bee because of its bigger eyes, shorter antennae, and better beard. Oh, and the beetle in its mouth.

Anyway, robber flies are incredible. While they tend to have long bodies, huge eyes, and a glorious beard (technically called a “mystax”), robber flies can vary quite a bit in their appearance. Famously, some of them are excellent mimics of bees, which is surely a huge advantage when hunting pollinators. Maybe it also deters predators as well, though I’m not sure a bee is any more intimidating than a bearded, bug-eyed surface-to-air missile with chemical weapons.

As I continue to learn about flies, though, I keep finding out about other predator groups. The first were the longlegged flies. I’d known about, and photographed longlegged flies for years before I discovered that those exquisite, shiny little creatures were eating other insects. I became aware of their predation when I photographed one carrying around part of a tiny ant (or some other small creature – I couldn’t tell for sure) in its mouth. When I rushed to the internet to learn more, I found out they are, indeed, predatory. I also learned there are thousands of species of them (because of course there are).

A longlegged fly (family Dolichopodidae) in my square meter plot this year. What a gorgeous creature, huh?
Longlegged fly with prey in my square meter photography plot this year.
Predators are also prey. This longlegged fly was the victim of another common predator in my square meter plot – the striped lynx spider.

I see longlegged flies frequently when I’m in the prairie, especially when I have a camera in my hands and my brain is switched to photographer mode. Usually, they’re hanging out on leaves of plants, maybe a foot or so below flowers. At least, that’s where I find them. They often move around pretty quickly, both by leg and wing, as they search (I assume) for itsy bitsy creatures to eat. I wonder if their prey is as surprised as I was that longlegged flies are predators? If so, at least they learn something before they die, I guess.

My most recent discovery of a fly predator was during this year’s square meter photography project. The discovery process was the same as with longlegged flies. I focused in on a fly and realized I was photographing two species at the same time – the fly and a leafhopper that it was sucking the life out of. Bugguide told me it was a tiger fly and that it’s taxonomic family, Coenosiinae, includes a couple thousand species of predatory flies. How is this not more widely known?

A tiger fly with a captured leafhopper in my square meter plot this year.

Once I knew what to look for, I started seeing both tiger flies and longlegged flies more often, with and without prey. It makes me wonder what other predatory flies are out there that I’m not yet aware of, let alone all the other amazing stories I’m ignorant of. That’s one of the reasons I’ve loved my square meter photography work. By focusing my attention more narrowly, I’ve made discoveries that have really broadened my understanding of how prairies work. Weird, huh?

I’d love to hear from anyone who knows of other predatory flies out there. One group I am aware of is the syrphid flies (aka flower flies and/or hover flies), which have predatory larvae. That’s pretty great, though I’ve yet to see and photograph one of those larvae eating aphids or other little creatures. I did photograph some syrphid larvae this summer, but they were feeding on pollen, not other animals.

While they’re technically categorized as parasitoids, not predators, the larvae of other fly species also feed on animals. Tachinid flies are a good example, and I see them around quite a bit. Parasitoids are the kind of creatures horror movies are inspired by. They lay their eggs on or in other creatures and their larvae hatch out and eat the victim’s insides while it’s still alive. They’re extraordinarily important, ecologically, but also creepy. And cool.

What other predator flies should I be on the lookout for? I know about bot flies, but, again, they’re parasites and it’s the larvae that are feeding on animals. Any other adult flies that capture and eat other creatures?

It’s an amazing world out there, huh?