That's Not an Ant?!

It is always best to avoid making assumptions about insects in the field. I was reminded of that on three seperate occasions this month, all of them involving mimicry of ants by small longhorned beetles in the family Cerambycidae. Early spring is apparently the season for these wood-boring insects, at least here in Kansas. Their resemblance to ants is uncanny, and highly convincing.

Cyrtophorus verrucosus

While exploring our yards and looking for insects on April 2, I noticed what I thought was a carpenter ant climbing on the exterior of our detached garage. I was about to dismiss it, but decided to document it anyway, as it would be new for the year. Imagine my surprise and delight to discover it was instead an adult Cyrtophorus verrucosus. This species has no common name, which is unfortunate given its uniqueness.

The beetle averages only 9 millimeters in body length. Larvae bore in a variety of host trees, including maple, hickory, oak, elm, birch, chestnut, dogwood, Eastern Redbud, American Beech, and even pine. The adult beetles are active from March to July, but are most abundant in early spring. Cyrtophorus verrucosus ranges over most of the eastern U.S., and Canada as far west as Alberta. This was a new species for our property.

I was invited to present the keynote address for a joint meeting of the Kansas Academy of Science, and Central States Entomological Society, in Wichita, Kansas on April 5, and Heidi and I stayed over to go exploring on Sunday, April 6. I reached out to a Facebook and iNaturalist friend, Ryan Philbrick, and we met at the Kansas Wildlife Exhibit in Central Riverside Park. From there we went to Keeper of the Plains, and walked the nice concrete trail along the Arkansas River. At one point, Ryan turned over a small, broken tree limb on the ground. He noticed millipedes and a small "ant" crawling over the bark.

Euderces pini

I trained my camera on the ant and discovered it was another species of longhorned beetle, Euderces pini. At only 7-8 millimeters, and slender, it is even more ant-like than other species. The larval host trees include Flowering Dogwood, Pecan, Winged Elm, and American Beech, but there are several other hardwoods on the menu.

The adult beetles are frequently found on flowers, feeding on pollen and nectar. Even dandelions will suffice until native trees start blooming. Euderces pini occurs in the southern half of the eastern United States, and appears as an adult from March to June.

Currant-tip Borer

Back home, yesterday, April 13, I happened to notice a small ant-like insect on the central air-conditioning unit in the back yard. It was even smaller than the other beetles, but at least it had a common name: the Currant-tip Borer, Psenocerus supernotatus.

Despite its name, this 4-millimeter beetle has been reared from plants other than currant: Green Hawthorn, Tulip Tree (aka Tulip Poplar, Yellow Poplar), and Black Willow. This beetle is widespread in North America, east of the Rocky Mountains, and the adults are found from April to June.

None of these three beetles are considered pests, maybe because they do not affect crops or orchard trees. Their earthtone colors make them rather cryptic, but they are quite active, running rapidly, and flying well.

An actual ant

It is interesting to note the convergence in body shape and coloration between the different genera. They each have white streaks or bars designed to suggest body segmenation where there is none. Ants show marked constrictions between the head, thorax, and abdomen. In these beetles, the thorax may be mistaken for the enlarged head of an ant, and the elytra (wing covers), "divided" into thorax and abdomen by those white markings. The true head of the beetle is relatively small, and in the case of the Currant-tip Borer, directed downward and nearly hidden by the thorax.

There are even more ant-mimicking longhorned beetle species. Give that "ant" a second glance, especially if you see it in isolation, with no other ants nearby. It may turn out to be a beetle instead.

Sources: Lingafelter, Steven W. 2007. Illustrated Key to the Longhorned Woodboring Beetles of the Eastern United States North Potomac, Maryland: Coleopterists Society, Special Publication No. 3. 206 pp.
Yanega, Douglas. 1996. Field Guide to Northeastern Longhorned Beetles (Coleoptera: Cerambycidae). Champaign, Illinois: Illinois Natural History Survey, Manual 6. 174 pp.

Raid of the Slave-makers

The insect world is full of drama, one of the major attractions for entomologists and naturalists and wildlife photographers. Among the more rarely-witnessed phenomena are raids by slave-making ants in the genus Polyergus, known as "Amazon ants."

Raiding party of Polyergus montivagus

I have had the privilege of seeing three separate slave raids while living here in Colorado Springs, Colorado. All have taken place in late afternoon or early evening. The latest was on July 1 of this year when my wife and I were hiking a trail in Cheyenne Mountain State Park. She happened to pause at what I initially dismissed as yet another harvester ant trail, worker ants bustling about with grass seeds.

Amazon ant workers carrying pupae of their "slaves"

The ants were actually carrying the pupae of their host, ants in the genus Formica. We traced their apparent destination to a Formica colony, but I have learned that this makes sense. The adult slaves on the receiving end of this raid represent members of a colony taken over by an Amazon queen that managed to dupe the workers of an existing Formica colony before killing its queen. Those workers will now set about rearing these most recent arrivals, "adopted" larvae and pupae.

Those jaws are ready for battle

The anatomy of Amazon ant females is such that they are obligatory warriors. Their bodies are sleek and shiny, their slick exoskeletons deflecting the attempted bites of their victims. The jaws of Amazon ants are sickle-shaped and designed to do only one thing well: pierce the heads of the worker Formica ants. Amazons cannot feed themselves, let alone excavate a nest, so they must depend on existing subterranean nests of their Formica hosts. The Formica workers at the receiving end of the Polyergus raid were already enslaved!

The efficiency of a Polyergus slave raid is stunning. One wonders if the victimized colony, pilfered of most of its juvenile workers, ever recovers from such devastation. These ant pirates show now mercy, except for the kidnapped larval and pupa offspring, the majority of which they somehow manage to transport without injury. Not that some of these soft-bodied juveniles will not end up as food instead of slaves, mind you.

Worker Amazon, Polyergus mexicanus

How this specialized lifestyle came to be is open to conjecture, though a reasonable theory comes from studies of ant evolution and genetic relationships. There are slave-making species in the genus Formica, some of them obligatory slave-makers, and others facultative. Facultative social parasites are species capable of existing and prospering in the conventional sense, but are also opportunistic slave-makers. It is surmised that Polyergus Amazon ants evolved from the obligatory slave-making Formica species. Indeed, they appear closely related.

An average colony of Polyergus is between 300-500 workers, surrounded by many hundreds of their slaves. Such a colony can include a mix of different Formica species. Raids on Formica colonies are frequent, and so Amazons need a robust population of host colonies that they can draw from.

Carting off the "booty"

The genus Polyergus is holarctic (found throughout the northern hemisphere), but reaches its zenith of diversity in the United States. There are fourteen species in the world, eleven of which are found only in the U.S. Polyergus mexicanus is likely the one I have been seeing here along the Front Range of the Rockies. Ant diversity in general is surprisingly great here in Colorado, so it is easy to assume you are only seeing a few species when in fact there are several.

Keep an eye out for interesting ant behaviors, and try and document them as best as you can. You could shed new light on our collective knowledge of these amazing social insects.

Sources: Anonymous. 2017. "Polyergus,"Antwiki
Holldobler, Bert and E.O. Wilson. 1990. The Ants. Cambridge, Massachusetts: The Belknap Press of Harvard University Press.732 pp.

Predator and Prey: Ants versus Termites

My neighborhood walk in Colorado Springs the other day, March 25, was like strolling through living confetti at some points. All the local termite colonies were launching swarms of winged males and potential queens (alates as scientists call them). The frail creatures were not ignored by other animals, either, especially ants. Closer inspection of the swarms revealed three species of ants preying on them.

Alate subterranean termites (Reticulitermes sp.) swarming

Termite swarms are not an indication of the impending collapse of your home or any other wooden structure. Yet, that is the first thought that enters the mind of the average person witnessing the spectacle. Such is the power of advertising for pest control companies. Now, a termite swarm inside your home should probably be cause for alarm. Outdoors, subterranean termites like these Reticulitermes sp. are vital to the recycling of decaying wood. They nest in the soil, as their common name suggests, and forage for wood and other dry cellulose in contact with the soil.

The synchronous nature of termite swarms is a marvel. All colonies in a given area need to liberate their reproductive castes at the same time in order to prevent inbreeding, but I have no idea how they "decide" when to do this. The day before we had snow and high winds. The alates issue from the tiniest of cracks in the soil, like toothpaste from the tube, the better to avoid easy detection. Eventually, enough of the insects appear that their gauzy wings reflect the sun and give away their presence. Soldier termites, and workers, too, escort them out and see them off.

Alate termites with workers and soldier (center) escorts

Hundreds, if not thousands, of winged termites begin filling the air. Few will survive the alert eyes and hungry mouths of birds, lizards, and other predators. The early season timing of swarms may in fact be tuned to precede the emergence of reptiles and the arrival of migrant birds. Ants, on the other hand, are already on the prowl.

Worker Formica sp. ant carrying termite prey

Both ants and termites are social insects, so it is fitting they would be deadly enemies and, one would think, well-matched foes. Watching one swarm happen on the edge of a driveway, I began noticing the appearance of worker ants, Formica sp., crossing the driveway. Eventually I saw one toting a winged termite back to the nest. The ant's nest. More ants followed suit.

Pavement ants (Tetramorium caespitum) killing alate termite (bottom) and worker termite (top)

Turning my attention back to where the termites were emerging, I noticed something even more frightening. Tiny "pavement ants," Tetramorium sp., were killing both alates and worker termites right at the termite nest opening. Whereas Formica ants are a bit larger than the termites, the pavement ants were smaller than their prey. How they avoided the menacing jaws of the soldier termites confounds me.

Formica ants near the entrance to their nest, with prey

Just up the street I noticed heavy ant activity originating at the base of a brick-and-mortar mailbox pillar. These were Formica pallidefulva ants, but appeared larger than the other ones I saw previously. It soon became apparent that they were also taking part in the Great Termite Massacre of 2017. Most of them were carrying wingless alates, though.

Ants (Formica pallidefulva) with termite prey

Alate termites, once paired, shed their wings easily. Both pairs of wings have a weak spot that allows the termites to break them off so they can quickly seek cover. The male ("king") termite follows on the heels of his mate (queen) as they form a two-car train in search of a potential nest site. They must do so quickly if they are to avoid the marauding ants.

Dealate queen with her mate trailing her in a "train"

Whether the honey pot ants were taking dealate (wingless reproductives) termites, or just seizing winged individuals and breaking off their wings, remains a mystery. They are certainly easier to transport without those cumbersome wings.

Worker Formica sp. ant carrying termite prey

As I turned the corner to go home, I caught sight of yet another ant, possibly Formica podzolica. It, too, was carrying a defeated termite. The ant seemed at least somewhat disoriented, and I eventually lost track of it in the thick grass at the edge of the curb.

So, termites are both integral to keeping soils fertile with their decomposition activities, and also a bounty for many other organisms that depend on them for food when other insect life is less plentiful. Ants are the lion kings and wolf packs of the macroscopic landscape, keeping termites and other insects from overrunning the planet. The ants are not immune, though, and in my next post we see them on the other end of the predator-prey equation.

Note: Special thanks to James C. Trager for identification of the ant species.

Flying Ants

I recently came across a male specimen of the "legionary ant" Neivamyrmex nigriscens while walking our dog in our Colorado Springs neighborhood. It was instantly recognizable to me, but it got me thinking about how most people would be hard-pressed to know what it was. Unfortunately, there are very few references for the identification of winged ants. This is a shame because it is often the "alates," males and winged queens, that are most obvious to the public.

Male Legionary Ant, Neivamyrmex nigrescens, Colorado

Indeed, worker legionary ants are mostly subterranean and nocturnal in their habits (nomadic, raid the nests of other ants to prey on the larvae and pupae), so hardly ever observed by the average person. Meanwhile, a homeowner may not notice he or she has an ant "problem" until the colony swarms, liberating a cloud of alates.

Alate queen carpenter ant, Camponotus sp., Colorado

Alates are typically larger than the worker caste of wingless, sterile females, so are more noticeable for that reason as well. In many cases, the winged reproductives resemble the workers in general appearance, but this is not always the case. The thorax of winged ants is frequently greatly expanded to accommodate the muscles that operate the wings, giving males in particular a distinctive "hump-backed" appearance.

Male carpenter ant, Camponotus sp., Massachusetts

Swarms are usually seasonal, and triggered by changes in day length, relative humidity, and air pressure, especially in the arid southwest U.S. where the onset of the monsoon rainy season sparks many ant species to swarm. These emergences can be spectacular events. Worker ants open new exits from nests in the soil, and scour the immediate vicinity to rout any potential predators and parasites.

Red Imported Fire Ant swarm, Solenopsis invicta, Georgia

Many swarm events take place in late afternoon, at dusk, after dark, or at dawn. Winged ants may be attracted to outdoor lights, which can lead to the assumption that the ants came from the house or building when that is not necessarily the case.

Colonies of a given species in a localized area swarm simultaneously such that members of different colonies can find each other and increase genetic diversity while decreasing the potential for inbreeding. Winds, and the insect's own muscle power, can take the ants far from their colony of origin.

Rough Harvester Ant swarm, Pogonomyrmex rugosus, Colorado

How do you know whether it is a winged ant or a winged termite? Please read my post on termite swarms for a concise explanation, and images of winged termites. Below is an image of a winged termite to compare to the ants illustrated here.

Alate dampwood termite, Zootermopsis laticeps, Arizona

What about wasp versus ant? That is a more problematic distinction, but most ants have distinctly "elbowed" antennae, whereas wasps often do not; or at least the first segment of the antenna is not as long as it is in ants. There are exceptions, of course, like the male Neivamyrmex ant shown at the top of this post that has no obvious elbow in the antennae.

Male Pavement Ant, Tetramorium sp., Colorado

Fortunately, I am not the only one who recognizes the need to pay more attention to alate ants in terms of research and public awareness. Laurel Hansen and Art Antonelli include a key to alates in their publication, listed below. Brendon Boudinot, in a guest post for Alex Wild's Myrmecos blog, extols the virtues of studying male ants for a clearer understanding of the phylogeny of the family Formicidae.

Alate queen thatching ant, Formica sp., Colorado

Sources: Boudinot, Brendon. 2013. "Male Ants Demystified," Myrmecos.Hansen, Laurel, and Art Antonelli. 2011. Identification and Habits of Key Ant Pests in the Pacific Northwest. A Pacific Northwest Extension Publication 624. Pullman, WA: Washington State University. 14 pp.
Houseman, Richard M. 2008. "Ants." University of Missouri Extension.