IMG_7733.JPG

cODING bEHAVIOR

Taking Data

All of the behavioral data John and the squad take in the field goes onto an interaction datasheet, where the time, location, individuals, and specific interaction are recorded on a single line. Prairie dogs are fast, and especially during the mating season when activity reaches its peak, pages and pages of these interactions may be taken in a single day, one after another after another in such quick succession our hands begin to ache. Looking at a datasheet, the writing may look like confusing coded alphanumerics, but like reading music all the lines together produce a story of what happened that day.

Getting to know and understand the prairie dogs is important. While simply writing down what you see happening may seem simple, understanding exactly what to write down is key. For example, a kiss can be friendly or hostile; or a kiss may actually be the start of a territorial dispute; one interaction following another might be directly related and such series of cascading reactions are indicated with arrows; et cetera. A particularly diligent field assistant will get to know the dogs well enough to anticipate certain behaviors from certain individuals, though "predictable" is not a word we would use to describe prairie dogs...more "consistent."

Key Interactions

Kiss (K)

To call the famous prairie dog mouth-to-mouth posture a kiss is a slight anthropomorphism, but the designation is simple and descriptive of what the behavior looks like. When two prairie dogs tilt their heads and touch mouths, usually touching tongues, it is coded as a KISS (K). While it is not fully understood what is being exchanged when two prairie dogs kiss (i.e. any sounds they are making that we can't hear) we know there are several purposes behind the famous kiss behavior in reinforcing social bonds: saying hello, identifying each other, and posturing during a dispute. In this way, kisses can be either friendly or hostile.

A friendly kiss; an adult prairie dog kisses a baby to say hello and determine if the offspring is hers. ©MRR 2017

Parting after a hostile kiss. ©John Hoogland 2010

During a friendly kiss (K), two prairie dogs are greeting and identifying each other - such as when a mother approaches a wandering litter and kisses every baby to determine if they are all hers; or when two females meet at a burrow entrance during their daily foraging. During a HOSTILE KISS (HK), one or both of the individuals involved will jerk or jump away from the kiss, may respond with an agitated squeak, or will slap the other prairie dog (usually the slapper is a female telling a male to go away). The most common hostile kiss is seen between males during a territorial dispute - in which case rather than coding the behavior HK, we will code the interaction TD for Territorial Dispute (more on that farther below).

All prairie dogs - male, female, adult, yearling, and juvenile - engage in kiss behavior.

 
 

Sniff (S)

The sniff is strictly a mating season behavior. During the mating season, males spend a considerable amount of time crawling up behind females to smell between their legs. They are looking for signs of estrus, eager to catch the female on her single day of receptivity. This behavior is coded SNIFF (S). Often the female will jump away from a sniffing male, but sometimes (especially when she is indeed in estrus) she will allow him to sniff for several seconds before walking off. Sniff behavior ends after all females have mated.

 

Male 2 comes up behind female Wide Apart to sniff between her legs. ©MRR 2017

 

Chase (C), HERD (h), Chase Herd (C,H)

Chasing is a hostile interaction among prairie dogs (unless they are playing, which is coded differently). A chase may occur as an isolated interaction and be coded simply as CHASE (C), or as part of a territorial dispute. Most often, the aggressor will chase the receiver into a burrow to establish dominance, and will proceed to stand for any number of seconds at the mouth of the burrow to ensure the receiver does not yet emerge, which is a HERD (H) behavior. When combined, the entire interaction is coded together as a CHASE HERD (C,H).

Interment Behavior (IB)

Occasionally after a Chase Herd, the aggressor will proceed to kick dirt into the mouth of the burrow wherein he's chased the receiver of his aggression. The activity will continue until the burrow entrance is buried or half-buried with dirt, and the receiver is "trapped" within - these plugs are easily broken through afterward. We code this interaction as INTERMENT BEHAVIOR (IB), and understand as a dominance-establishing activity.

A chase. ©John Hoogland 2009

Two feuding males nearly make contact during a high-speed chase. ©MRR 2017

 
 

Run away (RA), Jump away (JA)

Often the response to a hostile interaction, whether it be a kiss or just an approach, is for the receiver to run away from the aggressor, and we code this as a RUN AWAY (RA). Often the interaction sheet will read "C, RA", which happens when the aggressor bluffs a chase but does not carry through when the receiver runs off. Sometimes the response is a startled reaction and the receiver will actually jump in the air out of the way, and we code this a JUMP AWAY (JA). Jump Aways are most common during the mating season when the males are crawling up behind females to sniff between their legs as they check for signs of estrus. They also happen when a prairie dog feels cornered, as seen in the video below.

 
 

Territorial Dispute (TD)

Hostile kissing, chasing, posturing, and contact fighting are all elements of a TERRITORIAL DISPUTE (TD). Commonly seen between males (especially during the mating season) but also seen between females, territorial disputes are sometimes brief but usually drawn out arguments over territory and resources. No two TDs are the same; they could last a few seconds or several minutes; they could involve posturing without kissing, or drawn-out kissing; simply chasing and posturing, or escalating into a contact fight; et cetera.

 

A hostile kiss between two males during a territorial dispute. ©MRR 2017

 

While most TDs occur between males, females will often getting into equally violent disputes during the nesting season, when they are arguing over potential nursery burrows and the territory around those burrows. This hostility among females kicks in after each female has already mated and maternal behavior begins. A territorial dispute between a male and a female is rare.

 
 

Fight (F)

It is not very common for two prairie dogs to approach each other and break immediately into a contact fight; rather, in the midst of a particularly contentious territorial dispute, if a lunge is made, or a chase rolls over, and contact is made, the two disputers will break into a contact fight, coded simply as a FIGHT (F). Contact fights are brief, lasting anywhere from 2 to 5 seconds, rarely up to 10 seconds at the extreme. During this brief flurry, blows are delivered via biting, kicking, and clawing, as the two prairie dogs roll into a ball of flying fur, often going airborne with their momentum.

Two males in a contact fight. ©MRR 2017

A territorial dispute escalates into a contact fight. ©John Hoogland 2009

Contact fights between males are common during the mating season, where the risk of injury might be worth the chance to establish dominance and gain access to more receptive females. On the other end, once they have copulated and are past their day of estrus, females become more aggressive (as mentioned above), and contact fights over potential nursery burrows may happen. Establishing an optimally-located and suitable nursery burrow is key for pregnant females, and losing disputes can have grave consequences for the fitness of the mother and the survival of her upcoming litter.

Both Down (BD)

With a small number of exceptions every year, most copulations during the mating season occur underground. John and the squad use aboveground contextual cues to determine whether a copulation is going to occur, is likely occurring, or has occurred (see more in ABOVEGROUND CUES). The best indicator that two prairie dogs are going to mate on a given day is if both prairie dogs submerge underground for longer than 15 seconds together (typically after the male has chased the female, or sometimes even when the female has solicited him to follow). We code this time-sensitive submergence as a BOTH DOWN (BD). BDs can last anywhere from 15 seconds to several minutes, and there will usually be several BDs through a span of hours before the critical copulation occurs (during which both prairie dogs disappear underground for anywhere from several minutes to a couple of hours).

 
 

A BD is over when the first of the pair emerges back aboveground (typically the male will emerge first, and the female will come up a few seconds to a couple of minutes later). Though we always record the time of interactions on the datasheet, we are especially vigilant in recording seconds, minutes, or hours when it comes to coding BDs, as the amount of time spent underground will tell us if the critical copulation has occurred or is yet to occur. Visit our MATING BEHAVIOR GALLERY for photos of mating behavior.

LickS Penis (LP), LickS Vulva (LV)

A tell-tale sign of whether a copulation has occurred between two prairie dogs is when one or both emerge from their time underground and lick their genitals. Though this gesture can look similar to regular grooming, to lick between their legs both the male and the female have to bend over more dramatically, sometimes even falling over as they reach. When we witness this, we code it LICKS PENIS (LP) or LICKS VULVA (LV). Not every prairie dog will lick after copulating, but if it happens we can use it as part of the series of aboveground signs that a copulation has occurred.

Dust Bath (DB)

Along with LPs and LVs (or even absent of them), another sign that a copulation has occurred is if the male and/or female rolls in the dirt after emerging from the mating burrow. We code this behavior as a DUST BATH (DB). As with the other indicative behaviors, not every prairie dog will roll. It should be noted that dust baths can also occur after a contact fight, and are especially looked for after an infanticide (wherein the killer will roll in the dirt to clean him- or herself of blood).

 

Male 36 leans over to lick his penis after copulating. ©MRR 2017

 

LickS Front Paws (LFP)

Prairie dogs will lick their front paws after vigorously working their burrow mounds or their tunnels (to clear dirt from their nails), but we only take notice and record this behavior as LICKS FRONT PAWS (LFP) after a suspected infanticide event. If a prairie dog enters a burrow that is not her own, and in which we know babies are living, we watch and count the seconds or minutes the adult remains underground. If the time exceeds 20 seconds, we begin to suspect an infanticide may be occurring. If the prairie dogs emerges and licks her front paws, we are further suspicious. In the video below the LFP behavior is displayed, but in this instance this prairie dog was only working on her own tunnels. Context is critical to determining what an LFP is indicating.

 
 

Rubs Face in Dirt (RFD)

Another tell-tale sign of an infanticide is when the killer rubs his/her face in the dirt to remove blood. Though difficult to differentiate from a dust bath, when a prairie dog has committed an infanticide (or suspected infanticide) and rubs their face vigorously in the dirt, we code it as RUBS FACE IN DIRT (RFD). Like the licking of the paws, we only really take notice of RFDs when in the context of an infanticide. Prairie dogs may rub their faces in the dirt during dust baths as well, but when in the context of other indicators of infanticide, the RFD helps us to put together the bigger picture.

Additional Data

Takes Nesting Material

Every interaction datasheet has a blank right-hand column for recording nesting material (NM) behavior. Prairie dogs will gather nesting material (dried grass strands gathered like straw) all year-round (especially on inclement weather evenings), but the activity increases after the mating season when females are preparing their nursery burrows. We record how often a prairie dog brings nesting material into their burrow, as part of our behavioral analysis. One question this data hopes to answer, for example, is whether NM gathering affects offspring survival (does a mother who spends more time gathering nesting material have a higher chance of seeing her offspring to healthy emergence in the summer?). Sometimes a prairie dog will pilfer nesting material from one burrow and take that material to their own, and we record such pilfering behavior as well (though it is not very common).

Using her front paws to adjust stray strands of grass in her mouth. ©MRR 2017

A mother brings nesting material to her nursery burrow. ©John Hoogland 2007

 
 

Works Mound

On the bottom right of our datasheets is a blank column for recording mound work behavior. Prairie dogs will frequently work their mounds to clear rocks from the burrow entrance and to smooth the dirt, keeping ingress and egress easy and unobstructed. Among black-tailed prairie dogs especially, mound work also means building higher ground around the burrow entrance, called a rim crater, for better vigilance. The white-tail species like the Gunnison's prairie dog do not tend to build their mounds as high. Regardless, the prairie dog mound is one of the most prominent sights in the western grassland ecosystem, dotting the prairie as evidence of a healthy working landscape.

 

RACR works her mound, removing rocks and smoothing dirt. ©MRR 2017

 
 

©MRR 2016

 

Alarm Calling

At the bottom of every interaction datasheet is blank space for recording prairie dogs that "give alarm calls for no apparent reason." Often the perceived threat an alarm-calling prairie dog is responding to is not apparent to the researchers, no matter how much scanning of the colony we do. Prairie dogs can see, hear, and smell predators that humans cannot. If an alarm call is being sounded and the threat is not clear to us, we record the call. If a predator becomes visible, we will record the predator's presence and location on one of the lines of the interaction datasheet.

Targeted alarm call data is taken in the summer after the offspring of the year have emerged from their nursery burrows, and as the last task for John and the squad after all the new offspring have been marked and before the site is pulled for the season. We take this data using predator-pull experiments, which you can read more about in THE PREY ANIMAL. The datasheet is pictured below.

 

A predator-pull alarm-call experiment datasheet. ©MRR 2017

 

Censuses

Though not behavioral data, we take censuses frequently to keep track of the presence or absence of particular prairie dogs. These include Missing Male Lists (MMLs) which, despite the name, also contain females. The MML is used during the mating season and lists breeding males and any female who has not yet gone into estrus. Every 20 minutes, the researcher will search for every individual on the list, and if an individual is missing for more than two checks, it's possible he or she is copulating underground.

We also take censuses on a separate multi-use datasheet (pictured below) where throughout the year, we record times and locations of wakeups and beds for prairie dogs in every clan; and additionally, after emergence of new offspring in June, the same datasheet is used to count and record babies per litter to determine if any juveniles have gone missing from one day to the next.

 

A Wakeup/Bed and Baby Census datasheet. ©MRR 2016

 

It takes a bit of time and concentration for a new research assistant to learn how to take data efficiently, especially for tricky behaviors like Territorial Disputes and Both Downs. As the field season progresses, specific data becomes more or less common, and the note-taker is constantly learning new behaviors and new codes. By using an established glossary of behaviors and their codes, John has been able to guide his assistants over the decades toward taking consistent, significant data with little error. If you haven't already, be sure to visit the PURPOSE & METHODS page to learn more about the daily life of John Hoogland and the Prairie Dog Squad.