March weather had the rattlesnakes along the American River Parkway confused! As I have mentioned before, the rattlesnakes usually emerge and begin to bask when the average temperature at ground level remains at or above about 50F. They are usually deep enough underground during the winter to escape daily high and low fluctuations, so it is the average that eventually penetrates to them and signals that winter is likely over. But not this year.
Most telemetered rattlesnakes emerged in March but remained at their hibernation sites, basking on sunny days as they usually do before leaving their winter shelters. But rather than getting drier and warmer, the weather throughout April was a constant mixture of rain and cloudy cool days and nights. Of the few rattlesnakes that left their winter hibernacula, two returned while others became inactive at other sites. The rattlesnakes do just fine in the rain. In fact, they drink rain and dew when it’s available. And cloudy weather is not a deterrent to activity if it is accompanied by warm temperatures. But nighttime air temperatures below 50F followed by overcast cool days are not good for activity in animals like reptiles that depend on the environment for their body heat.
So, in late March and throughout April, the few stretches of several sunny days with relatively warm nights in between produced some sporadic rattlesnake activity. But it was not until the end of April that more consistent sun and heat produced vigorous widespread rattlesnake activity.
As expected, when we had activity, we observed lots of courtship. This is, after all, the peak of courtship activity, followed by almost no reproductive activity from June to mid-August before an untick in courtship again in the fall.
But by far the most interesting discovery this spring was by one of my UC California Naturalist students who found a small male rattlesnake with no rattle! (See photos below) Now I hesitate to mention this, for fear of starting a panic about rattleless rattlesnakes. Please remember several things:
Rattlesnakes missing their entire rattle are extraordinarily rare. So rare, in fact, that they usually rate a note in a biological journal. This one will!
No rattlesnake has a long pointed tail like our many harmless snakes.
Rattles can be missing due to either injury or genetic mutation.
This little snake appeared to be recently missing the end of his tail and had other serious but healing injuries. Apparently a close call with a predator!
Identify rattlesnakes by looking at the tail! Yes, rattlesnakes have elliptical pupils and heat-sensitive facial pits but some harmless snakes have elliptical pupils and both these characters are too small to be clearly seen from a safe distance (two-times the length of the snake). Some harmless snakes also have rather triangular heads, especially when they have been frightened and are behaving defensively. But no rattlesnake has a long pointed tail like our harmless snakes and all of California’s dangerous snakes are rattlesnakes.
Check out the following images…
To be sure, there are dangerous snakes in other parts of the United States that have long tails without rattles (cottonmouths, copperheads and coral snakes, in southern and eastern states). But in California and other northwestern states, the only dangerous snakes are rattlesnakes.
Look at the tail to identify rattlesnakes!
And, yes, Male 87 was processed and released, just like all the others. While he has no paint in his rattle, he won’t be hard to recognize!
It took only a few days of moderately warmer weather to stir our sleeping rattlesnakes. I haven’t downloaded my temperature logger recently but it has been easy to notice that it’s in the 50s instead of the 30s the past few mornings, while daytime highs have been in the high 60s.
And today, we found some rattlesnakes stirring. Until now, the body temps of all of the hibernating rattlesnakes have been in the single digits Celsius (less than 50F). Earlier today, ten of twelve telemetered rattlesnakes were in double digits despite a full thick overcast. Male 37 – still inaccessible on the bluff – produced a calculated body temperature of 21C (70F). (The bluff is always warmer because it is steep and faces south, so solar energy is more concentrated per unit of surface area.) Male 37 is clearly no longer deep underground.
Next was Male 46 at 16C (61F) and, sure enough, he was peering out from under the log where he spent the winter.
When we arrived at Female 41’s radio signal, we found a rattlesnake laying out in the grass and assumed it was her. The rattle was not clearly visible at the time but later examination of photographs disclosed white/green paint in the rattle. Female 41 is white/blue and the rattlesnake we observed turned out to be Male 49, marked and released without a transmitter exactly two years ago today, on 10 March 2015! Female 41 was very close but out of sight.
We also found Male 40 laying out in the same spot as last week (see the blog post on 6 March), plus we found a new animal laying in the grass next to another large log where two telemetered adults have been napping since last fall. He (or she) was captured and, while too small for a transmitter, will be measured, marked, and will donate a blood sample before being released as CROR81.
Well, it’s March and many recent days have been sunny and quite warm. Even though daytime highs have been listed in the 50s, that’s measured in shade and the temperature in the sun near the ground is always considerably warmer. So why were rattlesnakes basking in good numbers by late February last year but not yet this year?
There are two parts to the answer. First, most of the snakes spend the winter deep enough underground that they are insulated from daily temperature fluctuations. In other words, they do not feel daytime highs or nightime lows. Deep in their winter shelters, the temperature creeps up or down gradually in response to the mean (average) outside temperature. The second part of the answer is that natural selection tends to remove animals from the gene pool that don’t spend the winter deep enough and/or that venture out too early and get caught in freezing weather. The animals that select optimal winter shelters tend to live longer, and thus produce more offspring carrying the genes for those successful behaviors. (For those of you interested in the evidence for genetic influence on behavior, Time Love and Memory (2000) by Pulitzer Prize winning author Jonathan Weiner is an entertaining and astonishing summary, written in easy-to-understand language but with references to the scholarly research.)
So how do we know the temperature in the underground shelters?
Besides having recorded hourly temperatures for nearly four years with a data logger in a burrow 1 meter (39″) below ground in the Mohave Desert during a previous study, we can routinely calculate body temperatures of the hibernating snakes from the pulse interval of their transmitters during the winter.
The transmitters come from the manufacturer (Holohil Systems Ltd.; www.holohil.com) with a calibration graph showing the correlation between pulse rate and temperature for each transmitter (they vary slightly).
I routinely check the pulse rate in a water bath at room temperature before implanting a transmitter and adjust the mathematical conversion, if it differs slightly from the factory data. Then it is a simple matter in the field to time the pulse interval with a stopwatch and convert that measurement to temperature later. And since the transmitter is implanted inside the snake’s abdominal cavity, transmitter temperature equals core body temperature.
I also record hourly shade air temperatures with a data logger in the nature preserve at Effie Yeaw Nature Center. This gives me a standard local reference against which to compare rattlesnake behavior and look for correlations with weather conditions.
I can then compare rattlesnake behavior, body temperatures and air temperatures. This is especially interesting when one year is compared to another.
You can see about six weeks’ data from this year in the chart above. The fluctuations in mean (average) air temperature (open circles) reflect the daily changes based on cloudy verses sunny days and cold air masses moving through verses warmer southern air (the same factors discussed every day by meteorologists). The five filled circles are the mean body temperatures of the twelve telemetered rattlesnakes on the days that I recorded their data. There are more air temperature records because the data logger records automatically around the clock while rattlesnake body temperatures are only recorded when I visit the study site (which I do less frequently when the animals are inactive).
Now take a look at the corresponding data from last year:
Checking my behavioral observations from last year, I find a few rattlesnakes were starting to bask occasionally by the end of January and most were basking regularly on sunny days by the end of February. Regular basking continued through mid-March. Then, suddenly, between 17 and 19 March, most of the rattlesnakes left their hibernation shelters, as if someone had given them the “all clear” signal. This was multiple rattlesnakes leaving multiple shelters at the same time – pretty amazing! By the way, you may also notice that I visited and recorded data more frequently in February last year. That’s because there were rattlesnakes to see and behavior to record (besides the science; it’s just more fun when there are rattlesnakes to see!).
If I combine the raw data from 2016 and 2017 for comparison, the chart gets pretty messy and trends are hard to make out. But we can create mathematical models for those same data that present a much clearer picture.
You can see that mean February air temperatures started out much lower this year compared to 2016. And, in 2016, with mean air temperatures exceeding 50F by early February and staying there, the rattlesnakes began to bask and their body temps took off as a result. This year, air temps started nearly ten degrees lower, rose briefly, but then took a nosedive in late February, keeping the rattlesnakes cool and underground so far.
Of course, some individuals do not behave in the same manner as most others. I have heard some reports of a few rattlesnakes being sighted in the past few weeks and one dog reportedly bitten already. While none of the telemetered rattlesnakes at Effie Yeaw have emerged from hibernation yet, one rattlesnake without a radio has been basking early – just as he has done each of the past two years. My old skinny Male 40 has been frequently laying in the sun at the entrance to his winter shelter for more than a week.
As you may recall, Male 40 was implanted with a transmitter in 2014 but I removed his second radio last spring and released him without one because he was not producing useful behavioral data. It was not that he was just behaving differently than the others (after all, unusual behavior is interesting and valuable to record), he has been chronically and significantly underweight, appears to be quite old, moves very little during the summer, and has never been found courting a female. Given his poor body condition, I was afraid that he would die during hibernation and his transmitter would not be retrievable. But, even without a radio, he continues to be sighted regularly and has now made it through another winter.
But most others seem to be waiting for some sustained warmer weather before venturing out.
Well, we are entering that time of the season when we can expect to find baby rattlesnakes any day now. Of course, the youngsters will not be roaming around where they might encounter people for about 10-14 days after birth…so we usually don’t encounter them on trails and in yards until early September.
But our six telemetered females are displaying a variety of behaviors indicating that all may not be about to produce babies. Six weeks ago I reported that Females 39, 41, and 53 were all pregnant and in the gestation shelters they have used repeatedly to thermoregulate before giving birth in previous years. Furthermore, I could feel six fetuses in Female 80’s belly when I implanted her transmitter in June.
Since that mid-July report, Females 39 and 53 have behaved predictably for expecting moms, remaining in their gestation shelter, and both appear to be about to produce kids. In the photos below, note the abdominal girth of these females, with scales pulled widely apart.
This will be the third consecutive year that Female 39 has produced offspring and the second for Female 53, so far as we know. We were not monitoring them previously. Between the first of July and yesterday, 23 August, the average body temps for Females 39 and 53 have been 28C (82F) and 29C (84F), respectively.
Six weeks ago, Female 41 had just returned to the gestation shelter she used the previous two years, which led me to believe she was likely pregnant again. However, after staying only three weeks, she left and has been spending her days mostly out of sight in various ground squirrel burrows during August. I have not been able to get a good look at her in recent weeks but her behavior and average body temperature of 26C (79F) suggest that she may not reproduce this year. Although Female 41 produced kids in both 2014 and 2015, I want to remind readers that skipping a year or two between broods is far more common than annual reproduction for temperate-latitude pitvipers.
Our remaining three telemetered females are new to the study this year, so I have no data from previous years. Female 66 has behaved quite normally for a non-reproductive year: hunting continuously throughout the spring and summer. Her July-August average body temp has been 22C (72F).
Female 75, however, appears ready to produce babies, although she has been moving back and forth frequently between two ground squirrel burrows 17 meters (56 feet) apart. Her average body temperature during July and August has been 27C (81F).
Finally, there is Female 80. When I captured her and implanted her transmitter in June, I could clearly feel six fetuses lined up in her abdomen. Shortly after I released her at her capture site near the base of the bluff, she climbed up the hillside and has remained near the top ever since. On three occasions, I have climbed that steep, loose, and treacherous slope but have failed to locate her. Her radio signal seems to emanate from thick ornamental ivy growing down from a residential backyard under fig and valley oak trees. She is also just above an underground wasp nest. Just to be clear, I am far more comfortable with rattlesnakes than a nest of wasps – especially where I cannot easily run away! Since I have no previous data from her, she could be in her usual gestation shelter. However, her July-August body temperatures are not consistent with a gestating female, with an average of 23C (73F). I’m not sure how this is going to play out…
Just for comparison, the seven telemetered males have been moving around a lot, hunting and occasionally hanging out with the pregnant girls for a day or two at a time. Their average July-August body temps range from 17C (63F) to 26C (79F), with the average between the boys of 23C (73F). Note how closely these data match non-reproductive Female 66’s average body temp of 22C (72F) during the same period.
That’s it for now. Next post will almost certainly be (cute!?!?) baby pics!
We currently have transmitters implanted in six female rattlesnakes. We have been tracking Female 39 since 2014 and Females 41 and 53 since 2015 and all are currently pregnant and in their gestation shelters where they will likely thermoregulate for six weeks or more. Furthermore, they have all produced a brood during each season we have been following them, so this will be the third year in a row for Female 39 and the second in a row for 41 and 53. And we don’t know how many consecutive years they might have reproduced before that! Each has returned to the same gestation refuge each year, although 41 uses a different location than 39 and 53, who gestate together. We have also found pregnant unmarked females in both places in past seasons with these girls.
That’s not all. We have three new females this season, numbers 66, 75 and 80. These snakes do not appear to have settled into gestation shelters yet (and I don’t know where they were in previous years) but I just implanted a transmitter in Female 80 a few weeks ago and could feel six fetuses in her belly…it felt like she had swallowed six soft ping pong balls! I’m not sure about the reproductive status of 66 and 75, since their surgeries were earlier in the year and both had so much material in their intestines that it made identifying small embryos with confidence difficult.
Another opportunity to assess the health and growth of the rattlesnakes at Effie Yeaw Nature Center occurred day-before-yesterday (13 July) when Kelly came across an adult rattlesnake at the end of the Visitor Center building early in the morning. Per protocol, she expertly maneuvered it into the capture bag and deposited it in the holding barrel for me. It turned out to be Male 52, a rattlesnake previously captured, processed, and released without a transmitter early in May 2015. At that time, he was 30.7 inches in total length and weighed 11.6 ounces. He now measures 33.5 inches and weighs 18.9 ounces. While increasing 9% in length and 63% in mass in 14 months, he has shed three times (see photo below).
The constant growth of all the rattlesnakes being sampled and the annual reproduction of many of the females attests to the health of not just the rattlesnake population but the overall small animal community in the riparian habitat at Effie Yeaw Nature Center. While the region is undeniably in a severe long-term drought, enough local rainfall has occurred to keep the annual plants, shrubs, some trees, and the food web they support healthy.
Finally, I want to share with you a little bit about a presentation I made at the annual Joint Meeting of Ichthyologists and Herpetologists last week in New Orleans. I have reproduced the published abstract (summary) of my talk below. Please excuse the Latin names. Crotalus scutulatus is the scientific term for the Mohave rattlesnake and Crotalus oreganus is our own northern Pacific rattlesnake. While recent drought has not yet affected the rattlesnakes or their prey in the areas of northern California I have sampled, 2002 was a rainless year in the Mohave Desert, with no plant growth and a dramatic reduction in the availability of kangaroo rats and other small mammals that make up the great majority of the rattlesnakes’ diet there. During 2002, Mohave rattlesnakes changed their behavior very significantly, staying tightly coiled and avoiding wind and sun while moving very little and not courting or mating.
The take-home message I delivered in New Orleans was that (1) these animals are used to hot dry summers and get most of their water from their prey; (2) regional drought does not necessarily equal local drought; (3) rattlesnake behavior is probably not affected by drought until prey availability is affected; (4) water-stressed rattlesnakes minimize exposed skin by remaining coiled most of the time; (5) when water-stressed, they don’t move more, they move a lot less than usual; and (6) there is no evidence that they leave their normal home range during a drought.
In other words, there is zero evidence to support the frequent news media claims that drought drives rattlesnakes into yards.
Despite persistent cool mornings and alternating sunny and cloudy days, nine of eleven telemetered rattlesnakes have left their winter shelters and the remaining two have been basking regularly and will undoubtedly be on the move soon. We have also been finding plenty of new rattlesnakes basking and now have 27 animals processed and marked with colored paint in the rattles, in addition to the eleven with radios.
As I have already described, Female 41 was one of the first to leave her hibernation site and was soon joined by Male 49, a snake marked last season but not telemetered. Male 49 and Female 41 remained in close proximity at the same location from 16 February until 20 March and, although they were occasionally coiled in contact with one another, I never saw the male courting her. Male 49 followed when Female 41 moved more than 50 meters between 20 and 24 March but there has still been no witnessed courtship. However, the grass is very thick at their new location, making her hard to see and him impossible to spot if he’s not with her (and I don’t want to step on him… they’re very fragile!).
On 22 March, I came across Female 55, another animal processed last year and released without a transmitter, basking alone at the edge of a large log. But when I returned on 24 March, she had company: an unmarked (no paint in the rattle) male on top of her, jerking, chin-rubbing, and tongue-flicking – typical courtship behavior (you can view a 2015 clip of Female 41 with another male here).
We can expect the females to hunt for the next two-to-three months, after which pregnant females will retreat to their favorite gestation shelters to thermoregulate until their kids are born around the first of September. Non-pregnant females will continue to hunt through the summer. Males will spend most of their time looking for receptive females until late May/early June, after which they, too, will hunt full-time for voles and ground squirrel pups until courtship resumes in late summer.
After a couple of false starts during warm periods in mid and late February, a new series of cool storms kept the Effie Yeaw Nature Center rattlesnakes immobile through the first half of March. On days with a little sunshine, both telemetered and other rattlesnakes were frequently found basking at the entrance to their shelters but they did not venture away from them. Natural selection tends to remove animals from the gene pool that wander too early and get caught in a late-season freeze. As if to demonstrate that it was cool temperature and not the rain that was keeping them inactive, several rattlesnakes were found lying out in the rain on days when the temperature was in the high 60s, despite the clouds.
While the rattlesnakes continued to bask on sunny days, none of the telemetered animals moved between 29 February and 17 March. Then today, I discovered that 7 of the 9 telemetered rattlesnakes had left their shelters and moved significant distances in the past two days.
I haven’t seen courtship yet. Female 41 and Male 49 hung out together for a month (16 February through 15 March) but they were only rarely touching one another and were usually coiled a couple of feet apart with no courtship observed. She is now hunting several dozen yards away and Male 49, without a transmitter, has disappeared.
The important message, however, is that rattlesnakes are on the move and encounters between people and rattlesnakes will increase immediately. Unfortunately, I have already seen photos of a rattlesnake bite from a few days ago in the foothills. Watching carefully where you put your hands and unprotected feet and leaving snakes alone when you encounter them would prevent almost all bites. If you must walk in vegetation or rocks where you cannot be sure what you’re stepping on, wear high-top shoes or boots that cover your feet and ankles. Of course, boots that cover some of your calf are better but most accidental snakebites (where the person is not intentionally interacting with the snake) are to the ankle and below.
Remember, you can’t save all the cute animals, eat all the tasty ones, kill all the ones that scare you, and have a functional ecosystem, too!
After some limited basking by a couple of telemetered rattlesnakes and short movements by Female 41 and Male 49 two weeks ago, the return of cool cloudy weather sent these animals underground again. Then, despite several days of sunny weather last week, the rattlesnakes remained cool and out of sight – with the exception of Male 49. While Female 41 stayed underground and cold, he napped most afternoons in the sun just a couple feet away.
But yesterday, everything changed. Males 37 and 38 who spent the winter near the top of the bluff had moved down; one was out of sight under a log and the other was in an ambush coil in the grass under a small fig tree.
A rapidly pulsing transmitter told me before I arrived that my underweight geriatric Male 40 had survived another winter and was in the sun at his winter shelter.
Female 41 was in the sun with Male 49 laying partly on top of her, although I saw no active courtship during my brief visit.
Other telemetered snakes had moved short distances but were out of sight when I was there yesterday. But in my travels, I also sighted five unmarked rattlesnakes basking, including a beautiful little female sporting a food bolus about the size of a vole – so she has already fed successfully.
So for the next three months or so, females will be trying to eat as much as they can to nourish their next brood while males will be wandering all over the place (and hunting less) in search of receptive females. It is during this time that male rattlesnakes tend to turn up in yards and on trails, producing interactions with people.
Remember that rattlesnakes want nothing to do with something the size of a person. Leave them alone and they will be happy to avoid you, too!
As sunny 70+ degree days have been warming the ground recently, body temperatures of the telemetered rattlesnakes have been creeping up… but only from the 50-60F range into the higher 60s. Then, yesterday, I found Female 41 laying in the grass next to the log where she has spent the past two winters. But just being visible doesn’t constitute emergence.
I returned late this morning and found Female 39 visible for the first time, basking in dappled sun under the end of the log where she has spent the past two winters.
Then, not only did I find that Female 41 had moved about 20 feet from her winter shelter to a smaller nearby log, but I watched as Male 49, a large non-telemetered animal with white/green paint in his rattle, arrived at the same log and joined her. Although they coiled next to each other, I saw no actual courtship behavior during the 45 minutes I watched.
Body temperatures of Males 37 and 38, both of which spent the winter high on the bluff north of the preserve, were around 90F at midday today. Because their winter locations are almost inaccessible, I don’t know if they have actually left their hibernacula but they were both definitely in the sun today. Remember that ground temperature is much higher in the sun than the air temperature, particularly on south-facing slopes.
As of today, the other five telemetered rattlesnakes remain relatively cool and out of sight and the weather forecast looks like cloudy skies, cooler temps, and more rain over the next few days.
The study animals did not begin to leave their winter shelters until the end of the first week in March last year. Although this is three weeks earlier than last year, it is certainly a limited emergence with many snakes remaining underground and inactive. But the fact remains that some rattlesnakes have left their winter shelters, so “rattlesnake season” is definitely underway.
Remember that being careful where you place your unprotected hands and feet and leaving snakes alone when you find them would prevent almost all rattlesnake bites!
At the conclusion of my last post (29 October), I mentioned that it appeared that all of the rattlesnakes had settled into their winter shelters and, indeed, two days later, they had done so. Of the six telemetered rattlesnakes for which I have refuge locations from last winter, five of them returned to the same refuge this winter. The exception is Male 38, who spent last winter under a log in the flat floodplain but this winter is high on the hillside near the northern preserve boundary. On 31 October, I climbed the bluff and found Male 38’s location just a few meters away from Male 37, who had returned to his 2014/2015 location. Both animals’ radio signals were originating from under thick vegetation covered by wild grape vines high on the steep hillside. Both are undoubtedly underground, likely in ground squirrel burrows.
Of the eight telemetered rattlesnakes this winter, three are hibernating together in one refuge, two are together in another, and three are hibernating alone in different locations (including Males 37 and 38). Of course, there may be non-telemetered rattlesnakes with any or all of them but that’s hard to determine at this stage. I will do some burrow camera work soon but that is often inconclusive, as their shelters are often too deep or have too many sharp turns for the camera. Even if I reach them, I cannot visually identify the snakes without looking for paint in their rattles, which are usually covered by their coils. In warm weather I often poke them with the camera to get them to expose their rattles but I don’t want to disturb them when they are cold.
So how cold are they? The table below shows the ground surface temperature in sun and shade, as well as the coldest, warmest, and average body temperatures of the telemetered rattlesnakes over the last five weeks of 2015. I will continue to occasionally collect temperature data until spring emergence. That will be foretold by the snakes beginning to bask in the morning sun, probably sometime in March. Even if I don’t catch them basking, they won’t be able to hide their elevated body temperatures; then we’ll know they’re about to begin their 2016 season.
Brumation vs. hibernation
Several people have asked me about the term “brumation” and the difference between brumation and hibernation. According to Harvey Lillywhite’s Dictionary of Herpetology (2008, Krieger Publishing, Malabar, FL), brumation is:
“A condition of torpor during extended periods of low temperature (winter dormancy), intended to distinguish such states of inactivity of amphibians and reptiles from the term ‘hibernation’ that is used commonly in reference to birds and mammals. Term coined by W. Mayhew (Comp. Biochem. Physiol. 16:103–119, 1965)”
When used regarding endotherms (animals that regulate their body temperature internally, like birds and mammals), “hibernation” generally describes a state of deep sleep in which an animal lowers its body temperature and metabolism to conserve stored nutrients during harsh winters, such as many bears and bats. In reality, ectotherms like rattlesnakes do essentially the same thing with two exceptions: they remain responsive and do not “sleep” although they become very sluggish as they get cold and, after all, it is not a big physiological change for ectotherms because their body temperature varies with the environment year-round. As a result, many herpetologists ignore brumation and use hibernation to describe winter inactivity in reptiles and amphibians, too (e.g., F. Harvey Pough, et al. 2016. Herpetology, 4th ed., Sinauer Associates, Sunderland, MA).