vole – EYNC Rattlesnake Study

Will rattlesnakes be more abundant this year?

I’ve been getting this question a lot lately. And my response is, “It depends on where you are.”

Actually, it depends on the local weather in recent years where the question is being asked.

Many of the female rattlesnakes in the Effie Yeaw Nature Center preserve have been reproducing annually since at least 2014, including the drought years prior to this extraordinarily wet winter and spring. And, as I have written here before, that’s unusual – at least according to similar studies of other species of North American rattlesnakes. This far from the equator, most pitvipers only produce a brood every two or three years because of the time it takes females to replenish sufficient fat stores to support another pregnancy (they lose 30-50% of their body weight with each litter). But along the American River Parkway during 2014, 2015, and 2016, there was enough local rain each winter to get the annual plants growing and everything blooming, providing plenty of food for insects, squirrels, voles, and other small animals. The insects fed the abundant lizards that reproduced like crazy each year, and the rattlesnakes fed on the small mammals and lizards that were plentiful. So rattlesnakes and other mesopredators (mid-level predators in the food chain; i.e. they are eaten by bigger predators) have been thriving locally, despite the historically low reservoir levels and skimpy snowpack in recent years. And because rattlesnakes produce only one brood per year, they can’t do better than they’ve been doing.

On the other hand, when local drought reduces the availability of the rattlesnakes’ prey (i.e., reduced local rainfall significantly suppresses plant growth, which negatively affects everything else up the food chain), that changes the snakes’ behavior. Among other things, movement and courtship slows significantly and reproductive rates are almost certainly reduced.

So, in an area like the American River Parkway, this wet spring is not likely to increase the rattlesnake population because they can’t reproduce much better than before all the rain. However, in areas where reduced rainfall in recent years significantly impaired floral growth , rattlesnakes will be better fed and more active this year and they are more likely to successfully produce young.

But it is important to understand that what people interpret as “abundance” is usually just a change in behavior. I studied this phenomenon in Mohave rattlesnakes in the southern California desert for my MS thesis. In dry conditions, predators get most of their water from the body water of their prey (we’re all about 70% water). So, when rodents are abundant, rattlesnakes are well hydrated, behave normally, and males search constantly for females during spring and fall. But when their prey becomes scarce and other water sources are not available, they reduce water loss by remaining coiled and moving less, thus reducing the amount of skin exposed to the dry air. This is important because cutaneous (through the skin) evaporation accounts for about 75% of the snakes’ daily water loss.

As a result, during local drought, people encounter rattlesnakes less frequently – not because there are fewer rattlesnakes but because they remain hidden and move around a lot less (top photo, below). During non-drought times, people see more rattlesnakes because their behavior is not inhibited and they crawl around a lot more (bottom photo, below), especially the males during their courtship season.

So, if you live in an area like the American River Parkway, where there has been lush growth of grasses and other annual and perennial plants in recent years, this wet year should make little difference in the apparent abundance of rattlesnakes.

But if your neighborhood has been dry with little springtime plant growth in the last few years, you may see an increase in rattlesnakes this year for two reasons: the males may be searching for females much more than before, plus there may be more baby rattlesnakes in September and October this year (and likely next) as a result of females having plenty of rodents to eat this year.

Emphasis now on hunting and shedding

Before I launch into what’s been going on with the Effie Yeaw rattlesnakes over the past few weeks, I want to pass on a link to a recent interview with Dr. Bree Putman. Bree was a grad student with Matt Holding (lead author of the journal article I linked to in my last post) in Emily Taylor’s lab at Cal Poly San Luis Obispo before she moved on to Rulon Clark’s lab at San Diego State to finish her Ph.D. In Bree’s interview (click here), she talks about ground squirrels and northern Pacific rattlesnakes and she describes some of the very intriguing behavioral questions many of us would love to answer.

Back to EYNC rattlesnakes –

This female California ground squirrel was clearly nursing pups on 26 April at Effie Yeaw Nature Center. — This female California ground squirrel was clearly nursing pups on 26 April in the Effie Yeaw Nature Center preserve.

I have not seen a courting pair of rattlesnakes since 28 April, when three rattlesnake pairs were found together in different locations. Male 37 was found with unmarked adults that were likely females on 11 and 14 May but no courtship was observed. But as the spring courtship season wound down, California ground squirrels began producing pups and the emphasis of both rattlesnake sexes turned to hunting.

After not finding a telemetered rattlesnake in or very near a ground squirrel burrow during the first 400+ observations this year, Male 46 was found in an ambush coil facing an active burrow a foot away on 9 May. In the five weeks since, several rattlesnakes have been found close to or inside squirrel burrows on several occasions.

At the same time, the snakes have also been hunting heavily in vole (aka: meadow mouse; Microtus californicus) tunnels in the grass.

Male 46 visible in a vole tunnel on the morning of 3 June 2016. Original RAW IMG_2192.CR2 — Male 46 uncovered (arrow) in a vole tunnel in the grass on the morning of 3 June. Although you cannot see it here, voles construct a maze of above-ground tunnels in thick dry grass that protects them from most predators… but not rattlesnakes. I hope you can appreciate just how impossible it would be to learn about the habits of these snakes without radiotelemetry!

And since about the end of May, the rattlesnakes – especially males – have been shedding.

Pre-shed Male 38 on 14 June 2016 Original RAW IMG_2320.CR2 — The rattle of pre-shed Male 38 under a log yesterday, 14 June. The new rattle segment forming in the base of the tail is whitish and covered by the last couple rows of scales. He is easily identified visually by red/green paint in his rattle.

Periodically shedding the corneal layer of the skin (called ecdysis; for more info, click here) takes snakes out of commission for a week or more and males seem to put it off during the spring mating season. It’s a bit like race car drivers waiting for a yellow caution flag to make a pit stop!

Even more interesting is that the rattlesnakes have favorite places where they go during this process and it is not uncommon to find several pre-shed individuals of both sexes together this time of year. Like hibernation, there seem to be many logs and burrows where they could shelter while waiting to shed but they congregate in just a few of them. Those of us who study rattlesnake behavior would love to know why. What is so special about certain locations? Or is it something else… like family ties or some other social interaction?

Love is in the air

Rattlesnake love, that is!

I’ve been getting lots of questions after last night’s (17 April) KCRA News interview with a local “rattlesnake hunter” who makes his living keeping us all safe from rattlers. If you have been reading my blog, you know there is nothing new to be alarmed about. The last few hot days have not brought out the rattlesnakes. Indeed, general emergence from their winter dens occurred more than a month ago. Male rattlesnakes continue to be busy looking for and trying to mate with as many females as they can while the females hunt for mice, rats, and other prey. As sit-and-wait ambush predators, the females don’t move much and thus are seldom encountered by people. But the males have been and will continue to constantly search for females until late May or June, in the process turning up in yards and on trails and other places where they encounter people.

Once again, most bites can be avoided by watching where you put your unprotected hands and feet and leaving rattlesnakes alone when you encounter them.

We have been encountering numerous pairs of rattlesnakes in the Effie Yeaw Nature Preserve over the last few weeks. Here are brief video clips of a couple of examples: First, our telemetered Male 62 (blue/white paint in rattle), entwined with an unmarked female. These snakes were not moving during my brief visit but the males often accompany the females for many days but cannot keep courting them continuously (click here for video). You can see why in the video of Male 37 actively courting Female 41… it takes a lot of energy! (click here).

On 1 April, several hours after I videoed Male 37 courting Female 41, I returned and just missed a fight between two males over Female 41. As I approached, I briefly saw both males, with heads and necks high off the ground, twisted around one another. But one quickly fled, either from me or he had had enough of Male 37…who is a big healthy male rattlesnake! In any event, Male 37 was all excited and remained raised off the ground for a minute or more, guarding his girl (see photo below).

Male 37 still ready to fight shortly after another male retreated. Female 41 is coiled in the shade just below the male. I had videoed Male 37 courting Female 41 just a few hours before this scene. — Male 37 still ready to fight shortly after another male retreated. Female 41 is coiled in the shade just below the male. I had videoed Male 37 courting Female 41 at this spot just a few hours before this scene.

A video made a few years ago of two males fighting on the concrete in front of the Effie Yeaw Visitor Center doors is on the Nature Center website (or click here to see it).

Courtship has begun

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).

Female 55 basking alone in the afternoon sun on 22 March 2016. Note her rattle with red over yellow paint, visible through the grass on the left side of the photo. Without a radio, she had not been seen since June of last year.

type here — Then two days later, on 24 March, Female 55 is almost completely hidden by a new male on top of her; you can see his unmarked rattle in the middle foreground of the photo. Although the male is distracted by me in this picture, he had been vigorously courting the female just moments before. Unfortunately, the video camera auto-focused on the grass, making the resulting image unusable. Courtship continued at this location for at least three days but they were gone on 29 March, replaced by Male 35, who was enjoying the sun by himself yesterday afternoon. Although he would retreat under the log when I approached closely, he would re-emerge within minutes as soon as I backed away.

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.

The effect of drought on rattlesnakes

Field studies of rattlesnakes indicate that they cease most movement when water stressed and remain in their established home ranges, rather than migrating into developed areas in search of water, despite frequent claims to the contrary.

In my last post, I mentioned witnessing Females 41 and 47 feeding, as well as finding new Female 53, who was very heavy and likely pregnant. Since then, I have come across Female 41 eating another vole, found another new female (#54) that is heavy and definitely pregnant, and come across a fat but unidentified rattlesnake in the refuge where Females 41 and 43 had babies last year. I could only see the face and a bit of a flank of the unidentified animal (photo below) so I couldn’t even determine sex.

It could be a male that has just eaten a ground squirrel pup – but it is more likely another pregnant female. We now have five telemetered females (39, 41, 47, 53 and 54) and all are in great shape, with three either confirmed or likely pregnant and the others in good shape to reproduce although I have not yet had my hands on them this year to palpate for fetuses.

This brings up a timely point: This will obviously be a good year for rattlesnake reproduction in our area, despite being in the midst of an historic drought. Since the news media often quotes “experts” claiming drought “drives rattlesnakes out of the hills and into yards looking for water,” this is a great opportunity to set the record straight about how drought affects rattlesnake movement.

We live in a Mediterranean climate, historically characterized by warm dry summers and cool wet winters. Even during years with “normal” precipitation, vast tracks of mountains, foothills and many valley areas have no surface water between late spring and the return of winter rains in November or December – yet they support healthy populations of rattlesnakes. Herbivores (insects, rodents, etc.) get most of their water from the plants they eat and rattlesnakes get water from eating the herbivores. The bodies of terrestrial vertebrates are usually composed of 65–75% water, so eating a 100 gram (3.5 ounce) rodent is like drinking about 70 grams (2.5 ounces) of water for a rattlesnake (plus the nutrients and energy gained). Make no mistake, rattlesnakes suck droplets from various surfaces, including their own skin, deposited by rain and dew (photo, below) and they will certainly drink from standing water when it’s available. But especially during summer and fall, these other sources are not available and virtually all of the water a rattlesnake needs is obtained from its prey.

adult Crotalus s. scutulatus (Mohave Rattlesnake) drinking rain water near Apple Valley, San Bernardino County, California, USA [wild animal, in situ] — Mohave Rattlesnake drinking rain water from its own body. [Cardwell, M. D. 2006. Rain-harvesting in a wild population of *Crotalus s. scutulatus* (Serpentes: Viperidae). *Herpetological Review* 37:142-144.]

Rattlesnakes are models of low energy physiology. As ambush predators, they move comparatively little and rely largely on anaerobic metabolism. Their sedentary lifestyle combined with the corneal layer of their skin (full of water-blocking lipids) dramatically lowers the amount of water that passes into and out of their bodies – known as “water flux.” Nonetheless, multiple studies have shown that the most significant mechanism for water loss in terrestrial snakes is evaporation, with about 75% being lost through the skin and the remainder via exhaled breath.

During my four-year field study (2001–2004) of Mohave Rattlesnakes in southern California, I was able to compare behavior, including average daily movement and reproductive effort, between the severe drought year 2002 and 2003–2004, when rainfall returned to average or above average. I found that average daily movement during 2002 was less than one third of 2003–2004 averages. And while I encountered dozens of courting pairs during the two non-drought years, I observed a male courting a female on only one occasion in 2002. Yet these rattlesnakes continued to eat at a rate indistinguishable from the non-drought years, based on scats deposited in holding containers and later analyzed. These snakes were reducing exposed surface area (and, therefore, evaporative water loss) by remaining coiled and immobile, covering much of their skin within their coils. They even buried their coils partially in loose soil at times, covering additional skin area. Remaining stationary eliminated their ability to find and court mates but, as sit-and-wait ambush predators, it allowed them to continue to hunt – and obtain the body water of their herbivorous prey. They also positioned themselves behind vegetation and ground contours in 2002 to avoid wind and sun, both of which increase evaporation rates. You can find more details in my MS thesis.

We have seen similar behavior in Northern Pacific Rattlesnakes in recent years at my El Dorado Hills study site, where the rattlesnakes remained tightly coiled and stationary in deep chaparral on north-facing slopes during particularly hot dry summer weather. At Effie Yeaw, all of the rattlesnakes caught by staff around the ponds during the past year, as well as the telemetered rattlesnakes I have found there, have been males found during the courtship season. The females have remained in the woods, away from water sources. While the snakes will drink when they find the ponds (or other water sources), that’s not why the males are there… they’re wandering around looking for females! And, yet, the females are now fat and pregnant.

Drought probably does not affect rattlesnake movement until it becomes locally severe, as it did at my Mohave Rattlesnake study site in 2002. When the snakes start to become water stressed, they don’t set out into uncharted territory looking for surface water. Rather, they stop moving and hunker down where they can best reduce evaporative water loss while still striking any prey that wanders by. Currently, if the preserve at Effie Yeaw Nature Center is any indication, there is lots of annual plant growth and the vole and ground squirrel populations are thriving – and so are the rattlesnakes.

So when people find a rattlesnake in their yard during a drought, the most likely explanation is that it is a male looking for receptive females and the drought is not severe enough locally to stress the rattlesnakes. When they are truly water stressed, rattlesnakes move less – not more – than usual. Unlike most large mammals that have much higher metabolic and water flux rates and require standing water to drink, there is no evidence that rattlesnakes leave their established home ranges looking for water, despite the popular belief to the contrary. They do just the opposite.

29 March: Lots of courting pairs and a vole goes down!

We now have a total of 14 rattlesnakes marked at Effie Yeaw Nature Center, including 9 males and 5 females. This spring, I have processed, marked, and released 3 males and 2 females without transmitters (mostly too small for the transmitters). With prematurely failed transmitters in Males 36 and 37, we currently have working transmitters in 4 males (35, 38, 40 & 46) and 3 females (39, 41 & 47).

You almost need a scorecard to keep track of who has been with who over the past couple of weeks. Lots of the action has occurred at a small hollow log where Female 41 spent the last month or so of her winter slumber by herself. She departed on 13/14 March and just five days later Female 47 turned up there with Males 38 and 46. The three snakes were coiled next to and touching each other on 19 March in a narrow bit of shade. The late morning sun was hot and all three body temperatures were elevated (female = 86F and males = 91F & 93F), indicating they had recently been in the sun. The female disappeared into the log when I approached but the males were more concerned about each other than me. The smaller male, 46, was still excited and head-jerking (a common part of rattlesnake courtship) a little bit. Each time he would touch Male 38, the larger male would shove him away, pushing violently by thrusting a coil sideways. Male 46 would push back, reminding me of two brothers in the back seat on a long car ride. I suspect I may have missed some male combat earlier, which was probably cut short as their body temperatures approached dangerous levels and they were forced to get out of the sun. (Click here to see a video of male combat, shot in front of the EYNC Visitor Center in 2010)

Also on 19 March, Female 41 was found (by herself, as far as I could tell) in the refuge where Female 39 delivered her kids last year.

On 21 March, Female 47 and Male 38 were were still at the small hollow log, although laying a few inches apart and not actively courting when I was there. Male 46 was by himself several dozen meters away in the grass. At 674 mm (26.5 inches) snout-vent length, Male 46 was no match for Male 38, who measured 821 mm (32.3 inches) SVL at his recent transmitter replacement surgery. As you can see from the video mentioned above, male combat is a wrestling match and larger body size is a definite advantage. Snout-vent length or SVL is the common way biologists record body length in lizards and snakes; the tail is usually measured separately.

On 21 March, Female 41 had left the birthing refuge used by 39 last year and was coiled by herself under a pile of dry live oak branches. The following day, she had been joined by Male 46 and the two were copulating at about 11:20 AM. In the photo (below), Male 46’s rattle colors are green/red and Female 41’s are white/blue, although the blue is difficult to see through the brush.

Also on 22 March, Female 47 was still at the small hollow log but Male 38 had been replaced by Male 49, who was actively head-jerking, chin-rubbing, and tongue-flicking the female. Male 49 is not telemetered (but recognizable by white/green paint in his rattle) and at 767 mm SVL, he is not quite as long as Male 38, but he outweighs Male 38 by 28 grams (377 g vs. 349 g), Of course, I have no way of knowing if 38 and 49 even crossed paths; Male 38 could have departed before Male 49 arrived.

On 23 March, Female 41 had moved back to the birthing refuge used by Female 39 last year and had apparently been followed by Male 46; they were still together there on 24 March, although they were basking about 4 feet apart when I visited on both dates.

Female 47 was laying partly in the sun on the morning of 23 March with Male 49 nowhere in sight. Of course, without a transmitter, I had no way to find him. But they were laying together in the same place again on the next two mornings, so I suspect he was there on the 23rd, just not visible.

Female 41 remained at 39’s old birthing refuge on 25 and 26 March. Male 46 was still there on the 25th but they had been joined by Male 38. On this day, Female 41 and Male 46 were again basking apart from each other and male 38 was out of sight, betrayed only by his radio signal. The next day, 26 March, the female was basking, Male 38 was there but out of sight, and Male 46 was alone in a poison oak thicket some distance away.

By 27 March, Female 47 had left the small hollow log where she had been for nine days (with 3 males at various times) but she had been replaced by Female 41, leaving Male 38 apparently alone where he had been with Female 41 for the previous couple of days. As far as I could tell, Female 41 was also alone.

Also on the 27th, Male 35 was found a few minutes after 11 AM eating a California vole (aka meadow mouse, Microtus californicus) in thick knee-high grass next to the main trail, not far from the picnic area. The snake stopped swallowing and we were lucky that he did not spit out the rodent when he was disturbed, as rattlesnakes are quite defenseless with their mouth stretched around a meal. George Nyberg and I had to remain motionless for many minutes before the snake finally decided it was safe to continue swallowing. We shot a few photos as he finished his vole.

On the 28th, all of the telemetered snakes were coiled in vegetation, alone, and apparently hunting. Both places where most of the courtship had occurred over the past two weeks were empty. Are they finished courting? I doubt it; it’s not even April yet!