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.

unidentified Crotalus oreganus under log at Refuge 03 on 03 June 2015, Effie Yeaw Nature Center Origonal RAW IMG_7387.CR2

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.

More feeding, courting, and a new female

The first thing I encountered early in the afternoon of 13 May was a dead adult rattlesnake in the grass behind the Visitor Center. The carcass had completely putrefied and had been eviscerated by insects. It was an average-sized adult and had no paint in its rattle (i.e., it was not one of our study animals). I could not determine the sex. The head and rattle were intact, indicating it was unlikely a human had killed it. There was no obvious evidence of trauma elsewhere, either, but it was in such bad shape that it was difficult to tell. People often ask about the life expectancy of rattlesnakes and the answer is that, in captivity, they have been known to live more than 30 years. But in the wild, if they make it to adulthood (which few do), they are constantly threatened by raptors, coyotes, kingsnakes, people and their cars, temperature extremes, and other hazards. My guess is that few make it to ten years.

About 3 PM on the same day (13 May), I found the radio signal from Female 41 coming from dry grass on the hillside near the stairs on the trail behind the amphitheater. When I refined her location and parted the grass, I found her swallowing a small rodent – which she immediately spit out (this is common defensive behavior, as rattlesnakes are defenseless when swallowing prey). I had to remain completely still for five minutes or more before she decided it was safe to eat again. Because she was so deep in the grass and I had only a small opening through which to view her (photo below), I could not tell exactly what kind of mammal she was eating. It was uniformly gray with very fine fur, a gray belly, and was about mouse-sized. It was too large for a shrew and the lack of a light-colored belly ruled out most native mice. The dorsal fur looked too fine for a vole and I could not see the tail. What I could see looked like a house mouse, Mus musculus, but we will never know for sure as I did not want to bother her further and be the cause of her abandoning her kill.

adult female Crotalus oreganus (CROR41, in situ) swallowing an unidentified rodent (Mus or Microtis) at Effie Yeaw Nature Center on 13 May 2015 Original RAW ING_7074.CR2

When I next visited her two days later, she had moved only a few feet and was coiled next to a California Ground Squirrel burrow, which she retreated into when I arrived. This was the first time this season I have found a rattlesnake close to or in a ground squirrel burrow. I haven’t seen ground squirrel pups yet but I saw a pregnant female on 11 May, so I suspect there are pups in burrows by now and the rattlesnakes hunt the pups heavily in spring and early summer. (Click here and scroll to the bottom of the first page for more info on the fascinating interaction between Northern Pacific Rattlesnakes and California Ground Squirrels)

With all of our telemetered rattlesnakes alone and apparently hunting (and not having found a courting pair since 20 April), I was beginning to wonder if courtship might have concluded early this spring. But on 16 May, I found Male 38 on top of an unmarked female next to the large log in the meadow. She was heavy, healthy, and had no paint in her rattle – a beautiful rattlesnake, but she was also quick! As so often happens, the courting male was far less afraid of me (too much testosterone?!?!) than the female, who fled immediately and made it under the log before I could capture her.

I wish that people who fear rattlesnakes and think they are so malevolent could see how these animals really react to being approached by a person. If I had been able to catch all the unmarked rattlesnakes that have escaped from me this spring, our quota of seven males and seven females with transmitters would be full… but they are shy and very quick to flee into the grass, wanting nothing to do with something as big as a person. Remember, in their tiny primitive brains, rattlesnakes react to encounters based on three criteria: Can it eat me? Can I eat it? Can I mate with it? Clearly, we fall into the first category!

On 19 May, I found Female 47 in the meadow a little after 10 AM, crawling slowly through the grass, carefully tongue-flicking as she moved. I took a few photos and she crawled out of sight while I recorded my standard data. But when I started to depart, I came across her a couple of meters away with an alligator lizard, Elgaria multicarinata, in her mouth. She retreated a short distance to a small shrub with the lizard still in her mouth and, after I stood motionless once again for several minutes, she began to swallowed it (photo below).

Female Northern Pacific Rattlesnake (Crotalus oreganus; CROR47; in situ) eating Southern Alligator Lizard (Elgaria multicarinata) at Effie Yeaw Nature Center on 19 May 2015. Original RAW IMG_7254.CR2.

Two days later, while searching for Male 35 on 21 May, his signal led me to a large and very dense thicket of armpit-high Milk Thistle (Silybum marianum) in the meadow. I have avoided penetrating this thicket when various males have occasionally visited it this spring because the spines go right through all clothing and there is just no way to avoid them. But on this cool morning, when I peered over the thistle into a little clearing in the thicket, I saw a nice unmarked rattlesnake laying in the grass, probably hoping the clouds would clear and allow some sun to shine through (photo below).

CROR53 in situ at capture site. Original RAW IMG_7320.CR2

Hoping that she was a “she” and probably the reason that Male 35 was also in the thicket, I carefully retreated without spooking her and dropped all my gear in the grass except a cloth bag and snake hook. Stepping into the sea of spines, I slipped the snake hook under her and began to gently lift her before she reacted. I almost got her clear of the thistles before she wriggled off the hook but I was able to quickly catch her again before she completed her escape. Once clear of the thistles and in the dry yellow grass, she could no longer hide and I had her in the bag shortly thereafter.

Her sex was later confirmed and she became CROR53. (Click here for an explanation of CROR). She was very heavy for her length and full of shiny white abdominal fat when I implanted the transmitter. I could not be sure of her reproductive condition because of so much material in her bowel; she clearly had been feeding very successfully. Reproductive condition is determined by palpating her belly through her belly scales, feeling for yolk masses and, later in the season, for embryos. Because the transmitter incision is less than 3/4 of an inch long and made on the side, two scale rows up from the end of the belly scutes, it doesn’t help in determining reproductive condition. When she was released the morning after her surgery, she became our fourth telemetered female, along with four telemetered males (plus two additional males with failed transmitters I am still hoping to recapture). I am up to 413 recorded encounters with EYNC rattlesnakes this year… we are off to a great start!

Soon to come: an explanation of shedding and rattle growth.

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.IMG_5745

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.

CROR35 eating vole 27Mar15A  CROR35 eating vole 27Mar15B

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!