Nerodia fasciata fasciata, the Southern banded watersnake
Grand Bay Wildlife Management Area in Lowndes county, Georgia; 05 April 2012
Forgive me; this post is long. I’m working this stuff out through the act of writing…
So far, on our still-young phosTracks photoblog, we’ve seen a handful of Florida banded watersnakes, Nerodia fasciata pictiventris. This post presents the South Georgia representative of the species, the Southern banded watersnake, Nerodia fasciata fasciata. That being said, the snake you see here probably isn’t a Southern banded watersnake. Also, yes it is. And why so? Because it probably isn’t. Confused yet? Welcome to my world.
I find myself quite skeptical of most subspecies designations. In many cases, such subspecies divisions were originally driven by archaic observations based on phenotypic, observable characteristics such as colors, size, weight, and so on. In a sense, this potentially disregards simple regional variations seen from population to population — or, actually, it makes too much of them. Is such regional variation enough to warrant an actual phylogenetic division within a species? Well, maybe. Or maybe not. And what is a “sub”species anyway?
Linnaean Classification of dogs, from cnx.org.
As DNA sampling and phylogenetic analysis continues to advance, we live in an era where taxonomic classification as we know it is undergoing a significant evolution of its own. I do indeed believe that the Linnaean world I grew up in, the one where we learned Kingdom-Phylum-Class-Order-Family-Genus-Species, is on its way out. Dating back to the 1750s, Linnaeus founded our modernistic notion of binomial nomenclature and the taxonomic schema I was taught when I was a kid. It was nifty. Everything fit into nice little boxes (as seen in the educational image to the right). As time progressed, however, we started adding strata to the system to accommodate increasingly-perceived complexities in the real world of organic life and evolutionary diversity. We started creating suborders, infraclasses, suprafamilies, and so on. It turned out the world was far more complicated than the Linnaean system was originally designed for.
Picture this: Imagine you’re put in charge of building a writing center for new, local college from the ground-up. You don’t really know how many students you’ll ultimately serve, and you certainly don’t know what kinds of assignments they’ll ultimately need help with. Looking around, you think you have a pretty good idea of what to expect, so you build your writing center as best you can. Once the doors open and students start coming in, however, you soon discover there are far more students than you anticipated, and these students have a wide-ranging orders of concern. Some need help with commas, other with organization. Some have difficulty navigating digital technologies, others have no sense of audience. Further, you realize that different courses and disciplines have radically different types of assignments, and different teachers have radically different measures of assessments, goals, and outcomes. In short, you realize that the writing center you built is not optimally designed to deal with all of these issues as they exist in the actual, real, and fluid world. So, you have a choice: Do you add on to the already-flawed writing center infrastructure? Or do you build a new system to better accommodate your students and their ever-changing needs?
That’s sort of the dilemma with Linnaean taxonomy. For decades, we’ve been adding on to that flawed, closed system, keeping those little boxes in place. There is, however, a move to create a more-fluid, more-open system of cladistics — one that focuses on taxa as clades, subsections of taxa not necessarily bound to the pre-existing strata-titles of the Kingdom-Phylum-Class-Order-Family-Genus-Species box system. Life is, after all, rather complicated and ever-changing, and complicated systems rarely function with easily-divisible binaries. Cladistics recognizes that everything is still happening. In a way, Linnaean classification is based on categorizing things into their little boxes, whereas cladistics is about better understanding evolutionary relationships and the movement of species through evolutionary history. In Linnnaean taxonomy, to think that a static system will serve radically different taxa —such as the insects and the mammals— equally is, come to think of it, absurd. It’s like having two shoeboxes and then trying to jam radically different balls equally into each box. A tennis ball is not a golf ball. Anyhow, it is along these lines, the lines of the judicially nice and tidy boxes of Linnaean taxonomy, that my faith in subspecies designations begins to fall apart.
Cladogram of dogs from FractalFoundation.org.
Though it does sometimes occur, rarely are there natural borders that actually divide one subspecies from another (like, say, a massive river or a mountain range). Instead, you often end up with broad zones of intergradation — areas where two subspecies overlap and breed with one another. This is assuming that subspecies designations are valid divisions in the first place. Thinking about these “zones of intergradation” helps us keep and maintain our binary thinking with the subspecies: This is the Florida subspecies, and that is the Southern subspecies; it just so happens that sometimes they cross over. Sometimes they’re both.
But what if we just disregard that “subspecies” division altogether? What if we recognize that the phenotypic nature of species can range and “drift” not just in time but also in space? What if it’s not a matter of black and white, this vs. that? What if it’s all just shades of gray. This is why I’m attracted to cladistics/cladograms (seen in the insert to the right); it affords more value in the flowing of evolution history and diversity not just between species, but also within species, and it does so without trying to add labelled boxes onto the other labelled boxes. With cladograms, it’s easier to think of variation within species without feeling obligated to box and divide one subspecies from another.
So, that brings us to the banded watersnake we see here…
Nerodia fasciata is the recognized genus and species for the banded watersnake. In North America, we have three currently-recognized subspecies, two of which are in Florida and/or Georgia: The Florida banded watersnake, Nerodia fasciata pictiventris, and the Southern banded watersnake, Nerodia fasciata fasciata. In peninsular Florida, the default is always N. f. pictiventris. In central Georgia, the default is always N. f. fasciata. But what about the border in between? What about the broad zone of intergradation in far north Florida and extreme south Georgia? Truly, the “line” between the two species is somewhat a myth when you step into that fluid zone of intergradation.
In their book, North American watersnakes: A natural history (2003), J. Whitfield Gibbons and Michael E. Dorcas describe in great detail the natural histories of each of North America’s watersnake species. Regarding the two banded watersnake subspecies at stake in this post, Gibbons and Dorcas draw from a 1938 article by William Clay and summarize the differences as follows:
“Nerodia f. fasciata have dark ventral markings that are square, and they usually have more than 128 ventral scales. in contrast, N. f. pictiventris have ‘elongate dark areas near the anterior margins of the ventrals’ and fewer than 128 ventral scales’ (Clay 1938b). Both of these subspecies vary extensively in color pattern, having red, brown, or black crossbones dorsally on a lighter-colored yellowish, gray, or reddish background. Large individuals of either subspecies may become melanistic, obscuring the banding pattern, which may be obscure even in some smaller individuals.”
Elsewhere, in the Amphibians and reptiles of Georgia (2008), Camp, Gibbons, and Elliot write, “Of the three known subspecies, only the southern banded watersnake (N. f. fasciata) is found throughout most of the species’ Georgia range. The Florida banded watersanke (N. f. pictiventris) may occur in extreme southeastern Georgia.” This is further supported in Snakes of the southeast (2005), also by Gibbons and Dorcas, in which the authors show the only overlap of N. f. pictiventris into Georgia to be the tiniest tip of its southeastern and coastal edges.
The snake featured on this post was photographed at Grand Bay Wildlife Management Area in Lowndes county, Georgia, just northeast of Valdosta and perhaps two dozen miles from the Florida-Georgia border (well away from the coast). Judging by the range maps and distribution comments provided in Amphibians and reptiles of Georgia and Snakes of the southeast, this should be Nerodia fasciata fasciata, the Southern banded watersnake. Color pattern differentiation is essentially worthless in my experience because both subspecies are so heavily variable, so I decided to count the ventral scales on the snake. According to the Clay system (noted above), if the sum total of ventral scales is under 128, it should be the Florida banded watersnake. If it’s over 128, it should be the Southern banded watersnake. And what was the final count for the snake featured here? The snake that, according to the range maps, should be the Southern banded watersnake? The final count was 126 — two clicks into the Florida banded watersnake range. Ah, shit.
Which was it? Southern banded watersnake? Florida banded watersnake? Clearly this was an intergrade of the two, as were all the other banded watersnakes I photographed and worked with during my two years in southern Georgia. I repeated my ventral count a number of times with different individuals and came up with varying numbers between 126 and 130, both within the Florida and Southern subspecies ventral-count ranges.
So, yeah, the whole subspecies thing is a bit of a mess. So many of these subspecies divisions are based on fairly insignificant and erratic morphological differences between populations over space, but who’s to say that these physical differences necessitate such phylogenetic differentiation? After all, the Florida banded watersnakes of South Florida will likely continue to adapt differently than the Florida banded watersnakes of South Florida — as human development increases and as new, non-native species continue to arrive. In speciation and diversity, evolution never really shuts down. Adaptation doesn’t call it a day. A “species” as we know it today is simply another transitional form in the fluid flow of time and evolution. Perhaps general “subspecies” nomenclature can be helpful in that it helps us understand the general regionality of what we’re talking about, but I prefer to not think of it as black vs. white when it comes to subspecies. As far as I can tell, the differences between the Florida and the Southern banded watersnakes are negligible at best, and with such a broad zone of intergradation, we really are dealign with fifty-thousand shades of gray.
The act of writing this post has forced me to reconsider how I’m tagging and linking species on phosTracks. I’ve been using traditional and contemporary subspecies trinomials in both the posts and in the main species index on the right column of the main page. I’ll continue using subspecies trinomials within blog posts because that’s how so many of us are wired to think about these organisms, but I’m going to stick to the binomial (genus+species) model in the big species list on the right. For example, instead of having two entries in the big list, one for the Florida banded watersnake and one for the Southern banded watersnake, I’ll just have one single link for all Banded watersnake subspecies. That sounds like a decent compromise with myself. Heh.
phosTracks will be back in a few days. I want to retro-fix some metadata on the site in response to this post and then take a little bit of a breather!
Gibbons, J. W., & Dorcas, M. E. (2004). North American watersnakes: A natural history. Norman: UP Oklahoma.
Jensen, J. B., Camp, C. D, Gibbons, W., & Elliot, M. J. (2008). Amphibians and reptiles of Georgia. Athens: UP Georgia.
Gibbons, W., & Dorcas, M. (2005). Snakes of the southeast. Athens: UP Georgia.
The Southern Banded Watersnake (Nerodia fasciata fasciata), 05 April 2012 Nerodia fasciata fasciata, the Southern banded watersnake Grand Bay Wildlife Management Area in Lowndes county, …