Species Profile: Alligator Snapping Turtle

There are few more intimidating and mysterious animals in North America than the Alligator Snapping Turtle, Macrochelys temminckii. And few are more deserving of this aura than the mighty Alligator Snapping Turtle: Adults reach a whopping 175 pounds, are characterized by a row of nasty looking spikes down their back, and have a bite that is fully capable of breaking a broomhandle in half. Did we mention that one of their prey items is alligators (admittedly, small ones)?

Like all turtles, Alligator Snapping Turtles are long-lived. They do not achieve sexual maturity until approximately 11-13 years old (Dobie 1971), and they've been known to live at least 70 years (Gibbons 1987). I've always heard stories about a snapping turtle caught in the 1980s with a Civil War Era bullet lodged in its shell, but I'm essentailly unable to confirm that story (google has failed me).

Snapping turtles tend to attract amazing stories. According to unverified reports, the largest alligator snapping turtle ever was caught right here in Kansas, on the Neosho River, and weighed in at over 400 pounds. The largest verified specimen was 236 pounds, and I can't even imagine what I would do if I was dragging a seine net up and realized that a 200 pound snapping turtle was slowly crawling along the net towards me. I'm always a little freaked out by pulling up small common snapping turtles.

Oh, what was that? You didn't realize there were two kinds of snapping turtle? The far more numerous Common Snapping Turtle, Chelydra serpentina, is a totally different species. There are a lot of fairly obvious morphological differences (no spines on an adult common), you can see them side-by-side here. If you live in Kansas, the odds are you've never seen an Alligator Snapping Turtle in the wild. The common snapping turtle gets big (25-30 pounds in the wild), but not nearly as big, and frankly, that's part of what makes the alligator snapping turtle so much more awesome.

This is a 1-year old! (source)

So if you're a hundred pound turtle with no natural predators, what do you spend your time doing? Finding enough to eat! A study in Arkansas and Louisiana found a huge list of food items in the guts of 109 sampled individuals. By far the most abundant item was fish, but there were also crawfish, molluscs, other turtles, insects, Myocastor coypus (nutria), and even armadillos, possums, squirrels, hogs, snakes, eggs, and vegetation (adults are somewhat omnivorious; Elsey 2006). In all likelihood, several of these species were consumed post-mortem. Other studies have found acorns to be a significant source of food (see citations in Elsey 2006). In captivity, alligator snapping turtles have eaten just about anything put in front of them.

I wouldn't recommend trying this at home. (source)

Alligator snapping turtles are a uniquely North American species, and are restricted to river systems draining into the Gulf of Mexico. Which isn't to say these aren't extremely mobile guys. Unlike common snapping turtles, alligator snappers stick almost exclusively to large river systems. Instead of swimming they seem to mostly get around by just walking on the bottom of the river, which isn't the same as saying they don't swim at all. Individuals have been shown to move 27-30 km in a 3-year span and 16 km in a 2-month span (discussion in Shipman and Riedle 2008). That may not seem like a lot until you consider that individuals prefer to spend most of their time hiding out in cover, and get around by walking along river bottoms.

Despite all that movement, individual river systems seem to have genetically distinct populations of snapping turtles. Roman et al. (1999) found that 12 drainages had unique genetic characteristics, thereby justifying their management as separate units. This is probably explained by the fact that these turtles have never been found to move through terrestrial habitats (again, unlike common snapping turtles). In fact, common snapping turtles seem to be one large, genetically similar population across virtually all of North America.

Alligator snapping turtles are protected at the state level to some degree everywhere they occur, but have repeatedly failed to achieve federal status. This is borderline astonishing, since a wide range of data suggests that the species has been in rapid decline since the 1950s, continues to be threatened by the development of dams, and is considered a culinary delicacy in many parts of the United States. Overharvesting has been shown to dramatically affect the population structure of the species, even decades later. As with many other long-lived species, it is generally believed that the loss of reproductive adults will take a long time to recover (as I mentioned above, it takes over a decade for an individual to reach maturity). Within Kansas the Alligator Snapping Turtle is a Species in Need of Conservation.

Lit Cited:

Dobie, J.L. 1971. Reproduction and Growth in the Alligator Snapping Turtle Macroclemys temmincki (Troost). Copeia 4:645-658.

Elsey, R.M. 2006. Food habits of Macrochelys temminckii (Alligator Snapping Turtle) from Arkansas and Louisiana. Southeastern Naturalist 5:443-452.

Gibbons, J.W. 1987. Why do turtles live so long? BioScience 37:262-269.

Shipman, P.A. and J.D. Riedle. 2008. Status and distribution of the Alligator Snapping turtle (Macrochelys temminckii) in Southeastern Missouri. Southeastern Naturalist 7:331-338.

Species Profiles: The Flathead Chub

Threatened and Endangered species tend to fall into one of the following categories: Rare to the point where they are never seen, abundant in certain locations, or just damned hard to find. The Flathead Chub (Platygobio gracilis) falls into that first category. For example, some researchers at K-State have been sampling the Kansas River for most of the last three years. In 2007, they pulled out >36,000 fish, but didn’t find a single flathead chub (Eitzmann and Paukert 2007). Needless to say, this one is on the list of Kansas T&E species that I haven’t seen.

The flathead chub is a member of the minnow family (Cyprinidae), otherwise known as the largest family of freshwater fishes in the world (~286 species in N. America alone). Like most members of this family, flathead chubs are small (typically 9-16 cm). This species has a relatively broad, wedge-shaped head with long, sickle-shaped pectoral fins. Coloration is greenish or brown on top, with plain, silvery sides. The species was first described way back in 1836 (McPhail and Lindsey 1970), although the name has changed a lot (the genus flipped back and forth from Hybopsis to Platygobio a few times from 1950-1989, so you will see some references to H. gracilis here and there).

This species has occasionally been broken up into two subspecies: A large-river variety (H. gracilis gracilis; primarily the Missouri) and a smaller stream variety (H. gracilis gulonellus; primarily in the Ark River drainages). The transition zone between these two subspecies was Kansas, and the possibility of distinct genetic lineages was disputed (some people disagree with designating these as subspecies). That question will probably never be resolved due to the species’ afore-mentioned “rare to the point where they are never seen” status in Kansas.

The absence of this species isn’t terribly surprising, although that doesn’t make it somewhat depressing. Like the Pallid Sturgeon, the Paddlefish, the Chestnut Lamprey, and a ridiculous number of other fishes, the flathead chub has been eviscerated as a part of the natural community in large portions of its range by the construction of impoundments. The species occurs widely throughout North America, but is only endangered in the southern extent of its range (see the pretty map from Natureserve). The large rivers in the Missouri/Mississippi Drainage have been extensively regulated, thus wiping out avenues for upstream migration, and also reducing the productivity associated with flood events and backwater areas.

Meanwhile, less regulated areas of the flathead chub’s range continue to support viable populations. In Wyoming, the occurrence of the chub may be declining (Patton et al. 1998), but at least it is still out there. NatureServe actually lists them as being ‘secure’ throughout Colorado, Nebraska, Wyoming, Montana, South Dakota and much of Canada.

Little is known about the spawning habits of this species, although in Kansas most spawning occurs between July 1^st and August 15^th. As an agency, Kansas Department of Wildlife and Parks is supposed to have a recovery plan for this species, but as yet none has been written. I have a hard time imagining such a recover plan that didn’t include propagation efforts, and KDWP currently does not propagate any fish species for non-game purposes. At this point, the agency primarily protects this species by restricting activities in waters where the species potentially occurs (the Kansas and Arkansas Rivers) during the spawning period.

Want to read more about this species? I recommend this conservation assessment by Rahel and Thel (2007; pdf).

Lit cited:

Eitzmann, J.L. and C. Paukert. 2007. Annual performance report: Distribution and Abundance of fishes in the Kansas River.

McPhail, J.D. and C.D. Lindsey. 1970. Freshwater Fishes of Northwestern Canada and Alaska. Fisheries Research Board of Canada, Bulletin 173. Ottawa, Ontario, Canada.

Patton, T.M., F.J. Rahel, and W.A. Hubert. 1998. Using historical data to assess changes in Wyoming’s fish fauna. Conservation Biology 12:1120-1128.

Rahel, F.J. and L. A. Thel. 2004. Flathead Chub (Platygobio grcilis): A technical Conservation Assessment. Rocky Mountain Region, USDA Forest Service.

The Disappearing Eastern Spotted Skunk

There are few more interesting inclusions into the Kansas Threatened and Endangered species list than the Eastern Spotted Skunk (Spilogale putorius). The species is known by various names (e.g., polecat, civet cat) that indicate its distinction from its more common cousin skunks (the Latin name essentially means “stinking spotted weasel”). Where striped skunks (Mephitis mephitis) are more barrel-shaped and hefty, the spotted skunk is more weasel-like in appearance and small. A host of exceedingly detailed characteristics are used to distinguish S. putorius from S. gracilis (the Western Spotted Skunk), but hardly anyone honestly believed they were separate species until it was revealed that S. putorius breeds in Spring and has babies early summer, while S. gracilis breeds in the fall and has babies the following spring (Verts et al. 2001). Like all other skunks, the spotted skunk does have the skink-emitting, anal-scent glands so characteristic of skunks in general. In fact, Johnson (1921; described in Butfiloski and Swaygnham ), the spotted skunk will actually stand up on its front legs to more effectively direct its anal sacs towards a potential threat. I wish I could find a video of this behavior, but I did find a photograph (from Kinlaw 1995).


Both striped and spotted skunks have a long history as important furbearers. From the 1920s-1930s the spotted skunk was the third most harvested species in the Kansas fur-trade, averaging around 100,000 pelts a year (a level apparently believed to be sustainable), which was 3rd behind the striped skunk and possums (Roy 1997). The skunk appears to be a historic resident of Kansas, and most of the Great Plains, but some authors (e.g., Choate et al. 1973) have speculated that other spotted skunk species (i.e., the western Spotted Skunk, S. gracilis) may account for sightings prior to the agricultural settling of Kansas. In general these accounts refer to the spotted skunk as common throughout the state, even more common than striped skunks in some areas, up until about the early 1940s (Nilz and Fink 2008).

The late 30s/early 40s saw the number of skunk pelts declining significantly. By 1977 the season on these skunks was permanently closed and in 1982 the species was listed as threatened by KDWP. This wasn’t a uniquely Kansas phenomenon. Declines also occurred throughout the lower Midwest (Arkansas, Iowa, Missouri, Nebraska, Oklahoma; summarized in Nilz and Fink 2008) and potentially elsewhere in the species’ range (Reed and Kennedy 2000).

The skunk feeds on all kinds of small mammals, arthropods, birds and plant matter and is generally considered an opportunistic omnivore that benefits agriculture by eating rodents. There’s been some suggestions that the skunk moved west with agriculture and attained high densities only when benefiting from old, relatively wasteful, agricultural practices. The advent of modern farming, with less waste, chemical methods to control pests (rodents) and less ‘fringe’ habitats seems to correspond to the decline of the species. As a result, modern agricultural practices are generally considered to be at the heart of the skunk’s decline.

And decline is the right word. The spotted skunk’s preferred habitat is…just about anything native. According to the available data (which is admittedly scarce), the skunk does well in woodlands, prairies, riparian areas, scrub-land, and around abandoned farmhouses. Yet despite the huge range of habitats and food options available to the skunk, it’s almost totally absent from the Kansas landscape. Almost all the recent data comes from road kills and trappers. Nilz and Fink (2008) summarized the recent data and demonstrated that the distribution of the known occurrences of this species has continued to decline in the last thirty years. Almost nobody I’ve ever met has even seen one.

There’s a lot more out there to talk about, for instance, quite a bit is known about the evolutionary origins of this species. And apparently the skunks are in the same family as ‘stink badgers’, which I’d never known even existed. Unfortunately, I’m out of time for this week.


Choate, J.R., E.D. Fleharty, and R.J. Little. 1973. Status of the spotted skunk, Spilogale putorius, in Kansas. Transactions of the Kansas Academy of Science 76:226-233.

Johnson, C.E. 1921. The “hand-stand” habit of the spotted skunk. Journal of Mammology 2:87-89.

Kinlaw, A. 1995. Spilogale putorius. Mammalian Species 511:1-7.

Nilz, S.K. and E.J. Finck. 2008. Proposed recovery plan for the Eastern Spotted Skunk (Spilogale putorius) in Kansas. KDWP.

Reed, A.W. and M.L. Kennedy. 2000. Conservation status of the Eastern Spotted Skunk Spilogale putorius in the Appalachian Mountains of Tennessee. American Midland Naturalist 144:133-138.

Roy, C.C. 1997. 1995-1996 Kansas furbearer surveys and investigations project summary. Kansas Department of Wildlife and Parks, Unpublished report, 70 pp.

Verts, B.J., N. Leslie, and A. Kinlaw. 2001. Spilogale gracilis. Mammalian Species: 674:1-10.

The Chestnut Lamprey

Lampreys are a fairly unique class of aquatic vertebrates with an apparently ancient lineage. Every description I’ve read identifies them as superficially similar looking to eels, but lacking a jaw and instead having a round, sucking disk mouth (Cross and Collins 1995, Wikipedia, the Great Lakes Fishery Commission, etc.). That is to say that they are elongated, have no scales, and have spine-less fins. However, the resemblance is pretty much completely superficial. In fact, lampreys are not even true fish (Cross and Collins 1995, Tree of Life), and in 2006 researchers found a fossilized lamprey that was virtually indistinguishable from the modern animal (article about the find) from 360 million years ago. That is way back there, when fish were just evolving jaws.

Lamprey life-history is fairly straightforward...for an amphibian (which it isn’t). The larvae are called ammocoetes, and live in mucky sediments consuming microorganisms (at one point, it was thought these were a totally different species). After 5-7 years of this, they grow eyes and a new mouth, get bigger and learn how to swim. Then they spent ~18 months swimming around and sucking the blood out of fish large enough for them to latch onto (this varies tremendously by species, Chestnut lampreys, the Kansas Threatened species I’m gradually getting to, do not survive more than a year and are adults only a very short amount of this time). Adults then move upstream into smaller streams and reproduce. Reproduction occurs by spawning (egg and sperm release) into shallow, excavated depressions in gravel beds, and the hatched offspring move downstream and spend much of their lifespan in the benthos.

Lampreys are typically thought of as blood-suckers. Indeed, many of the most well-known members of this are fish parasites that kill their host and cause untold economic damage. The introduction of sea lampreys into the Great Lakes by the construction of locks and canals for shipping essentially destroyed the fishing industry in the lakes and devastated the native trout species (probably more economic cost from that than all the benefit they’ve provided). According the Great Lakes Fishery Commission “before sea lampreys entered the Great Lakes, Canada and the United States harvested about 7 million kgs. (15 million lbs.) of lake trout in lakes Huron and Superior annually. By the early 1960s, the catch was only about 136,000 kgs. (300,000 lbs.). The fishery was devastated.”

With all that context, why is there a lamprey species listed as a Threatened species in Kansas? Well, I’m still not sure. The Chestnut Lamprey (Ichthyomyzon castaneus) appears to occur sporadically, but widely, across the Eastern United states. Early literature describes them as “little known” (Jordan 1918). Hall and Moore (1954) considered them rare, and were apparently surprised to be able to find them easily to conduct some fairly technical taxonomic studies (apparently a lot has been made about their teeth?). In 1973, the Kansas Academy of Sciences (KAS) published a list of Rare, Endangered and Extirpated Species in Kansas, and included the Chestnut lamprey as a species “Endangered in Kansas but not nationally” because its status was uncertain and no records had been collected since the 1950s. They recommended stabilizing flow and minimizing turbidity on small, high-gradient streams. Kansas T&E laws date from 1975, and I suspect but cannot confirm yet that the KAS list was adopted en masse when the state law went into effect. So apparently the species was being found with relative frequency prior to the 1950s, then stopped being found anywhere in Kansas except in the main-stem of the Missouri and Kansas Rivers. There’s a nice-looking map on Natureserve that seems to indicate the species is imperiled several places in its range, but I’ve got a touch of color-blindness and therefore I can’t really interpret the map very well (feel free to do so yourself).

Hall and Moore (1954) also described the Chestnut Lamprey’s ecology and habits in Oklahoma, suggesting that this might be one of those native species that actually benefits from the construction of large reservoirs. This is because large fish moving upstream are forced to stop at the barriers presented by reservoirs. Where they congregate, the Chestnut lamprey is able to find abundant food, and their larvae are able to survive better because of the removal of silt. However, the construction of those dams in Oklahoma also corresponds to the species disappearing in Kansas (~1950s-60s). I have a hypothesis that the lamprey throughout the Ark basin was a single, massive metapopulation that became fragmented after the construction of the dams and was subsequently wiped out by random, local, events (i.e., feedlot runoff in the Neosho basin killed hundreds of thousands of fish in the Neosho Basin).

These days, the species if found in Kansas only in the Missouri River and the Kansas River below a dam in Lawrence. That’s an extremely limited range. Cross and Collins (1995) also indicate an active record in Cherokee County (extreme SE Kansas on the Ozark Plateau). That’s actually an area of intense heavy-metal pollution due to mining activities over the past hundred or so years, and I suspect if any lampreys exist, they survive in Missouri and only occasionally wander into Kansas. The only designated critical habitat for this species in Kansas is the Missouri River, and as far as I know, KDWP is not taking any special actions or precautions for the species. This is one of those species I don’t know a lot about, so as I read more, I may write more on it.

Jordan, D.S. 1918. The Freshwater Lamprey’s of the United States. Copeia, 64: 93-96

G.E. Hall and G.A. Moore. 1954. Oklahoma Lampreys: Their Characterization and Distribution. Copeia, 2:127-135

Kansas Academy of Science. 1973. Rare, endangered and extirpated species in Kansas. I. Fishes. Transactions of the Kansas Academy of Science 76:97-106.

The Hidden Beauty of the Neosho Midget Crayfish

If there’s a group of animals that gets my complete affection without reservation it is the crayfish. I’m a alum of David Lodge’s lab at the University of Notre Dame, and although David has now moved very strongly into the realm of scientifically informing policy, back in the day he was a huge crayfish guy. Several members of his lab still work pretty exclusively on Orconectes rusticus (the dreaded Rusty Crayfish), and so I spent a lot of my graduate career thinking about crayfish. Plus they are just awesome.

The Neosho Midget Crayfish (Orconectes macrus [Greek, meaning long, in reference to long gonopods]) is a small crayfish found exclusively in the Neosho River Basin, and really almost exclusively in the Spring River Basin (Pflieger 1996). The historic range was probably more extensive, but despite its name, I think it is unlikely this crayfish occurred throughout the greater Neosho River Basin. In Kansas, this species occurs in the Spring River basin, although apparently very rarely. Beasley and Branson (1971) found these crayfish in the Missouri portion of the Spring River, but no where else in the drainage basin. Pflieger (1996) describes the species as “common…in most streams of the Neosho drainage except for North Fork of Spring River in Jasper and Barton counties.” I don’t want to suggest Pflieger is wrong, but if this is accurate, these are the only places on Earth the crayfish is common. Durbian et al. (1994) surveyed crayfish in Cherokee County Kansas (including the Spring River) and found no individuals of this species. Taylor et al. (2004) in their review of Oklahoma crayfish referred to the species as rare or uncommon and did not find it in the Spring or Neosho main-stem. The Kansas Department of Wildlife and Parks did find two individuals in the Spring River in 1996 (Ghedotti 1998), at a location they had been found at historically, but this apparently was the result of considerable sampling effort. More recently, consultants sampling around a power plant in the Spring River collected 12 individuals labeled O. rusticus. Further inspection of a voucher specimen revealed it to be O. macrus (whew!), so the species does still occur in Kansas (KDWP, pers. comm).

The Neosho Midget Crayfish is a Species in Need of Conservation (SINC) in Kansas, which means selling or take of this species are illegal, but does not afford it the kind of protection granted Threatened and Endangered species. For example, projects occurring in the habitat of Neosho Midget Crayfish do not require a Kansas Department of Wildlife and Parks (KDWP) Action Permit. However, KDWP does devote some special consideration to research projects involving SINC species, and we happen to know of a graduate student at Fort Hays State who is conducting a state-wide crayfish survey with some special emphasis on this species (or at least this area).

Like most crayfish, this species is considered to be opportunistic omnivores who will eat decaying vegetation and animal matter, as well as predate on invertebrates and small fish. According to Pflieger (1996) the species behaves more like Cambarus spp. in Missouri than like other Orconectes spp. Having not worked with Cambarus spp. in the past, I’m not really sure what he means, but this species appears to be pretty sedentary, spending most of its time under rocks or in tunnels. According to Pflieger “…not an agile or strong swimmer, and seldom takes evasive action when attempts are made to capture it by hand.” Reproduction apparently occurs in March or early April, and females virtually disappear from sampling collections during April (presumably because they were sequestered in burrows and thus unable to be collected).

The Spring River in Kansas has been the brunt of some fairly severe pollution impacts over the course of its history. The EPA has a 115 mile square Superfund site in this county (see here and here .pdf) from a history of heavy zinc and lead mining, and the Spring River is heavily polluted. Likely due to this pollution history and some fairly extensive development in SE Kansas, a number of species have become endangered or rare from the Spring River drainage. The Neosho Midget Crayfish appears to consist of several isolated populations in tributary streams that are kept disconnected by poor quality habitat in the main-stem of the Neosho and Spring Rivers. In particular, my feeling is that the Spring River in Kansas is a particularly crucial connection between apparently abundant subpopulations in Missouri and isolated populations in Arkansas, Oklahoma and Kansas. Without a strong avenue to re-connect these isolated populations, the entire species is at risk from localized impacts and genetic drift.

Lit cited:

Beasley and Branson. 1971. A partial biological survey of the Spring River drainage in Kansas, Oklahoma, and Missouri Part III. The Crayfishes. Transactions of the Kansas Academy of Science 74:228-233.

Durbian, F.E., B.J. Frey, and D.W. Moore. 1994. Crayfish species from creeks and rivers of Cherokee County, Kansas. Transactions of the Kansas Academy of Science 97:13-17.

Ghedotti, M.J. 1998. An annotated list of the crayfishes of Kansas with first records of Orconectes macrus and Procambarus acutus in Kansas. Transactions of the Kansas Academy of Science 101:54-57.

Pflieger, W.L. 1996. The Crayfishes of Missouri. Missouri Department of Conservation. 152 pp.

Taylor, C.A., S.N. Jones, and E.A. Bergey. 2004. Crayfishes of Oklahoma revisited: New state records and checklist of species. The Southwestern Naturalist 49:250-255.

The Mysterious and Elusive Blackside Darter (Percina maculata)

And with that, Analyze Everything is back. Expect a weekly post on Thursday or Friday, focused initially on Kansas T&E and SINC species. Other posts will pop up sporadically. Hope you enjoy, and feel free to ask questions.

State and federal Threatened and Endangered Species lists are riddled with species who are so rare that they may easily be considered extirpated (or even extinct; see Eskimo Curlew). Although apparently facing no global threat of extinction (even in 1973 the darter was considered endangered in Kansas but not nationally), the Blackside Darter has become a whisper of a ghost in Kansas. According to the NatureServe distribution map (see it here), the existing (or recently existing) Kansas population is confined to the Mill Creek Watershed. This is interesting because the next closest population is in Missouri or Oklahoma, between 4-8 HUC8s away.

During the Summer of 2008, KDWP conducted a survey of the Mill Creek looking for Topeka Shiners (more on them elsewhere). There was some optimism that the Blackside Darter would be found at the same time, since unlike many darters, the Blackside is a common mid-column pool resident (Cross and Collins 1995). At this point, no Blacksides have been found (although some collected specimens may be identified as such later), and as far as I can tell, none have been found in Kansas since 1974! (Drenner and Cross 1981) Actually, now that I dig around the KDWP Stream Survey database I see a record of 3 Blackside Darters found on a site in 1994. A return visit in 2000 didn’t find any, however.

I wonder if the Blackside Darters are becoming extirpated in part due to their hybridization with logperch (P. caprodes). The two species have overlapping ranges, and have been observed to hybridize in Kansas (Drenner and Cross 1981), with hybrids having intermediate physical characteristics. Winn (1958) observed logperch males chasing female P. maculate (the latter being a small portion of the overall darter population even when relatively common). Hybridization may have some weird implications for conservation of rare species (c.f. Perry 2001). I did see a logperch in Mill Creek in July of 2008 that looked ‘funny.’ I wonder if that individual has a Blackside ancestor (might explain why we caught it in a pool).

The Blackside Darter is sexually mature in their second year (Becker 1983), and females may continue breeding until they are 4 years old (Bart and Page 1992). This darter consumes primarily benthic invertebrates (NatureServe 2008), although Cross and Collins (1995) also mentioned that the Blackside Darter “sometimes rises to the surface for food.” Among Kansas darters, the Blackside is unique for not clinging to the stream bottom. In general the species tends to prefer streams that one would consider good for mayfly and stonefly species (i.e., cool, clear streams with moderate currents and gravel substrates). Many Flint Hills streams fit this description, but despite fairly extensive sampling efforts in the Northern Flint Hills (mostly hunting the Topeka Shiner), the species is only known to occur in Mill Creek.

Bart, H. L., Jr., and L. M. Page. 1992. The influence of size and phylogeny on life history variation in North American percids. Pages 553-572 in R. L. Mayden, editor. Systematics, historical ecology, and North American freshwater fishes. Stanford Univ. Press, Stanford, Calfiornia. xxvi + 969 pp.

Becker, G. C. 1983. Fishes of Wisconsin. Univ. Wisconsin Press, Madison. 1052 pp.

Cross, F.B. and J.T. Collins. 1995. Fishes in Kansas. University Press of Kansas, Lawrence KS. p 238.

Drenner, R.W. and F.B. Cross. 1981. A natural hybrid fish, Percina maculate × P. caprodes, from Kansas. Transactions of the Kansas Academy of Sciences 84:61-62.

NatureServe. 2008. NatureServe Explorer: An online encyclopedia of life [web application]. Version 7.0. NatureServe, Arlington, Virginia. Available http://www.natureserve.org/explorer. (Accessed: July 21, 2008 ).

Perry, W.L., J.L. Feder, and D.M. Lodge. 2001. Hybridization and introgression between introduced and resident Orconectes crayfishes; implications for conservation. Conservation Biology.

Soil Conservation Commission (SCC). 1973. Rare, Endangered and Extirpated Species in Kansas. I. Fishes. Transactions of the Kansas Academy of Science 76:97-106.