Thursday, February 6, 2014

A brief meditation on the joys of packrat middens

One more trip to the Quaternary, for a personal favorite...

Imagine someone has asked you to describe what an area was like hundreds, thousands, perhaps tens of thousands of years ago. They're interested in the ecology, the climate, and so forth. What could you use to accomplish this? If there are lakes and marshes, you might start by taking sediment cores and looking for pollen, which tell you what kinds of plants were present, and how the flora changed over time. If you're on the coast, you might look for deposits of shells, which are useful for describing the conditions of the water (salinity, temperature, how energetic the setting was) and for stable isotope analyses. If there's a lot of ancient wood, you might get into tree-ring data. In dry protected places of western North America, there is another type of paleoecological indicator: middens constructed by packrats.

Good try, guys, but not quite; needs more juniper and less fabric.

Packrats, also known as woodrats and sometimes trade rats, belong to the genus Neotoma. They are brown or gray and, as the name suggests, rat-like in appearance, although the tail is sometimes quite furry. Packrats gather materials such as plant fragments, bones, and dung to build dens in sheltered areas like caves and crevices. Their less-common name "trade rat" comes from their habit of dropping what they are carrying if they find something they like better (such as shiny things). There are numerous species spread across North America, some of which have stronger den-building behaviors than others, and some of which apparently prefer certain types of den-building materials. Packrats have unwittingly left us excellent records of the plants, animals, and ecology of the areas around their dens. The secret ingredient, and I warn you that if you are eating, you may perhaps want to pause for the next few paragraphs, is rat pee.

"Who, me?" A bushy-tailed woodrat (Neotoma cinerea) at Wind Cave, posing for a National Park Service photo.

Fossil middens, "resembling blocks of asphalt with the consistency and mass of an unfired adobe brick" (Spaulding et al. 1990), represent only part of an active den. Packrats will typically use a portion of their den as an outhouse, and unused and discarded plant fragments that accumulate in these areas become saturated in rat urine. In dry climates, the urine crystallizes into a substance called "amberat" which has a number of useful properties. Most obviously, it binds the midden materials. It is self-sealing to an extent: it rehydrates under humid conditions and becomes sticky, so dust and dirt become trapped on the outside surfaces and prevent moisture from penetrating more deeply. Saturating plant material with amberat is comparable to packing it in salt, protecting it from decay. Amberat also appears to prevent lichen growth and insect feeding. Finally, it affixes material to shelter walls and ceilings, so it is not uncommon to find "hanging" middens where underlying materials have eroded, providing one way to examine changes in cliff retreat (Spaulding et al. 1990). Packrats are solitary animals (Vaughan 1990), and individual middens are thought to represent a few years of activity. However, choice settings are reused over and over again, so large fossil middens spanning thousands of years can be produced. There can be millimeter-scale layers in a midden deposit, which may represent "melted" remains of individual middens (Spaulding et al. 1990). The foraging range of a packrat has a radius of about 100 to 160 ft (30 to 50 m) (Vaughan 1990), so their middens are local records. This is where a long-term midden deposit is quite useful: as conditions change and different plants become more or less abundant, enter an area, or disappear from it, the changes will be recorded by generations of rats.

Rat-tested, rat-approved (a photo from Nevada borrowed from Wikipedia, originally from a no-longer accessible USGS page).

Packrat middens are common, and we have reports of them being encountered (although not necessarily understood) since the mid-1800s, one of the most entertaining and stomach-turning being a party of '49ers running across some in the Papoose Lake area of Nevada. They encountered an odd glossy material resembling candy in niches in a cliff, some of them were feeling peckish... Nobody died, fortunately. True scientific investigation (as opposed to spitballing about their origin) did not begin until the 1960s, Wells and Jorgensen (1964) being the key early reference. So, what can we learn from this stuff?

Packrat middens are excellent sources of pollen, plant fragments, body fossils of arthropods (insects, spiders, millipedes, etc.) and (typically) small vertebrates, and fossil rodent droppings (coprolites). They span at least 40,000 to 50,000 years and are particularly well-represented in the deserts from western Texas to eastern California, areas that are lacking in other sources of paleoecological fossils like lakes and marshes. By far the most popular aspect of fossil middens has been their plant fossils. Researchers reconstruct changes in plant assemblages over time, which can be used as a proxy for climate. Many other topics have been studied, though. To borrow from a National Park Service inventory from a couple of years ago with a familiar lead author (Tweet et al. 2012):
"Studies following the distribution of a particular plant taxon through time are common (Lanner, 1974; Wells and Hunziker, 1976; Spaulding and Van Devender, 1977, 1980; Van Devender and Hawksworth, 1986; Van Devender et al., 1990; Lanner and Van Devender, 1998; Hunter et al., 2001). Various categories of fossils outside of plant macrofossils have also been studied, including pollen (Thompson, 1985; Davis and Anderson, 1987), arthropods (Hall et al., 1989, 1990; Elias and Van Devender, 1990, 1992; Elias et al., 1992), small vertebrates (Mead et al., 1983, 2003; Van Devender and Bradley, 1990; Van Devender et al., 1991), and rodent coprolites, which have been used to study rodent size and response to climate (Smith and Betancourt, 1998). At least one taxon has been named from midden fossils (the rabbitbrush Chrysothamnus pulchelloides, from a midden at Chaco Culture National Historical Park; Anderson, 1980). Middens have been used to study the rate of erosion in the Grand Canyon (Cole and Mayer, 1982), the extinction of ground sloths (Phillips, 1984), the disappearance of people from Chaco Canyon (Betancourt and Van Devender, 1981; Betancourt, 1990) and to provide points of comparison for resource management at Capitol Reef National Park (Cole and Henderson, 1993; Cole et al., 1997; Cole and Murray, 1999) and Glen Canyon National Recreation Area (Fisher et al., 2006, 2009). Middens have also been incorporated into cultural resource studies (Emslie et al., 1995)."
Not bad for heaps of plant matter and other things cemented by rat pee.

Distribution of midden sites from the old USGS-NOAA midden database (no longer functioning).

References:

Anderson, L. C. 1980. A new species of fossil Chrysothamnus (Asteraceae) from New Mexico. Great Basin Naturalist 40:351–352.

Betancourt, J. L. 1990. Late Quaternary biogeography of the Colorado Plateau. Pages 259–292 in Betancourt, J. L., T. R. Van Devender, and P. S. Martin, editors. Packrat middens: the last 40,000 years of biotic change. University of Arizona Press, Tucson, AZ.

Betancourt, J. L., and T. R. Van Devender. 1981. Holocene vegetation in Chaco Canyon, New Mexico. Science 214:656–658.

Cole, K. L., and N. R. Henderson. 1993. The presettlement vegetation of Capitol Reef National Park reconstructed with fossil packrat middens. Page 33 in Santucci, V. L., editor. National Park Service Research Abstract Volume. NPS Geological Resources Division, Denver, CO. Technical Report NPS/NRPO/NRTR-93/11.

Cole, K. L., and L. Mayer. 1982. Use of packrat middens to determine rates of cliff retreat in the eastern Grand Canyon, Arizona. Geology 10:597–599.

Cole, K. L., and L. K. Murray. 1999. Middle and late Holocene packrat middens from Capitol Reef National Park. Pages 3–24 in Riper, C. v., III and M. A. Stuart, editors. Proceedings of the Fourth Biennial Conference of Research on the Colorado Plateau. U.S. Geological Survey, Reston, VA. Report Series USGSFRESC/COPL/1999/16.

Cole, K. L., N. Henderson, and D. S. Shafer. 1997. Holocene vegetation and historic grazing impacts at Capitol Reef National Park reconstructed using packrat middens. Great Basin Naturalist 57:315–326.

Davis, O. K., and R. S. Anderson. 1987. Pollen in packrat (Neotoma) middens: pollen transport and the relationship of pollen to vegetation. Palynology 11:185–198.

Elias, S. A., and T. R. Van Devender. 1990. Fossil insect evidence for late Quaternary climatic change in the Big Bend region, Chihuahuan Desert, Texas. Quaternary Research 34:249–261.

Elias, S. A., and T. R. Van Devender. 1992. Insect fossil evidence of late Quaternary environments in the northern Chihuahuan Desert of Texas and New Mexico: comparisons with the paleobotanical record. The Southwestern Naturalist 37:101–116.

Elias, S. A., J. I. Mead, and L. D. Agenbroad. 1992. Late Quaternary arthropods from the Colorado Plateau, Arizona and Utah. Great Basin Naturalist 52:59–67.

Emslie, S. D., J. I. Mead, and L. Coats. 1995. Split–twig figurines in Grand Canyon, Arizona: new discoveries and interpretations. Kiva 61:145–173.

Fisher, J. F., K. L. Cole, and R. S. Anderson. 2006. Using packrat middens to assess how grazing influences vegetation change in Glen Canyon National Recreation Area, Utah. U.S. Geological Survey, Reston, VA. Open File Report 2006-1183.

Fisher, J., K. L. Cole, and R. S. Anderson. 2009. Using packrat middens to assess grazing effects on vegetation change. Journal of Arid Environments 73:937–948.

Hall, W. E., C. A. Olson, and T. R. Van Devender. 1989. Late Quaternary and modern arthropods from the Ajo Mountains of southwestern Arizona. Pan Pacific Entomologist 65:322–347.

Hall, W. E., T. R. Van Devender, and C. A. Olson. 1990. Arthropod history of the Puerto Blanco Mountains, Organ Pipe National Monument, southwestern Arizona Pages 363–379 in Betancourt, J. L., T. R. Van Devender, and P. S. Martin, editors. Packrat middens: the last 40,000 years of biotic change. University of Arizona Press, Tucson, AZ.

Hunter, K. L., J. L. Betancourt, B. R. Riddle, T. R. Van Devender, K. L. Cole, and W. G. Spaulding. 2001. Ploidy race distributions since the Last Glacial Maximum in the North American desert shrub, Larrea tridentata. Global Ecology & Biogeography 10:521–533.

Lanner, R. M. 1974. A new pine from Baja California and the hybrid origin of Pinus quadrifolia. The Southwestern Naturalist 19:75–95.

Lanner, R. M., and T. R. Van Devender. 1998. The recent history of pinyon pines in the American Southwest. Pages 171–182 in Richardson, D. M., editor. Ecology and biogeography of Pinus. Cambridge University Press, Cambridge, United Kingdom.

Mead, J. I., T. R. Van Devender, and K. L. Cole. 1983. Late Quaternary small mammals from Sonoran Desert packrat middens, Arizona and California. Journal of Mammalogy 64:173–180.

Mead, J. I., L. L. Coats, and B. W. Schubert. 2003. Late Pleistocene cave faunas in the eastern Grand Canyon, Arizona. Pages 64–86 in Schubert, B. W., J. I. Mead, and R. W. Graham, editors. Ice Age cave faunas of North America. Indiana University Press, Bloomington and Indianapolis, IN.

Phillips, A. M., III. 1984. Shasta ground sloth extinction: fossil packrat midden evidence from the western Grand Canyon. Pages 148–158 in Martin, P. S., and R. G. Klein, editors. Quaternary extinctions: a prehistoric revolution. University of Arizona Press, Tucson, AZ.

Smith, F. A., and J. L. Betancourt. 1998. Response of bushy-tailed woodrats (Neotoma cinerea) to late Quaternary climatic change in the Colorado Plateau. Quaternary Research 50:1–11.

Spaulding, W. G., and T. R. Van Devender. 1977. Late Pleistocene montane conifers in southeastern Utah. The Southwestern Naturalist 22:269–271.

Spaulding, W. G., and T. R. Van Devender. 1980. Late Pleistocene montane conifers in southeastern Utah. Pages 159–161 in Jennings, J. D., editor. Cowboy Cave. University of Utah, Salt Lake City, UT. Anthropological Papers 104.

Spaulding, W.G., J. L. Betancourt, L. K. Croft, and K. L. Cole. 1990. Packrat middens: their composition and methods of analysis. Pages 59–84 in Betancourt, J. L., T. R. Van Devender, and P. S. Martin, editors. Packrat middens: the last 40,000 years of biotic change. University of Arizona Press, Tucson, AZ.

Thompson, R.S. 1985. Palynology and Neotoma middens. American Association of Stratigraphic Palynologists Contributions Series 16:89–112.

Tweet, J. S., V. L. Santucci, and A. P. Hunt. 2012. An inventory of packrat (Neotoma spp.) middens in National Park Service areas. Pages 355–368 in Hunt, A. P., J. Mil├án, S. G. Lucas, and J. A. Spielmann, editors. Vertebrate coprolites. New Mexico Museum of Natural History and Science, Albuquerque, NM. Bulletin 57.

Van Devender, T. R., and G. L. Bradley. 1990. Late Quaternary mammals from the Chihuahuan Desert: Paleoecology and latitudinal gradients. Pages 360–362 in Betancourt, J. L., T. R. Van Devender, and P. S. Martin, editors. Packrat middens: the last 40,000 years of biotic change. University of Arizona Press, Tucson, AZ.

Van Devender, T. R., and F. G. Hawksworth. 1986. Fossil mistletoes in packrat middens from the southwestern United States. Madrono 33:85–99.

Van Devender, T. R., L. J. Toolin, and T. L. Burgess. 1990. The ecology and paleoecology of grasses in selected Sonoran Desert plant communities. Pages 326–349 in Betancourt, J. L., T. R. Van Devender, and P. S. Martin, editors. Packrat middens: the last 40,000 years of biotic change. University of Arizona Press, Tucson, AZ.

Van Devender, T. R., A. M. Rea, and W. E. Hall. 1991. Faunal analysis of late Quaternary vertebrates from Organ Pipe Cactus National Monument, southwestern Arizona. The Southwestern Naturalist 36:94–106.

Vaughan, T. A., 1990. Ecology of living packrats. Pages 15–27 in Betancourt, J. L., T. R. Van Devender, and P. S. Martin, editors. Packrat middens: the last 40,000 years of biotic change. University of Arizona Press, Tucson, AZ.

Wells, P. V., and J. H. Hunziker. 1976. Origin of the creosote bush (Larrea) deserts of southwestern North America. Annals of the Missouri Botanical Garden 63:843–861.

Wells, P. V., and C. D. Jorgensen. 1964. Pleistocene wood rat middens and climatic change in Mohave desert: a record of juniper woodlands. Science 143:1171–1174.

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