Macrofossil analysis of wood rat (Neotoma) middens as a key to the quaternary vegetational history of arid America

Wood rat (Neotoma) deposits preserved in dry rock shelters have radiocarbon ages extending from close to the present to >40,000 BP, thus providing elaborate samples of changing vegetation during the climatic shifts of the late Pleistocene and Holocene. The established record extends geographically from Oregon and Wyoming (at 45°N) south to Baja California, Sonora, and to Tehuacan (at 18°N) in southern Mexico. Several hundred ancient middens have been uncovered, and over 130 have been radiocarbon-dated.There are about 20 extant species of Neotoma with a combined range extending from the Yukon and New England to Nicaragua and Florida. Hence, a wider application for the method seems likely, wherever dry caves favor preservation. The acquisitive rats accumulate an incredibly detailed inventory of the local flora and fauna within a small home range, measured at about 100 m or less in radius. The biomass spectrum of a modern wood rat deposit was compared with associated pollen spectra and source vegetation. The most dominant arborescent species, as determined by a quantitative study of the local vegetation, were proportionately represented in the midden, but not in the pollen rain. Less dominant species were variable in proportionality.A novel approach to the comparative ecological physiology of long-dead plants has been demonstrated with macrofossils from ancient wood rat deposits. The ratio of the stable isotopes of carbon (¹³C/¹²C) is more altered from the atmospheric proportion during CO₂ fixation by C₃ plants than it is by the different (PEP) carboxylating enzyme of heliophile or xerophytic, C₄ or CAM plants. Mass spectrometry of the carbon of macrofossils enables the distinction to be made in the past.The desiccated, allelochemic urine, or amberat, indurates and preserves the middens by cementing the loose debris of macrofossils into a tough, coherent mass of surprising strength and rigidity, that adheres tenaciously to rock surfaces. Water softens the most indurated middens by dissolving the crystallized urine, causing them to dislodge and to fall apart; it also removes the osmotic and allelochemic deterrent to decay by fungi, or to consumption by termites, crickets, or other decomposing herbivores. A secure position in a dry rock shelter is therefore essential to preservation of the macrofossils for time periods on the order of 10⁴ years. Attrition with time is seen in the frequency distribution of radiocarbon dates obtained on 132 Neotoma middens. The distribution is skewed toward the end of the Wisconsin glacial and the Holocene, and there is a decline in frequency with age. Cave erosion, including destructive rockfalls and new crevices that admit seepage and cause deterioration and dislodgement of the middens, is probably the main factor in the progressive attrition with increasing age.