Life history attributes of the threatened Australian snake (Stephen's banded snake Hoplocephalus stephensii, Elapidae)

Whether or not a species is threatened by habitat change is influenced by its life-history traits as well as by the nature and severity of the threatening process. Detailed studies of declining taxa can clarify reasons for their vulnerability, both in proximate terms (e.g., taxa with slow growth, delayed maturation, low fecundity and infrequent reproduction will be poorly suited to withstand increases in adult mortality rates) and in an evolutionary perspective (i.e., what attributes of the environment have favoured the evolution of certain life history traits?). We obtained data on life history traits during an intensive radio-tracking study on the threatened arboreal elapid snake Hoplocephalus stephensii from forest habitats of eastern Australia. Based on 39 field-collected specimens, these long, slender snakes (mean adult snout-vent length 73 cm, mass 114 g) feed predominantly on mammalian prey such as Bush Rats (Rattus fuscipes) and Pygmy-possums (Cercartetus nanus); but juveniles also take lizards. Foraging modes are diverse, from active searching through to ambush. Gape-limitation prevents snakes from ingesting adults of the most abundant local rodent (R. fuscipes), limiting the snakes to specialise on sub-adult rats which are only seasonally available. Rates of feeding, growth and reproduction are low; for example, none of 21 wild-caught females was reproductive. However, snakes given access to abundant prey in captivity, rapidly gained in body condition, grew significantly, and subsequently reproduced. Litters consisted of a few (1-9), relatively large (25 cm snout-vent length, 7.5 g) offspring. The “slow” life-history traits that have contributed to endangerment of this taxon may include proximate as well as evolutionary responses to the low temperatures and limited prey availability in the forest habitat of H. stephensii.     

Decline of an isolated timber rattlesnake (Crotalus horridus) population: Interactions between climate change, disease, and loss of genetic diversity

Extinction of populations from anthropogenic forces rarely has a single cause. Instead, population declines result from a variety of factors, including habitat loss, inbreeding depression, disease, and climate change. These impacts often have synergistic effects that can lead to rapid decline in isolated populations, but case studies documenting such processes are rare. Here, we describe the recent decline of the last known population of timber rattlesnakes (Crotalus horridus) in the state of New Hampshire. We used polymorphic nuclear DNA markers to compare genetic diversity of this population to other populations in the region that are not isolated. We also compare results from ongoing field monitoring of these populations. Genetic analyses reveal that the New Hampshire population lacks genetic diversity and exhibits signs of a recent bottleneck. New Hampshire snakes also exhibited high levels of morphological abnormalities (unique piebald coloration, amelanistic tongues) indicative of inbreeding depression. Furthermore, after a year with exceptionally high summer rainfall, a skin infection of unknown etiology caused significant mortality in the New Hampshire population, whereas other surveyed non-inbred populations were unaffected. This case study demonstrates how different anthropogenic impacts on natural environments can interact in unexpected ways to drive threatened populations toward extinction.     

Ecology and conservation of an endangered rattlesnake,Sistrurus catenatus,in Missouri,USA

The life history and ecology of the massasauga Sistrurus catenatus, an endangered rattlesnake, was studied from 1979 to 1983 at the Squaw Creek National Wildlife Refuge, Missouri, USA. Except for tail length, this population exhibits little sexual dimorphism. Massasaugas are active from April to October, and are primarily diurnal, except in summer. Snakes are found mainly in a cordgrass prairie in spring and autumn, utilising drier, upland areas in summer. Mean brood size for this population was 6·35, and there was a significant positive relationship between brood size and female body size. Although this species apparently demonstrates considerable geographic variation in reproductive potential, the significance of this variation cannot be currently assessed. Growth rates, estimated from size-frequency data, suggest an age of maturity of 3–4 years for females. Massasaugas at Squaw Creek feed mainly on rodents and other snakes. Current refuge practices, such as controlled burning of the prairie and unrestricted visitor usage, may have significant negative impacts on Sistrurus populations. Recommendations for mitigating these impacts are provided.     

Physiological evidence for reproductive suppression in the introduced population of brown tree snakes (Boiga irregularis) on Guam

Introduced species are an increasingly pervasive problem. While studies on the ecology and behavior of these pests are numerous, there is relatively little known of their physiology, specifically their reproductive and stress physiology. One of the best documented introduced pest species is the brown tree snake, Boiga irregularis, which was introduced onto the Pacific island of Guam sometime around World War II. The snake is responsible for severely reducing Guam's native vertebrates. We captured free-living individuals throughout the year and measured plasma levels of stress and sex hormones in an effort to determine when they were breeding. These data were compared to reproductive cycles from a captive population originally collected from Guam. Free-living individuals had chronically elevated plasma levels of the stress hormone corticosterone and basal levels of sex steroids and a remarkably low proportion were reproductively active. These data coincide with evidence that the wild population may be in decline. Captive snakes, had low plasma levels of corticosterone with males displaying a peak in plasma testosterone levels during breeding. Furthermore, we compared body condition between the free-living and captive snakes from Guam and free-living individuals captured from their native range in Australia. Male and female free-living snakes from Guam exhibited significantly reduced body condition compared to free-living individuals from Australia. We suggest that during the study period, free-living brown tree snakes on Guam were living under stressful conditions, possibly due to overcrowding and overexploitation of food resources, resulting in decreased body condition and suppressed reproduction.     

Snake species distributions and temperate grasslands: A case study from the American tallgrass prairie

Many snake species are declining globally, yet data unavailability in temperate grasslands hinders snake conservation in one of the world’s most endangered biomes. To encourage and inform the conservation of snakes in these regions, I examined snake species diversity and abundance during 2 years of a mark-recapture study at 22 sites located in six American tallgrass prairie preserves in northern Illinois, USA. I emphasized landscape-scale relationships after accounting for covariance with environmental factors at both finer (microhabitat) and broader (regional) scales. A total of 120 snakes representing seven species was captured using drift fence arrays associated with funnel traps and sheet metal cover. The low numbers and diversity of snakes captured, when compared to historic evidence, indicate that since the 1930s Midwestern snake populations have declined. Non-metric multidimensional scaling and Mantel tests demonstrate that differences in snake species composition in remnant and degraded prairies are discernable along gradients of urban and agricultural land cover. However, different patterns of abundance for individual snake species indicate that the relative importance of specific landscape factors differ among species, which has significant conservation implications. Here, I recommend that conservationists use a species-specific approach to manage snake populations. I also stress the need for thorough inventory of snake populations and studies of snake-habitat relationships to advance our understanding of snake ecology and conservation within the little studied temperate grassland habitats of South America and eastern Eurasia.     

Survival rates of sea snakes caught by demersal trawlers in northern and eastern Australia

This paper examines the survival rates of sea snakes caught in research and commercial trawls in the main fishing grounds of northern and eastern Australia. Observers recorded whether the snake was alive or dead when it was landed on deck, and (if possible) its species, its length and weight, the duration of the trawl and the total weight of the catch. To estimate long-term survival, captured sea snakes were kept in sea water in drums on a large research vessel and monitored after 1, 6, 12, and 24 h then daily for up to 4 days. Up to 28% of all sea snakes landed on deck from commercial prawn trawling were found to be dead. Both duration of the trawl and catch weight had significant (P<0.001) positive effects on within-trawl (short-term) mortality of these sea snakes. About 30% of all sea snakes from 30 min duration trawls died during the 4-day survival experiment, with Hydrophis ornatus showing the highest mortality (< 50% alive after 96 h in a tank). Hydrophis elegans appeared to have the lowest mortality rate after capture (94% alive after 96 h). Overall, mortality from research and commercial prawn trawling was estimated as 48.5% of snakes caught. Visible damage to sea snakes by trawling was relatively low (5.4%). The introduction of Bycatch reduction devices, such as square mesh and fisheye windows, should improve the survival of sea snakes by reducing the catch weight and the number of sea snakes being caught.     

Evidence of indirect impacts of introduced trout on native amphibians via facilitation of a shared predator

Hyperpredation occurs when non-native prey facilitate invasive predators, which then suppress native prey. Direct impacts of introduced fish on amphibians are well studied, but the role of fish in supporting shared predators has not been considered. We present evidence for indirect effects of trout on amphibians through snake predation. Analyses of the diet, distribution and density of the Pacific coast aquatic garter snake (Thamnophis atratus) relative to the sympatric common garter snake (Thamnophis sirtalis) in the Klamath Mountains of California suggest that trout introductions facilitated expansion of T. atratus by providing alternative prey. T. atratus diet included trout and amphibians whereas T. sirtalis preyed solely upon amphibians. The distribution and density of T. atratus matched that of introduced trout instead of native amphibians. Populations of T. atratus could reach high densities in the absence of high densities of amphibians. When the snakes opportunistically prey upon amphibians whose numbers are already directly impacted by trout, they can cause significant additional declines. When T. atratus was present in lake basins, native Cascades frogs (Rana cascadae) were rarer than in basins without T. atratus. This case differs from other hyperpredation studies because the two prey species also interact via intraguild predation. Given the worldwide practice of stocking fish into aquatic habitats, it is important to understand the consequences of the practice on food-web structure and ecosystem functioning. Bottom-up impacts of introduced predators should be considered as well as top-down so that managers can incorporate the range of ecosystem-level effects into conservation goals and decisions.     

Beyond the wetland border: Estimating the impact of roads for two species of water snakes

We used models integrating road maps, traffic volume, and snake movements to examine the potential for roads to contribute to mortality in two species of water snakes that differ in their vagility, use of terrestrial habitats, and conservation status. Road networks and traffic volumes typical of three regions in Indiana, USA, may account for mortality of 14-21% of the population per year in the more vagile, terrestrial, and imperiled copperbelly water snake (Nerodia erythrogaster neglecta) but only 3-5% mortality in the more sedentary, aquatic, and common northern water snake (Nerodia sipedon). The majority (>91%) of road crossings and associated mortality are predicted to occur during overland migrations to other wetlands, suggesting roads bisecting travel routes between wetlands may function as mortality sinks. Our models highlight the proportionately greater risk of mortality for the more vagile and imperiled species, N. e. neglecta, and suggest current wetland conservation strategies that focus on the wetland alone are unlikely to adequately protect wetland biodiversity from certain types of anthropogenic habitat modification. What is needed is a landscape approach to wetland conservation that considers not only the quality of wetlands and nearby terrestrial habitats, but also ensures that terrestrial corridors between wetlands remain permeable and offer safe passage for wildlife.     

Fungal colonization as affected by litter depth and decomposition stage of needle litter

The present study was designated to evaluate the relative effects of litter depth and decomposition stage of needles on fungal colonization of needle litter in field experiments. The experiment was carried out in coniferous temperate forests in central Japan. Needle litter of Chamaecyparis obtusa and Pinus pentaphylla var. himekomatsu at two decomposition stages (recently dead and partly decomposed) were placed into the organic layer at two depths (on the surface of and beneath the litter layer). Fungal colonization of needles after 1 year was examined in terms of hyphal abundance and frequency of fungal species. Total and live hyphal length on needles were affected by the litter depth and (or) the decomposition stage of needles. Length of darkly pigmented hyphae on needles was 1.7-2.6 times greater beneath the litter layer than on the litter surface regardless of the decomposition stage of needles. Length of clamp-bearing hyphae in Pinus pentaphylla was 5.0-5.2 times greater in partly decomposed needles than in recently dead needles regardless of the litter depth. Frequencies of Pestalotiopsis spp. and Cladosporium cladosporioides were higher on recently dead needles than on partly decomposed needles and (or) were higher on the litter surface than beneath the litter layer. Frequencies of Trichoderma, Penicillium, and Umbelopsis species generally were higher on partly decomposed needles than on recently dead needles and were higher beneath the litter layer than on the surface.     

Geographic isolation, genetic divergence, and ecological non-exchangeability define ESUs in a threatened sky-island rattlesnake

We utilized nine microsatellite loci and 865 bases from two mtDNA genes to estimate demographic parameters and visualize historic/contemporary connectivity among populations of a sky-island rattlesnake (New Mexico ridge-nosed rattlesnake, Crotalus willardi obscurus). This taxon is listed as threatened under the United States Endangered Species Act (ESA) and is distributed patchily within three borderland mountain ranges [Animas (ANM), Peloncillo (PEL), Sierra San Luis (SSL)] of southeastern Arizona, southwestern New Mexico and north-central México. Molecular data support a hypothesis of northward range expansion from México, with subsequent isolation on sky-islands through vicariant desertification that transformed intervening wooded valleys and low elevation passes into inhospitable habitat. Historic and recent movements have been within rather than among mountains. Although populations are linked via ancestral polymorphism, they do not represent a single mtDNA gene pool. All are genetically bottlenecked, with the Peloncillo reflecting deepest/sharpest declines and fewest captures per unit effort. Most recent population declines occurred ∼4 kybp (thousands of years before present). Thus, population reductions are historic and environmental rather than contemporary and anthropogenic. Our data demonstrate that PEL is ecologically non-exchangable with other sky-island populations, and thus comprises one ‘evolutionary significant unit’ (ESU), while SSL and ANM comprise ‘management units’ (MUs) within a second ESU. All three meet legal criteria for recognition as ‘distinct population segments’ (DPS) under the ESA.     

Fungal ingrowth on forest floor and decomposing needle litter of Chamaecyparis obtusa in relation to resource availability and moisture condition

Fungi play an important role in litter decomposition in forest ecosystems and studies are needed to follow the changes in hyphal abundance during litter decomposition and examine the factors regulating the ingrowth of hyphae in litter. The purposes of the present study are to demonstrate the patterns of needle decomposition of Chamaecyparis obtusa in terms of the vertical distribution of fungal biomass and chemical properties within litter horizons (L1, L2, F, and H layers) and fungal ingrowth and succession in relation to organic chemical and nitrogen dynamics during needle litter decomposition over a one-year period. A further aim is to assess the effect of moisture and availability of organic matter on live hyphal length, during 1 year of decomposition. Live hyphal length was correlated to holocellulose concentration in four litter horizons. In a litter bag experiment, total (live plus dead) hyphal length increased during decomposition which was correlated to the concentrations of nitrogen, lignin, holocellulose, and soluble carbohydrate in the litter. The 12-month period over which decomposition was measured was divided into four seasons and the correlation between the water content and live hyphal length was evaluated for each period. The length of live hyphae was correlated to the water content of litter in all four periods. The slopes of regression lines between the water content and live hyphal length were positively correlated to the mean concentrations of soluble carbohydrate of each period, suggesting that the growth of live hyphae was highly dependent on the moisture condition of litter, and under moist conditions on the availability of soluble carbohydrate in the litter. The decrease in the slopes during decomposition can be ascribed to ecophysiological traits of fungi responsible for decomposition in these periods.     

A set of nonlinear regression models for animal movement in response to a single landscape feature

The study of individual animal movement in relation to objects in a landscape is important in many areas of ecology and conservation biology. Yet, many of the models used by ecologists do not account for landscape features and thus may not be conducive to analysis of animal movement data. This article develops a set of nonlinear regression models for both move angles and move distances in relation to a single object in the landscape. Our models incorporate the concept of perceptual range from theories of animal movement behavior. We describe numerical methods for obtaining the maximum likelihood estimates of the model parameters. For illustration, we show results from both computer simulated data and real movement data collected for a red diamond rattlesnake (Crotalus ruber) via radio telemetry field techniques.     

Terrestrial vertebrate communities at black-tailed prairie dog (Cynomys ludovicianus) towns

We capitalized on a regional-scale, anthropogenic experiment—the reduction of black-tailed prairie dog (Cynomys ludovicianus) towns across the Great Plains of North America—to test the hypothesis that decline of this species has led to declines in diversity of native grassland vertebrates of this region. We compared species richness and species composition of non-volant mammals, reptiles and amphibians at 36 prairie dog towns and 36 paired sites in the Panhandle Region of Oklahoma during the summers and falls of 1997, 1998 and 1999. We detected 30 species of mammals, 18 species of reptiles and seven species of amphibians. Comparisons between communities at prairie dog towns and paired sites in the adjacent landscape indicated that while richness per se was not necessarily higher in towns, they did harbor significantly more rare and imperiled species. Species that were positively associated with prairie dog towns during one or both seasons (summer and fall) included badgers (Taxidea taxus), eastern cottontails (Sylvilagus floridanus), coyotes (Canis latrans), grasshopper mice (Onychomys leucogaster), swift fox (Vulpes velox), pronghorn antelopes (Antilocapra americana), striped skunks (Mephitis mephitis), white-tailed deer (Odocoileus virginianus), cattle, thirteen-lined ground squirrels (Spermophilus tridecemlineatus), black-tailed jackrabbits (Lepus califonicus), barred tiger salamanders (Ambystoma tigrinum), plains spadefoot toads (Scaphiopus bombifrons), Great Plains toad (Bufo cognatus), Woodhouse's toad (Bufo woodhousii), prairie rattlesnakes (Crotalis viridis), western plains garter snakes (Thamnophis radix), Texas horned lizards (Phrynosoma cornutum), and ornate box turtles (Terrapene ornata).     

Seasonal variation in nitrogen isotopic composition of bog plant litter during 3 years of field decomposition

In this study, we describe the seasonal variation in ¹⁵N abundance in the litter of two Sphagnum species and four vascular plant species during 3 years of field decomposition in an Italian Alpine bog. Litter bags were periodically retrieved at the end of summer and winter periods, and the δ¹⁵N in residual litter was related to mass loss, litter chemistry, and climatic conditions. In Sphagnum litter, higher rates of decomposition during summer months were associated with an increase of δ¹⁵N probably due to the incorporation of microbial organic compounds rich in ¹⁵N. The litter of Eriophorum vaginatum and Carex rostrata was characterized by a decrease of δ¹⁵N, so that the final signature was significantly lower than in initial litter. On the other hand, the residual litter of Potentilla erecta and Calluna vulgaris was characterized by a final δ¹⁵N higher than in initial litter. Our data reported a seasonality of ¹⁵N abundance in the residual litter of Sphagnum species, but not in that of vascular plant species, thus highlighting the role of differences in litter chemistry.     

Potential causes of population declines in forest fragments in an Amazonian frog

Forest fragmentation results in population declines and extinctions for many forest vertebrates, but little is known about the mechanisms causing declines in fragments. We investigated potential causes of declines in forest fragments for an Amazonian forest frog (Colostethus stepheni) at an experimental fragmentation study site in central Amazonian Brazil using field estimates of abundance and vital rates coupled with population simulations. Although adult male survival was not reduced by fragmentation, mean clutch size was reduced by 17%. Population simulations demonstrate that a reduction in clutch size of this magnitude is sufficient to cause the observed magnitude of population declines in fragments. Female snout-vent length was also reduced in fragments and may be related to the observed reduction in clutch size.     

Palatability trials on hardwood leaf litter grown under elevated CO2: a stable carbon isotope study

The palatability to isopods and microbes of a broad range of hardwood leaf litter, derived from three field CO₂-enrichment experiments in the USA, was investigated, using δ¹³C, to trace the C flow from litter to isopods and to CO₂ respired by microbial decomposition. Leaf litter grown under elevated CO₂ had δ¹³C values ranging from −39 to −45‰, which were significantly different from ambient litter δ¹³C values of around −30‰. Litter palatability to isopods of the Porcellio sp. was tested by incubating ambient- and elevated-CO₂ litter, and a mixture of the two, in the presence of isopods for 14 days, under environmentally controlled conditions; δ¹³C was measured on litter and isopods' body before and after incubation. In an additional experiment, litter was incubated in the absence of fauna for 30 days, and on five occasions the δ¹³C of the CO₂ respired from litter was measured. The ¹³C label was clearly carried from the litter source to the isopods' bodies, and their faeces. For microbial-respired CO₂, δ¹³C was significantly higher than that of the litter source, suggesting preferential degradation of substrates enriched in ¹³C as compared to those in the overall litter. With the exception of Quercus myrtifolia leaf litter, elevated CO₂ did not affect the palatability to isopods nor the microbial degradation of any of the litters, possibly as a result of unaltered litter N concentration. However, significant differences in litter palatability and decay rates were observed among the different species. With this study, the use of isotopically labelled litter material was confirmed as a key methodology that can significantly contribute to the advancement of the understanding of litter decomposition and of the quantification of C fluxes in the process.     

Decomposing ability of interior and surface fungal colonizers of beech leaves with reference to lignin decomposition

Fungi were isolated from interior and surface of beech (Fagus crenata Blume) leaf litter by surface sterilization and washing methods. Species composition differed between the interior and surface of the leaves. Xylaria sp. (anamorph) was a major interior colonizer, while Pestalotiopsis spp. and Trichoderma spp. were predominantly surface colonizers. Ascochyta sp. was isolated from both the interior and surface of leaves. Leaf litter decomposing abilities of all isolated species were assessed by pureculture decomposition tests. Changes in lignin, carbohydrate and polyphenol amounts in the litter were investigated. Percent loss of original weight of sterilized beech leaf litter ranged from 0.24 to 11.16 % in the decomposition test. Interior fungi such as Xylaria sp. (anamorph) had the highest abilities to bleach leaf litter and decompose lignin. Most surface fungi had limited ability to decompose leaf litter and lignin. The difference in the decomposing abilities between the interior and surface fungi is discussed in relation to the difference in organic-chemical compositions between interior and surface of the litter.     

A test of the hierarchical model of habitat selection using eastern massasauga rattlesnakes (Sistrurus c. catenatus)

Understanding the process animals follow to select habitat, rather than just documenting the habitat they use, will improve our ability to predict how the animals use habitat in other locations and how they will respond to changes in habitat. Animals are usually assumed to select habitats hierarchically, preferentially using specific macrohabitats at a landscape scale and specific microhabitats within the preferred macrohabitats. We used four years of telemetry data from 34 individuals to test this hierarchical model of habitat selection with eastern massasauga rattlesnakes (Sistrurus c. catenatus) in Ontario. Snakes were selective at the microhabitat scale, preferentially using locations with closer retreat sites and shrubs than random. Gravid females were most selective, using sites with more rock cover and less canopy closure than sites used by males and nongravid females. Snakes preferred forested habitats for hibernation and steadily increased their use of open, wetland, and edge habitats to a peak in mid-summer. Landscape-scale habitat preferences were generally mild and could be explained by the relative availability of suitable microhabitat within habitats, suggesting habitat selection was primarily driven by microhabitat preferences. The lack of selectivity at the landscape scale may be a consequence of fine-grained differences between habitats that allow massasauga rattlesnakes to find suitable microhabitats in all available macrohabitats. For species that select habitat primarily at the microhabitat scale (e.g., the rattlesnakes we studied), landscape-scale modeling of habitat use will only be effective to the extent habitats reflect the availability of suitable microhabitat within.     

Photodegradation of plant litter in the Sonoran Desert varies by litter type and age

Predicting litter decay rates in arid systems has proved elusive and sunlight (photodegradation) is a potentially important but poorly understood driver of litter decay in these systems. We placed three litter types (Cynodon dactylon, Larrea tridentata leaves, and L. tridentata twigs) in envelopes whose tops either transmitted all solar radiation, filtered UV-B, filtered all UV, or filtered all UV and visible solar radiation, on the soil surface of the Sonoran Desert and assessed mass loss over 14 months. Regardless of treatment, final mass loss was greatest in C. dactylon litter and least in L. tridentata twig litter, consistent with initial litter characteristics of presumed litter quality; C. dactylon had the lowest lignin concentration and lignin:N, and the highest cellulose:lignin and area:mass. Compared to litter in sunlight, excluding solar UV, or UV-B, slowed mass loss of all 3 litter types, and UV-B appeared more effective than UV-A in photodegradation. The relative contribution of UV photodegradation to mass loss increased with litter age. After 14 months, litter exposed to solar UV lost 1.2 (C. dactylon), 1.3 (L. tridentata twigs) and 1.4 (L. tridentata leaves) times as much mass as litter not exposed to UV radiation. The relative contribution of UV photodegradation to mass loss increased with the initial C:N ratio of litter, but was not related to initial lignin concentration or optical properties (i.e. UV and visible absorbance and transmittance) of litter. Within all litter type by treatment combinations, there was a strong positive correlation between litter mass loss and ash concentration. In some cases, a discontinuity in this relationship was detected, suggesting a threshold ash concentration, above which further mass loss was negligible. We expected these thresholds to be most prevalent in sunlight, because soil films could prevent sunlight from reaching litter and thereby minimize photodegradation. Contrary to expectations, thresholds were more common in shade or UV filter treatments, suggesting that reductions in photodegradation attributable to soil films were not typically responsible. The effect of shading, which likely enhanced microbial degradation via higher relative humidity due to lower temperatures, depended on litter type and time. Compared to litter in sunlight, mass loss of shaded litter was greater over the initial 3 months in all litter types, illustrating that microbial degradation in shade was greater than photodegradation in sunlight. These differences in mass loss between shaded and sunlit litter increased over the 14 month experiment in L. tridentata twigs, declined in L. tridentata leaves, and disappeared within 6 months in C. dactylon, illustrating that the timing of this shift in the dominance of photodegradation versus microbial degradation was highly dependent on litter type. In a second experiment, we reduced microclimate differences among sun and shade treatments, pre-sterilized litter to reduce microbial degradation, and examined the mass loss of young and old and L. tridentata leaf litter after 53 days outdoors. Consistent with our first experiment, mass loss attributable to photodegradation was greater in old than young litter. Unsterilized litter exposed to sunlight (UV and visible) lost 1.3 (young) and 1.5 (old) times as much mass as shaded litter. Pre-sterilized litter exposed to sunlight lost 11.4 (young litter) and 45.9 (old litter) times as much mass as shaded litter. These large differences in pre-sterilized litter were the result of the very small mass loss of shaded litter (≤0.2%), together with modest losses of sunlit litter (<5%). Taken together, our findings suggest that as litter aged, microbial degradation became a weaker driver of mass loss, while photodegradation became stronger.