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Historically, ecosystem models have treated rainfall as the primary moisture source driving litter decomposition. In many arid and semi-arid lands, however, non-rainfall moisture (fog, dew, and water vapor) plays a more important role in supporting microbial activity and carbon turnover. To date though, we lack a robust approach for modeling the role of non-rainfall moisture in litter decomposition. We developed a series of simple litter decay models with different moisture sensitivity and temperature sensitivity functions to explicitly represent the role of non-rainfall moisture in the litter decay process. To evaluate model performance, we conducted a 30-month litter decomposition study at 6 sites along a fog and dew gradient in the Namib desert, spanning almost an eightfold difference in non-rainfall moisture frequency. Litter decay rates in the field correlated with fog and dew frequencies but not with rainfall. Including either temperature or non-rainfall moisture sensitivity functions improved model performance, but the combination of temperature and moisture sensitivity together provided more realistic estimates of litter decomposition than relying on either alone. Model performance was similar regardless of whether we used continuous moisture sensitivity functions based on relative humidity or a simple binary function based on the presence of moisture, although a Gaussian temperature sensitivity outperformed a monotonically increasing Q10 temperature function. We demonstrate that explicitly modeling non-rainfall moisture and temperature together is necessary to accurately capture litter decay dynamics in a fog-affected dryland system and provide suggestions for how to incorporate non-rainfall moisture into existing Earth system models.

Non-rainfall moisture (fog, dew, and water vapor; NRM) is an important driver of plant litter decomposition in grasslands, where it can contribute significantly to terrestrial carbon cycling. However, we still do not know whether microbial decomposers respond differently to NRM and rain, nor whether this response affects litter decomposition rates. To determine how local moisture regimes influence decomposer communities and their function, we examined fungal communities on standing grass litter at an NRM-dominated site and a rain-dominated site 75 km apart in the hyper-arid Namib Desert using a reciprocal transplant design. Dominant taxa at both sites consisted of both extremophilic and cosmopolitan species. Fungal communities differed between the two moisture regimes with environment having a considerably stronger effect on community composition than did stage of decomposition. Community composition was influenced by the availability of air-derived spores at each site and by specialization of fungi to their home environment; specifically, fungi from the cooler, moister NRM Site performed worse (measured as fungal biomass and litter mass loss) when moved to the warmer, drier rain-dominated site while Rain Site fungi performed equally well in both environments. Our results contribute to growing literature demonstrating that as climate change alters the frequency, magnitude and type of moisture events in arid ecosystems, litter decomposition rates may be altered and constrained by the composition of existing decomposer communities.

Models assume that rainfall is the major source of moisture driving decomposition. Non-rainfall moisture (NRM: high humidity, dew, and fog) can also induce standing litter decomposition, but there have been few standard measurements of NRM-mediated decompositions across sites, and no efforts to extrapolate the contribution of NRM to larger scales to assess whether this mechanism can improve model predictions. Here we show that NRM is an important, year-round source of moisture in grassland sites with contrasting moisture regimes using field measurements and modeling. We first characterized NRM frequency and measured NRM-mediated decomposition in sites on the extreme dry and wet end of grassland systems: at two sites in the Namib Desert, Namibia (hyperarid desert) and at one site in Iowa, USA (tallgrass prairie). NRM was frequent at all sites (85-99% of hours that litter was likely to be wet were attributed to NRM) and tended to occur in cool, high-humidity periods for several hours or more at a time. NRM also caused respiration of standing litter at all sites when litter became sufficiently wet (>5% for fine litter and >13% for coarse), and contributed to mass loss, even in the Namib West site that had almost no rain. When we modeled annual mass loss induced by NRM and rain, and extrapolated our characterization of NRM decomposition to a final site with intermediate rainfall (Sevilleta, New Mexico, semiarid grassland), we found that models driven by rainfall alone underestimated mass loss, while including NRM produced estimates within the range of observed mass loss. Together these findings suggest that NRM is an important missing component in quantitative and conceptual models of litter decomposition, but there is nuance involved in modeling NRM at larger scales. Specifically, temperature and physical features of the substrate emerge as factors that affect the common microbial response to litter wetting under NRM across grasslands sites, and require further study. Hourly humidity can provide an adequate proxy of NRM frequency, but site-specific calibration with litter wetness is needed to accurately attribute decomposition to periods when NRM wets litter. Greater recognition of NRM-driven decomposition and its interaction with other processes (e.g. photodegradation) is needed, especially since fog, dew, and humidity are likely to shift under future climates.Manuscript highlightsNon-rainfall moisture (NRM; humidity, fog, dew) induces decomposition in grasslandsNRM decomposition depends on substrate type, and occurs at colder times than rainIncluding NRM (instead of rain alone) improved predictions of litter decomposition

Welwitschia mirabilis is an ancient, endemic gymnosperm found in numerous disjunct populations in western Namibia north of the Namib Sand Sea to the Bentiaba River in southwestern Angola. Based on six plants grown in the Berlin-Dahlem Botanical Garden, Leuenberger [2001, Welwitschia mirabilis (Welwitschiaceae), male cone characters and a new subspecies. Willdenowia 31: 357–381] differentiated two groups of plants using eight male inflorescence characters. He concluded that the two groups represented geographical subspecies: W. mirabilis subsp. mirabilis in Angola and W. mirabilis subsp. namibiana in Namibia, based on putative location data of the seed sources for the plants. Leuenberger called for field data collection, especially in northern Namibia, to verify his results. However despite this lack of data, numerous authors have cited his work as definitive in scientific papers and online plant descriptions and keys. We analysed five distinguishing male cone characters from 13 Namibian plant populations ...

For the past fourteen years, two Grinnell College graduates have been selected each year for a competitive one-year Grinnell Corps Service Fellowship at the Gobabeb Research and Training Centre in Namibia. A recent survey of the 28 Fellows that completed the program revealed that this is an invaluable experience for the participants, with more than half revealing that it was “life-changing” and a further 43% reporting that it re-affirmed their life goals. Most Fellows felt that their most valuable contribution to Gobabeb was teaching and mentoring Namibian secondary and tertiary students. All Fellows are now either in graduate school or employed in a diverse array of positions reflecting their professional interests that in some fashion “support the common good”, an integral part of Grinnell's mission statement. Synergistic activities that have arisen from this Grinnell–Gobabeb relationship include visiting lecturer opportunities at Grinnell for Namibians; internships for Grinnell students and service fel...

Ephemeral rivers have been largely excluded from previous attempts to classify global hydrologic regimes, or to assess the role of hydrologic characteristics in regulating ecological processes and patterns in fluvial ecosystems. The Namib Desert's ephemeral rivers are amongst the most hydrologically variable fluvial systems yet described. The coefficient of variation for mean annual runoff (CVMAR) among 28 stations, representing 7 Namib rivers, averages 1.55, compared with a global mean of ∼0.45. Distinct curvilinear relationships exist among hydrologic characters and longitudinal position along the rivers. In particular, peak discharge, annual flow volume, and days of flow per annum exhibit a marked decline in the lower reaches of the rivers, after a mid-catchment peak. These longitudinal gradients exert strong controls over the composition of vegetation, invertebrate, and fungal communities; the availability and structure of various micro-habitats; and the rates of ecological processes such as decomposition. Flood pulses, although variable in their timing and magnitude, play a critical role in regulating organic matter transport and deposition, and primary and secondary production. Despite the tolerance of the biota to harsh and variable abiotic conditions, these ecosystems are highly sensitive to hydrologic alterations as water is acutely limiting for many organisms and ecological processes. Accordingly, alterations to flow regimes can quickly degrade ecological integrity. Managers must seek to maintain existing hydrologic regimes, a challenge amplified by increasing human water demand and changing regional climates.

Welwitschia mirabilis is a monotypic member of the family Welwitchiaceae which, along with Ephedra and Gnetum species, comprises the gymnospermous order Gnetales. While the monophyly of this order is now widely accepted, the relationship of the Gnetales to other seed plants is still contentious. Despite the unique phylogenetic position of W. mirabilis and its extraordinary physiological and anatomical adaptations, little is known about the plant's phylogeny or its current distribution in isolated locations throughout the Namib Desert. As a preliminary step in the design of an more extensive phylogeographic study, we analyzed 37 random amplified polymorphic DNA (RAPD) loci from 59 plants distributed among five sites separated by distances of 6-440 km. Cluster analysis and analysis of molecular variance (AMOVA) revealed significant levels of variation within and between populations and little evidence of inbreeding. Genetic differences between populations reflect the geographic distances separating them. Three of the populations formed discernable genetic clusters, suggesting that little gene flow occurs between populations separated byor = 18 km. In contrast, gene flow is occurring between two populations separated by only 6 km, supporting previous observations that pollen dispersal is primarily local and that seeds are not readily wind-born over the large distances separating most W. mirabilis populations. As a working hypothesis, we propose that W. mirabilis had a continuous distribution across its current range as much as 105 million years ago, and that as a consequence of subsequent drying trends and physical disturbance, populations became progressively isolated, accounting for their current distribution.

The hyper-arid western Namib Sand Sea (mean annual rainfall 0-17 mm) is a detritus-based ecosystem in which primary production is driven by large, but infrequent rainfall events. A diverse Namib detritivore community is sustained by minimal moisture inputs from rain and fog. The decomposition of plant material in the Namib Sand Sea (NSS) has long been assumed to be the province of these detritivores, with beetles and termites alone accounting for the majority of litter losses. We have found that a mesophilic Ascomycete community, which responds within minutes to moisture availability, is present on litter of the perennial Namib dune grass Stipagrostis sabulicola. Important fungal traits that allow survival and decomposition in this hyper-arid environment with intense desiccation, temperature and UV radiation stress are darkly-pigmented hyphae, a thermal range that includes the relatively low temperature experienced during fog and dew, and an ability to survive daily thermal and desiccation stress at temperatures as high as 50°C for five hours. While rainfall is very limited in this area, fog and high humidity provide regular periods (≥ 1 hour) of sufficient moisture that can wet substrates and hence allow fungal growth on average every 3 days. Furthermore, these fungi reduce the C/N ratio of the litter by a factor of two and thus detritivores, like the termite Psammotermes allocerus, favor fungal-infected litter parts. Our studies show that despite the hyper-aridity of the NSS, fungi are a key component of energy flow and biogeochemical cycling that should be accounted for in models addressing how the NSS ecosystem will respond to projected climate changes which may alter precipitation, dew and fog regimes.

Summary 1 The chemical characteristics of floodwaters in ephemeral rivers are little known, particularly with regard to their organic loads. These rivers typically exhibit a pronounced downstream hydrological decay but few studies have documented its effect on chemical characteristics and material transport. To develop a better understanding of the dynamics of floods and associated material transport in large ephemeral rivers, floods of the ephemeral Kuiseb River in south-western Africa were tracked and repeatedly sampled at multiple points along the river's lower 220 km. 2 We quantified the composition and transport of solute and sediment loads in relation to longitudinal hydrological patterns associated with downstream hydrological decay. Source and sink areas for transported materials were identified, and the composition and transport dynamics of the organic matter load were compared to those described from more mesic systems. 3 Concentrations of sediments and solutes transported by floods in the Kuiseb River tended to increase downstream in association with pronounced hydrological decay. The contribution of particulate organic matter to total organic load is among the highest recorded, despite our observation of unusually high levels of dissolved organic matter. Hydrological decay resulted in deposition of all transported material within the lower Kuiseb River, with no discharge of water or materials to the Atlantic Ocean. 4 Our results suggest that longitudinal variation in surface flow and associated patterns of material transport renders the lower Kuiseb River a sink for materials transported from upstream. The downstream transport and deposition of large amounts of labile organic matter provides an important carbon supplement to heterotrophic communities within the river's lower reaches.

Soils were examined along three ephemeral rivers in the Namib Desert to assess the influence of their hydrologic characteristics on soil properties. Soils consisted of layers of fluvially deposited, organic-rich silts, interstratified with fluvial and aeolian sands. The most significant influence of the ephemeral hydrologic regime upon soils was related to the downstream alluviation associated with hydrologic decay. This alluviation increased the silt proportion of soils in the lower reaches of the rivers. Organic carbon, nitrogen and phosphorous were correlated with silt content, and silt deposition patterns influenced patterns of moisture availability and plant rooting, creating and maintaining micro-habitats for various organisms. Localized salinization occurred in association with wetland sites and soluble salt content tended to increase downstream. Because of the covariance between silt and macronutrients, and the influence of silt upon moisture availability and habitat suitability, alluviation patterns associated with the hydrologic regime strongly influence the structure, productivity, and spatial distribution of biotic communities in ephemeral river ecosystems.

The spatiotemporal patterns and ecological significance of the retention of coarse particulate organic matter and large woody debris have been intensively studied in perennial rivers and streams but are virtually unknown in ephemeral systems. We examined the influence of 2 features characteristic of ephemeral systems, downstream hydrologic decay and in-channel tree growth, on the distribution, transport, and retention of woody debris following a flood having a ∼2.6-y recurrence interval in the ephemeral Kuiseb River in southwestern Africa. A total of 2105 pieces of wood were painted at 8 sites along the river channel to measure retention patterns. The flood had a peak discharge of 159 m 3 /s at the upper end of the study area, decaying to 3 /s by 200 km downstream. Downstream export of wood from marking sites totaled 59.5% (n = 1253). Transport distances ranged from 1 to 124 km, and 34.8% (n = 436) of the exported wood was recovered. Marked wood retained within marking sites was sig...

In September 1990, 74 sporocarps of Battarrea stevenii were observed on the floodplain of the ephemeral Kuiseb River in western Namibia. Herein we report subsequent studies of the distribution, abundance, nutritional role, phenology, and sporocarp development of this fungus in the hyper-arid Namib Desert. Included are full descriptions of developing and mature sporocarps. B. stevenii is a common associate of riparian forests on silty floodplain terraces, but does not form mycorrhizal associations with the dominant woody species, Faidherbia albida or Tamarix usneoides. Rather, clamped mycelium extends throughout floodplain soils decomposing coarse and fine particulate organic material (4—7% of soil dry weight). Sporocarp production occurs 4.5—12 mo post-flooding in response to soil desiccation at depths of 20—35 cm. The extensive mycelium, duration of vegetative growth post-flooding, and large size and abundance of B. stevenii sporocarps suggest that it is an important component of the subsurface decomposer community in the Namib's ephemeral rivers. Given that the fungus has also been recorded from floodplain soils of Angola, Hungary, and New Mexico (U.S.A.), and is known to have a world-wide distribution, we predict that further biogeographical studies will reveal that B. stevenii is a characteristic element of the riparian biota in dryland rivers, which drain approximately one-third of the earth's land surface.

Because of the aridity of the Namib Sand Sea, it has long been assumed that decomposition of buried plant material was largely independent of rainfall. Losses were attributed to consumption by detritivores that forage year-round. Moisture-limited micro-organisms were reported to occur in low densities in Namib sands, supporting the assumption that rainfall was insignificant in regulating decomposition. Observations of abundant macrofungal fruiting from buried plant material and herbivore dung, following a 12 mm rain, suggested the importance of rain-induced decomposition had been underestimated. We used cellulose substrates to compare material loss during dry periods and following differing amounts of rain. Strips of cotton cloth and filter paper, buried at 10 cm depths at five disjunct locations, were sequentially removed over 10 months. A period of at least 170 days elapsed before rains, ranging from 4–46 mm, fell at all locations. Material loss during the dry period averaged 8·2% (range 0–16·7%), and was attributed to macrodetritivore consumption. In marked contrast, an average of 84·1% of the material (range 64·7–97·2%) was lost following rains greater than 9 mm. Wet substrates were heavily colonized by fungi, and termites and tenebrionid beetle larvae were observed feeding on fungus-colonized substrates. Stepwise regression analysis revealed that rainfall, rather than duration of burial, was the primary factor determining substrate loss in the Namib Sand Sea. Although rain events are infrequent and ensuing periods of moist soil are brief, substrate loss following rains is highly significant relative to that occurring in the absence of rain. In contrast to more mesic deserts, rainfall is an important trigger of decomposition in the Namib Sand Sea where soils are too dry to support significant decomposition, except when episodically moistened by rain.

A selection of soils were sampled in the Central Namib Desert to investigate the importance of landscape and associated plant communities on soil nutrient status. Soils were sampled in an ephemeral river, a wash, and at two dune sites. The soils in the lower landscape positions, the Kuiseb River and an associated wash, had significantly higher nutrient levels than those at the dune sites. These lower landscape positions also had the most stable plant communities, Faidherbia albida and Welwitschia mirabilis, respectively. The two dune soils, which both support perennial herbaceous species, had extremely low nutrient levels. Only the wash showed nutrient enrichment of the soil under plants, as measured by a relative enrichment factor. This community is the most established, with plants being hundreds of years old; this time factor may be crucial for nutrient accumulation by plants in this extremely arid system. The lack of enrichment under the F. albida community suggests that the fluvial inputs and exports, both organic and inorganic, tend to homogenize the nutrient levels within this system. These results indicate that soil nutrient levels in the Namib Desert are closely linked to landscape position and less so with the associated plant communities.

The arid central dune field of the Namib Desert is characterized by a pronounced rainfall gradient across its west–east, 160 km breadth, and a correlated increase in sand stability and grass community complexity. In addition to these macro-gradients, micro-gradients of sand stability and available moisture across each dune slope result in stratified grass communities on the dunes. The effects of abiotic factors and plant associations on the community structure of VA-mycorrhizal fungi in a naturally arid and unstable grassland could thus be investigated. Mycorrhizal fungal communities associated with five grass species were sampled at sites located across the gradients. Diversity and abundance of spores, as well as percent mycorrhizal colonization of plant roots were used to characterize the fungal communities and their plant specificity. Five Glomus species (Glomales) were associated with grasses at all sites, but no plant specificity was observed. Rather, the fungal communities varied in diversity and abundance both within a dune site and across the dune field. Regression analyses showed that spore abundance and colonization levels were significantly affected by abiotic factors. Sand stability affected spore abundance and thus determined the limits of distribution of the fungal community in the dune grassland. In contrast, colonization levels were primarily affected by moisture availability, and fungal growth and spore production following an isolated rain event were closely associated with moisture availability. A rapid and opportunistic growth response to moisture, production of resilient spores in response to declining moisture, and lack of plant symbiont specificity are characteristics which allow mycorrhizal fungal communities to function under hyperarid conditions.

Assessing genetic variation within populations and genetic exchange between populations requires an understanding of the distribution and abundance of individual genotypes within the population. Previous workers have used somatic incompatibility testing to distinguish clones or individuals in natural populations of ectomycorrhizal fungi. However, somatic incompatibility tests performed with isolates of Suillus granulatus from a natural population revealed a lack of transitiveness, which brought into question the validity of this method. Subsequent studies of genetic identity of these isolates, using randomly amplified polymorphic DNA (RAPD) markers, conclusively showed that somatically compatible isolates are not necessarily genetically identical. RAPD marker analysis is more reliable and provides higher resolution of genotype distribution in natural populations than does somatic incompatibility testing. This is of particular importance in populations of organisms such as ectomycorrhizal fungi in which the mating systems are incompletely known.

Mycorrhizal synthesis studies revealed that allopatric populations of Suillus granulatus differ in host specificity. Isolates of Suillus granulatus originated from Nepal (associated with Pinus wallichiana), Korea (Pinus densiflora), and the U.S.A. (Pinus strobus). Functional compatibility of the mycorrhizal association was used to measure host specificity. Useful characters for measuring mycorrhizal compatibility were defined as those which revealed statistical differences between the uninoculated controls and mycorrhizal treatments and which were stable within one population. Useful characters, as defined by these two tests, were shoot wet weight, total wet weight, percentage mycorrhizal short roots, shoot dry weight, total dry weight, and Hartig net penetration. Data for these characters were subsequently used in anova tests to determine whether allopatric populations of Suillus granulatus differ in functional compatibility. Isolates associated with Pinus strobus were most compatible with Pinus strobus and least compatible with the other trees, whereas isolates from Pinus wallichiana and Pinus densiflora showed variable compatibility with all three trees. Phenetic analyses used to determine how similarly the isolates responded with the hosts confirmed these results. We conclude that isolates associated with Pinus strobus are most specific for this host, whereas isolates from the other two trees are not host specific. The implications of these results are discussed in light of future investigations regarding the relatedness of these tree-associated isolates. Key words: Boletaceae, Suillus, Pinus, ectomycorrhizae.

Aspergillus niger is an asexual, haploid fungus which infects the seeds of Namibia's national plant, Welwitschia mirabilis, severely affecting plant viability. We used 31 randomly amplified polymorphic DNA markers to assess genetic variation among 89 A. niger isolates collected from three W. mirabilis populations in the Namib Desert. While all isolates belonged to the same vegetative compatibility group, 84% were unique genotypes, and estimates of genotypic evenness and Simpson's index of diversity approached 1.0 in the three populations. Analysis of molecular variance revealed that 78% of the total variation sampled was among isolates from individual W. mirabilis plants. Lower, but significant, amounts of variation detected among isolates from different plants (12%) and different sites (10%) also indicated some site- and plant-level genetic differentiation. Total gene diversity (H(T) = 0.264) was mostly attributable to diversity within populations (H(S) = 0.217); the relatively low level of genetic differentiation among the sites (G(ST) = 0.141) suggests that gene flow is occurring among the three distant sites. Although sexual reproduction has never been observed in this fungus, parasexuality is a well-known phenomenon in laboratory strains. We thus attribute the high levels of genetic variation to parasexuality and/or wind-facilitated gene flow from an as of yet undocumented broader host range of the fungus on other desert vegetation. Given the apparent ease of transmission, high levels of genetic diversity, and potentially broad host range, A. niger infections of W. mirabilis may be extremely difficult to control or prevent.