PHOTOPERIODIC ADAPTATION IN DESERT POPULATIONS OF XANTHIUM STRUMARIUM

Reproductive adaptation to photoperiod is diverse among desert populations of Xanthium. Chihuahuan Desert populations require dark periods of 9.5–10.5 hr for reproduction, and Sonoran Desert populations require 9–10.5 hr. Many Chihuahuan populations from western Texas two weeks from sowing need only 10 cycles of 11-hr nights to produce 100% flowering, but Sonoran populations from western Mexico four weeks from sowing need 18 cycles or more. Some Sonoran plants produce buds only at a cooler temperature program, 24–15 C, but Chihuahuan plants produce them more readily under the warmer program, 30–24 C. Chihuahuan plants that were germinated under 11-hr nights and four different temperature programs were induced to flower in each condition. Differences in photoperiod and ripeness-to-flower (maturity) responses were also demonstrated under natural day lengths in central Texas. Although desert populations occurring at approximately the same latitude in either the Chihuahuan or Sonoran Desert are exposed to similar day lengths, each population may be adapted to different photoperiod cues that maximize its utilization of the local growing conditions.     

ECOTYPIC VARIATION IN LARREA DIVARICATA

Data on rate of germination, rate of growth, and temperature and moisture responses of croosotebush (Larrea divaricata Cav.) seedlings from the Sonoran Desert and Chihuahuan Desert suggest that the species as it occurs in North America is comprised of at least two major genetic population systems or ecotypes. Compared to the Chihuahuan Desert ecotype, that of the Sonoran Desert is characterized by a taller, more erect and more open growth form, more slender and less incurved leaflets, slower germination rate, slower initial growth rate of the seedling, less tolerance for low temperature, and greater tolerance for low moisture.     

A phytogeographical characterization of the vine flora of the Sonoran and Chihuahuan deserts

Aim This study presents a phytogeographical characterization of the vine flora of two lower North American desert regions as a biogeographical framework for further ecological inquiry into desert vines. Location The phytogeography of the vine flora of the Sonoran and Chihuahuan Deserts was c haracterized based on 263 known species. Methods Checklists of the vines of each desert were developed. Represented genera were then grouped into 10 phytogeographical elements based on worldwide distribution patterns. To compare the floristic composition of the desert floras, an index of species similarity was calculated. Results About a third more species of vines occur in the Sonoran Desert than in the Chihuahuan Desert. Based on the analysis, cosmopolitan genera are the only group more numerous in absolute terms in the Chihuahuan Desert than in the Sonoran Desert. Tropical elements are represented in about the same proportion in each desert as the number of species, however, nearly twice as many pantropical and neotropical genera are represented in the Sonoran Desert as in the Chihuahuan Desert. Proportionately, more genera of temperate elements occur in the Chihuahuan Desert than in the Sonoran desert, although the absolute number of genera is slightly higher in the latter. Main conclusions As these deserts are relatively recent ecological formations and as vines evolved in forest ecosystems, the composition of the desert vine floras is the result of the interaction between historical vegetation types, their constituent taxa and climatic and geological history. The main differences in the vining floras of the present‐day Sonoran and Chihuahuan Deserts appear to be the result of greater historical influence in the Sonoran Desert of (1) tropical vegetation types and (2) the emergence of the Gulf of California. The Chihuahuan Desert vine flora seems to be the result of (1) a more pronounced historical temperate vegetation, (2) the lack of an important isolating event, such as the creation of the Baja California peninsula, and (3) a cooler climate with shorter growing seasons.     

Effects of plant size and weather on the flowering phenology of the organ pipe cactus (Stenocereus thurberi)

Background and Aims

Flowering phenology is a critical life-history trait that influences reproductive success. It has been shown that genetic, climatic and other factors such as plant size affect the timing of flowering and its duration. The spatial and temporal variation in the reproductive phenology of the columnar cactus Stenocereus thurberi and its association with plant size and environmental cues was studied.

Methods

Flowering was monitored during 3 years in three populations of S. thurberi along a latitudinal gradient. Plant size was related to phenological parameters. The actual and past weather were used for each site and year to investigate the environmental correlates of flowering.

Key Results

There was significant variation in the timing of flowering within and among populations. Flowering lasted 4 months in the southern population and only 2 months in the northern population. A single flowering peak was evident in each population, but ocurred at different times. Large plants produced more flowers, and bloomed earlier and for a longer period than small plants. Population synchrony increased as the mean duration of flowering per individual decreased. The onset of flowering is primarily related to the variance in winter minimum temperatures and the duration to the autumn–winter mean maximum temperature, whereas spring mean maximum temperature is best correlated with synchrony.

Conclusions

Plant size affects individual plant fecundity as well as flowering time. Thus the population structure strongly affects flowering phenology. Indications of clinal variation in the timing of flowering and reproductive effort suggest selection pressures related to the arrival of migrating pollinators, climate and resource economy in a desert environment. These pressures are likely to be relaxed in populations where individual plants can attain large sizes.Key words: Flowering phenology, optimal timing, plant size, Sonoran Desert, Stenocereus thurberi, temperature     

Tracing Genetic Lineages of Captive Desert Tortoises in Arizona

Abstract: We genotyped 180 captive desert tortoises (Gopherus agassizii) from Kingman (n = 45), Phoenix (n = 113), and Tucson (n = 22), Arizona, USA, to determine if the genetic lineage of captives is associated with that of wild tortoises in the local area (Sonoran Desert). We tested all samples for 16 short tandem repeats and sequenced 1,109 base pairs of mitochondrial DNA (mtDNA). To determine genetic origin, we performed assignment tests against a reference database of 997 desert tortoise samples collected throughout the Mojave and Sonoran Deserts. We found that >40% of our Arizona captive samples were genetically of Mojave Desert or hybrid origin, with the percentage of individuals exhibiting the Mojave genotype increasing as the sample locations approached the California, USA, border. In Phoenix, 11.5% were Sonoran–Mojave crosses, and 8.8% were hybrids between desert tortoise and Texas tortoise (G. berlandieri). Our findings present many potential implications for wild tortoises in the Sonoran Desert of Arizona. Escaped or released captive tortoises with Mojave or hybrid genotypes have the potential to affect the genetic composition of Sonoran wild populations. Genotyping captive desert tortoises could be used to inform the adoption process, and thereby provide additional protection to native desert-tortoise populations in Arizona.     

Genetic diversity of the endangered endemic Agave victoriae-reginae (Agavaceae) in the Chihuahuan Desert

Long-lived perennials are a species-rich, ecologically important component of the North American deserts, yet we know little about their genetic structure, information important for their conservation. Agave victoriae-reginae is an endemic of the Chihuahuan Desert of northern Mexico that is endangered by collection for the ornamental trade. We examined levels and patterns of variation at ten polymorphic allozyme loci in ten populations representing the range of the species. Levels of genetic variation (mean H_e= 0.335) and differentiation (mean F_ST = 0.236) were high. Phenetic clustering suggested the existence of at least three distinct groups of populations. If this pattern of variation is representative of other long-lived desert perennials, it may explain the species richness of this group and will pose a real challenge to gene conservation efforts.     

Implications of Anthropogenic Landscape Change on Inter-Population Movements of the Desert Tortoise (Gopherus agassizii)

In the Sonoran Desert of North America, populations of the desert tortoise (Gopherus agassizii) occur in rocky foothills throughout southwestern Arizona and northwestern Mexico. Although tortoise populations appear to be isolated from each other by low desert valleys, individuals occasionally move long distances between populations. Increasingly, these movements are hindered by habitat fragmentation due to anthropogenic landscape changes. We used molecular techniques and radiotelemetry to examine movement patterns of desert tortoises in southern Arizona. We collected blood samples from 170 individuals in nine mountain ranges and analyzed variability in seven microsatellite loci to determine genetic differentiation among populations. Gene flow estimates between populations indicate that populations exchanged individuals historically at a rate greater than one migrant per generation, and positive correlation between genetic and geographic distance of population pairs suggests that the limiting factor for gene flow among populations is isolation by distance. Life history traits of the desert tortoise, a long-lived species with delayed sexual maturity, may severely constrain the ability of small populations to respond to disturbances that increase adult mortality. Historic gene flow estimates among populations suggests that recovery of declining populations may rely heavily on the immigration of new individuals from adjacent mountain ranges. Management strategies compatible with the evolutionary history of gene flow among disjunct populations will help ensure the long-term persistence of Sonoran desert tortoise populations.     

Life history variation in Campanula americana (Campanulaceae): population differentiation

Genetic variation in life history traits has important consequences for life-history evolution. Here we report the results of a greenhouse experiment investigating the broad sense genetic basis of variation in life history traits within and among five populations of Campanula americana distributed along a latitudinal gradient. The populations exhibit differentiation for a number of morphological traits (seed weight, number of branches, final plant size, number of capsules) and the phenological traits, days to emergence, days to bolting, the onset of flowering, and the duration of flowering. Families within populations differed only in days to emergence and seed weight. These results suggest that the life history differences among populations are genetically based. In addition, two life history types—winter annuals and biennials—have previously been reported from natural populations of Campanula americana. This experiment identified a third type—summer annuals from the Florida population.     

Effects of altered temperature and precipitation on desert protozoa associated with biological soil crusts

Biological soil crusts are diverse assemblages of bacteria, cyanobacteria, algae, fungi, lichens, and mosses that cover much of arid land soils. The objective of this study was to quantify protozoa associated with biological soil crusts and test the response of protozoa to increased temperature and precipitation as is predicted by some global climate models. Protozoa were more abundant when associated with cyanobacteria/lichen crusts than with cyanobacteria crusts alone. Amoebae, flagellates, and ciliates originating from the Colorado Plateau desert (cool desert, primarily winter precipitation) declined 50-, 10-, and 100-fold, respectively, when moved in field mesocosms to the Chihuahuan Desert (hot desert, primarily summer rain). However, this was not observed in protozoa collected from the Chihuahuan Desert and moved to the Sonoran desert (hot desert, also summer rain, but warmer than Chihuahuan Desert). Protozoa in culture began to encyst at 37 degrees C. Cysts survived the upper end of daily temperatures (37-55 degrees C), and could be stimulated to excyst if temperatures were reduced to 15 degrees C or lower. Results from this study suggest that cool desert protozoa are influenced negatively by increased summer precipitation during excessive summer temperatures, and that desert protozoa may be adapted to a specific desert's temperature and precipitation regime.     

Pollinator availability as a determinant of flowering time in ocotillo (Fouquieria splendens)

Summary Ocotillo, a perennial shrub of Sonoran and Chihuahuan Deserts, produces its red tubular flowers in spring. This timing coincides with northward migration of hummingbirds through desert areas. Observations of visitors, pollen collections, and seed set reductions following exclusion of different flower visitors indicate that both hummingbirds and solitary bees pollinate ocotillo in southern Arizona. Seed sets of flowers on marked plants varied considerably within and between years, and this variation was related to the temporal match between flowering and hummingbird migration. This suggests that selection acts on plants to synchronize flowering with periods of pollinator abundance.     

Phenological responses to nitrogen and water addition are linked to plant growth patterns in a desert herbaceous community

Increases in nitrogen (N) deposition and variation in precipitation have been occurring in temperate deserts; however, little information is available regarding plant phenological responses to environmental cues and their relationships with plant growth pattern in desert ecosystems. In this study, plant phenology and growth of six annuals in response to N and water addition were monitored throughout two consecutive growing seasons in 2011 and 2012 in a temperate desert in northwestern China. The effects of N and water addition on reproductive phenology differed among plant species. N and water addition consistently advanced the flowering onset time and fruiting time of four spring ephemerals; however, their effects on two spring‐summer annuals were inconsistent, with advances being noted in one species and delays in another. N and water addition alone increased plant height, relative growth rate, leaf number, flower number, and individual biomass, while their combinative effects on plant growth and reproductive phenology were dependent on species. Multiple regression analysis showed that flowering onset time was negatively correlated with relative growth rate of two species, and negatively correlated with maximum plant height of the other four species. Our study demonstrates that phenological responses to increasing precipitation and N deposition varied in annuals with different life histories, whereby the effects of climate change on plant growth rate were related to reproductive phenology. Desert annuals that were able to accelerate growth rate under increasing soil resource availability tended to advance their flowering onset time to escape drought later in the growing season. This study promotes our understanding of the responses of temperate desert annuals to increasing precipitation and N deposition in this desert.     

Habitat Use by Sonoran Desert Tortoises

ABSTRACT The distribution of desert tortoises (Gopherus agassizii) spans a wide range of biotic and abiotic conditions in the southwestern United States and northwestern Mexico, with physical and behavioral differences distinguishing tortoises inhabiting the Mojave Desert from those inhabiting the Sonoran Desert. Relative to tortoise populations in the Mojave Desert, populations in the Sonoran Desert have not been well-studied. To assess how habitat use of desert tortoises in the Sonoran Desert was influenced by topography, vegetation, geomorphology, and soil, we surveyed 40 randomly located 3-ha sites for presence of adult tortoises within a site-occupancy framework. We modeled both occupancy and detection probability as a function of environmental features, and compared those results with a logistic regression model that assumed detection probability was equal to 1. Results from both approaches agreed, suggesting that habitat selection of tortoises in the Sonoran Desert was influenced primarily by topographic and geomorphologic features rather than by vegetation. Specifically, tortoises were more likely to occupy sites that were steep (we detected tortoises on 29% of sites with mean slope <5° and 92% of sites with mean slope >15°) and predominantly east-facing (53% of sites with <5% of site facing E and 92% of sites with >20% facing E), and less likely to occupy north-facing slopes (100% of sites with <10% of site facing N and 14% of sites with >60% facing N). Our results contrast with patterns of habitat use in the Mojave Desert where tortoises primarily occupy valley bottoms. Habitat use of tortoises in Sonoran and Mojave Desert populations differ considerably, contributing to the mounting body of evidence suggesting that these geographically distinct populations may represent separate species.     

Assortative mating by flowering time and its effect on correlated traits in variable environments

Reproductive timing is a key life‐history trait that impacts the pool of available mates, the environment experienced during flowering, and the expression of other traits through genetic covariation. Selection on phenology, and its consequences on other life‐history traits, has considerable implications in the context of ongoing climate change and shifting growing seasons. To test this, we grew field‐collected seed from the wildflower Mimulus guttatus in a greenhouse to assess the standing genetic variation for flowering time and covariation with other traits. We then created full‐sib families through phenological assortative mating and grew offspring in three photoperiod treatments representing seasonal variation in daylength. We find substantial quantitative genetic variation for the onset of flowering time, which covaried with vegetative traits. The assortatively‐mated offspring varied in their critical photoperiod by over two hours, so that families differed in their probability of flowering across treatments Allocation to flowering and vegetative growth changed across the daylength treatments, with consistent direction and magnitude of covariation among flowering time and other traits. Our results suggest that future studies of flowering time evolution should consider the joint evolution of correlated traits and shifting seasonal selection to understand how environmental variation influences life histories.     

Genetic diversity in Chihuahuan Desert populations of creosotebush (Zygophyllaceae: Larrea tridentata)

We examined isozyme variation in the dominant Chihuahuan Desert shrub, Larrea tridentata (creosotebush), to determine the genetic variation within and among populations, the biogeographic relationships of populations, and the potential inbreeding in the species. We surveyed 17 populations consisting of 20 to 50 individuals per population along a 1600-km north-south transect across the Chihuahuan Desert. The southernmost population was near Villa Hidalgo, Mexico, and the northernmost near Isleta Pueblo, New Mexico. All 12 isozyme loci examined were polymorphic (H(t) = 0.416), with up to nine alleles per locus. Despite high levels of variation, we detected moderate inbreeding in L. tridentata populations. Most variation was found within rather than among populations (G(ST) = 0.118). Furthermore, recently established populations in the northern limits of the Chihuahuan Desert did not show decreased levels of genetic variation (H(o) = 0.336). A significant correlation was found between pairwise genetic and geographic distances (r = 0.305). Larrea tridentata showed and continues to show a massive range expansion into the arid and semi-arid regions of the American Southwest, but as shown by the high genetic variation, this expansion took place as a wave, rather than a series of founder events.     

Adaptive phenology of desert and Mediterranean populations of annual plants grown with and without water stress

Summary The dynamics of vegetative and reproductive growth were compared in matched pairs of Mediteranean and desert populations of three unrelated annual species, Erucaria hispanica (L.) Druce, Brachypodium distachyon (L.) Beauv. and Bromus fasciculatus C. Presl., under high and low levels of water availability in a common-environment experiment. Plants of all desert populations showed earlier switches to reproductive development and to subsequent phenophases, and the transition to flowering occurred at smaller plant sizes. Water stress had no effect (E. hispanica) or slightly accelerated the transition to flowering in B. fasciculatus (by 1–2 days) and in B. distachyon (by 4–6 days). Plant senescence was strongly enhanced by water stress, and this enhancement was greater in desert populations than in corresponding Mediterranean ones. Duration of life cycle was greatly shortened by water stress in all three species. Desert and Mediterranean populations of the three species exhibited small differences in their relative response, i.e. phenotypic plasticity, to water stress for phenological and plant size parameters. In E. hispanica and B. fasciculatus the population x water regime interaction amounted to less than 3% of total variance. By contrast, the Mediterranean population of B. distachyon was much more plastic in its response to water stress than the desert population in its transition to plant senescence. Plants from the desert populations appeared to be adapted to shorter, more compact growth cycles, culminating in earlier dates of seed maturation and plant senescence. In addition, they showed larger phenotypic plasticity in the transition to plant senescence, which trait was enhanced or magnified by sustained or repeated lack of water. By contrast, plants from Mediterranean populations delayed switchover from one phenophase to the next, seeming thus to bet on more water being forthcoming.     

Architecture of coastal and desert Encelia farinosa (Asteraceae): consequences of plastic and heritable variation in leaf characters

The shrub Encelia farinosa (Asteraceae) exhibits geographic variation in aboveground architecture and leaf traits in parallel with environmental variation in temperature and moisture. Measurements of plants occurring across a natural gradient demonstrated that plants in desert populations produce smaller, more pubescent leaves and are more compact and branched than plants in more mesic coastal environments. This phenotypic variation is interpreted in part as adaptive genetic differentiation; small size and pubescence reduce leaf temperature and thus increase water-use efficiency but at the cost of lower photosynthetic rate, which results in slower growth and more compact growth form. We explored the basis of phenotypic variation by planting seed offspring from coastal and desert populations in common gardens in both environments. Phenotypic differences among populations persisted in both common gardens, suggesting a genetic basis for trait variation. Desert offspring outperformed coastal offspring in the desert garden, suggesting superior adaptation to hot, dry conditions. Herbivore damage was greater for all offspring in the coastal garden. Phenotypic characters also showed plastic responses; all offspring had smaller, more pubescent leaves and more compact growth form in the desert garden. Our results confirm that leaf size and pubescence are heritable characters associated with pronounced variation in plant architecture.     

QUANTITATIVE GENETICS OF SIZE,SHAPE, LIFE-HISTORY,AND FRUIT CHARACTERISTICS OF THE SEED-HETEROMORPHIC COMPOSITE HETEROSPERMA PINNATUM. I. VARIATION WITHIN AND AMONG POPULATIONS

We document phenotypic and genetic variation within and among populations of the seed heteromorphic species Heterosperma pinnatum Cav. (Compositae) in the production of seed morphs and in a variety of life-history and morphological characteristics that might be correlated with seed and head traits. Each trait is found to have significant genetic variance in most or, usually, all populations. Significant among-population genetic variation exists for all traits except number of achenes per head and seedling shape, although some traits have much less genetic variation among than within populations. Number and percentage of intermediate achenes per head, total number of achenes per head, and lengths of central and peripheral achenes had little among-population genetic variation compared to within-population variation. Most traits had slightly less genetic variation among than within populations; however, some traits (percentage of central achenes, length of awns, date that the first flowering head opened, date that the first fruiting head opened, and death date) had more among-population genetic variation. The proportions of achene morphs produced had high broad-sense heritabilities and high among-population genetic variance, except in the case of intermediate achenes. All phenological variables had high among-population genetic variation. Within-population heritabilities were high for dates of first flowering head and fruiting head but low for death date and reproductive interval. Family and population means measured in the greenhouse for traits having high broad-sense heritability or among-population genetic variance were closely correlated with field means for the corresponding families or populations. The amounts of phenotypic variation were similar for traits that were measured in both the field and the greenhouse. These lines of evidence suggest that greenhouse results provide reasonable estimates of genetic variation in the field for this species. Numerous studies have reported variation in the proportion of seed morphs for different heteromorphic-seeded species and have discussed adaptive scenarios for the evolution of seed proportions; however, our investigation is one of only a few that have documented the amount of phenotypic and genetic variation within and among populations.     

Drought Leads to Collapse of Black-Tailed Prairie Dog Populations Reintroduced to the Chihuahuan Desert

ABSTRACT Recently, a conservation strategy developed to restore populations of black-tailed prairie dog (Cynomys ludovicianus) suggested reintroducing animals into the Chihuahuan Desert grasslands of the southwestern United States. Rainfall in desert habitats is lower and more variable compared to rainfall near the center of the prairie dog's range. Additionally, peak rainfall comes months after prairie dogs reproduce in these desert systems. Thus, southwestern populations may be less prolific and fluctuate more than those found in northerly climes. Using mark-recapture and mark-resight techniques, we estimated reproduction and monthly survival from 577 individuals inhabiting 6 reintroduced colonies from 2003 to 2005 in the northern Chihuahuan Desert. During 2003 precipitation was 64% of the long-term average, whereas both 2004 and 2005 had near-average precipitation. Probability that a female became pregnant, number of juvenile prairie dogs emerging from maternity burrows, and date of emergence were all correlated to adult female body mass. Adult monthly survival decreased from >0.95 during spring to 0.70 in summer 2003, following a rapid loss in adult body mass that coincided with low precipitation. In 2003 monthly juvenile survival was near zero on 2 of the 3 largest colonies and growth rates of juveniles were half that of subsequent years. Estimated population size declined by 68% (range = 18–91%) from 2003 to 2004, and 5 of 6 populations declined an average of 75% from their original introduction size. Prairie dog populations in desert environs may have a high risk of extirpation caused by weather patterns indicative of desert climates. Our results are important for those managers involved in the conservation of prairie dogs and we suggest that regional differences should be carefully considered prior to any reintroduction effort.     

The influence of Prosopis canopies on understorey vegetation: Effects of landscape position

Abstract. The influence of canopy trees and shrubs on under‐storey plants is complex and context‐dependent. Canopy plants can exert positive, negative or neutral effects on production, composition and diversity of understorey plant communities, depending on local environmental conditions and position in the landscape. We studied the influence of Prosopis velutina (mesquite) on soil moisture and nitrogen availability, and understorey vegetation along a topographic gradient in the Sonoran Desert. We found significant increases in both soil moisture and N along the gradient from desert to riparian zone. In addition, P. velutina canopies had positive effects, relative to open areas, on soil moisture in the desert, and soil N in both desert and intermediate terrace. Biomass of understorey vegetation was highest and species richness was lowest in the riparian zone. Canopies had a positive effect on biomass in both desert and terrace, and a negative effect on species richness in the terrace. The effect of the canopy depended on landscape position, with desert canopies more strongly influencing soil moisture and biomass and terrace canopies more strongly influencing soil N and species richness. Individual species distributions suggested interspecific variation in response to water‐ vs. N‐availability; they strongly influence species composition at both patch and landscape position levels.     

On the relative importance of history and ecology in structuring communities of desert small animals

Several recent studies have compared small mammal community structure across multiple deserts on different continents. These studies have tacitly assumed that variation in community structure was greater between continents than within, and so have not evaluated variation across desert regions within continents. I evaluated several metrics of community structure and a model of community assembly for four desert regions in North America - the Great Basin, Mojave. Sonoran. and Chihuahuan Deserts in order to explicitly compare these metrics across these deserts. Additionally. I compared these results with similar analyses conducted on two desert regions in central Asia - the Gobi Desert and the Turan Desert Region to evaluate the relative magnitude of intra- vs inter-continental variation. Although the patterns observed are complex, they demonstrated marked heterogeneity in desert small mammal communities within North America. However, this heterogeneity is much less than that observed in inter-continental comparisons, in which Asian and North American deserts differ markedly. These results agree with other recent studies providing limited or no support for the existence of substantial convergence in community characteristics or ecological function across geographically distant regions. Rather, the results support the hypothesis that the common evolutionary history of faunas in globally disjunct landmasses has had a stronger influence on the evolution of communities and faunas than do regional variations in climate, physiography, etc. Whereas a common ecological setting may have large impacts on some facets of organismal structure (e.g., bipedalism in desert small mammals), common evolutionary history appears to have a more profound influence on local dynamics.