The ploidy races of Atriplex confertifolia (Chenopodiaceae)

Previous accounts of polyploidy in the North American salt desert shrub Atriplex confertifolia (shadscale) have dealt with the distribution of polyploidy and the morphological and secondary chemical differences between races. The present study amplifies these studies and reveals additional ploidy-flavonoid races, with ploidy levels known to extend from 2x to 12x, and all except 2x and 12x represented by races with and without 6-methoxylation of flavonol compounds. Results of this study show that diploids across their range have about 113% as much DNA per genome as do polyploids and that parallel variation in monoploid genome size between diploids and accompanying polyploids can be shown in different parts of the species’ range. Polyploidy, therefore, appears to have developed independently in several areas of the western United States. Hexaploids are generally not as common as octoploids in shadscale, which could be an indication of diploidization of older tetraploid races. Informes anteriores sobre la poliploidía en el arbusto Atriplex confertifolia (“shadscale”) del desierto de sal norteamericano, han tratado la distribución de la poliploidía, las diferencias químicas secundarias y las diferencias morfológicas entre razas. El presente estudio amplía los estudios previos y revela razas ploidía-flavonoides adicionales, con niveles de ploidía que se sabe que van de 2x a 12x, y todos, con excepción de 2x y 12x, están representados en razas con y sin 6-metoxilación de compuestos de flavonoles. Durante el estudio descubrí que los diploides a lo largo de su rango de distribución tienen alrededor del 113% más ADN por genoma que los polploides. También descubrí que la variación paralela en el tamaño del genoma monoploide entre los diploides y sus poliploides correspondientes, se puede manifestar en diversas partes del rango de distribución de la especie. La poliploidía, por lo tanto, parece haberse desarrollado de forma independiente en varias regiones del occidente de los EE.UU. Los hexaploides son generalmente menos comunes que lo octoploides en el shadscale, lo cual podría ser indicio de diploidización de razas tetraploides más antiguas.     

Comparative habitat and community relationships of Atriplex confertifolia and Sarcobatus vermiculatus in central Utah

Thirty-four study sites were established in shadscale ( Atriplex confertifolia [Torr. & Frem.] Wats.) and greasewood ( Sarcobatus vermiculatus [Hoov. Torr. in Emory) communities bordering Utah Lake in central Utah. Differences in species composition, vegetation, and soil characteristics were assessed. Significant differences in soil factors between the two communities were found for sand, calcium, manganese, zinc, and copper. Soluble salts and sodium concentrations were generally higher in the greasewood type, but differences were not significant. Major differences were found in understory species, with burr buttercup ( Ranunculus testiculatus Grantz) showing significantly greater cover in the shadscale community and cheatgrass ( Bromus tectorum L.) showing significantly greater cover in the greasewood community.     

Natural polyploidization within tetraploid and hexaploid populations of the desert shrub Atriplex confertifolia

Shadscale ( Atriplex confertifolia ) is a wind-pollinated dioecious shrub of western North America with an unusual development of apparently autoploid races, showing all even ploidy levels from 2x to 12x (base x = 9). Of these, tetraploid races are the most frequently encountered, with octoploids the next most common, and hexaploids being much less common. In this study, the occurrence of neopolyploid individuals within tetraploid and hexaploid populations of shadscale was examined at natural sites in order to investigate the pathways by which octoploids may be formed and to find a possible explanation for the predominance of octoploids over hexaploids. The frequencies of apparent neopolyploid individuals among adults and the frequencies of their backcrosses were tabulated, as was the occurrence of neopolyploidy among seeds of female plants from some of the same populations. The neopolyploids encountered in these surveys were almost exclusively those expected from unions involving single unreduced gametes. 6x presumed neopolyploid plants were observed in tetraploid populations at a frequency of 0.20%, and 6x seeds were observed at a frequency of 1.91%. Within examined hexaploid populations, 9x adults were not encountered, but 9x seeds were observed at a rate of 0.52%. Evidence was obtained for the operation of some postulated autoploid pathways for generation of octoploids, but these routes did not operate at rates comparable to those for the production of hexaploids. Although the question needs to be studied by genetic methods, the most probable reason for scarcity of hexaploids in nature appears to be that some tetraploid races are diploidized, which would result in irregularities of meiosis in their neohexaploids and a failure to form new races due to infertility and inviability. El “shadscale” ( Atriplex confertifolia ) es un arbusto dioico del oeste de Norteamérica polinizado por el viento que exhibe el desarrollo inusual de razas aparentemente poliploides, con todos los niveles pares de ploidía desde 2x hasta 12x (x base = 9). De éstos, los más comunes son los tetraploides y luego los octoploides. Los hexaploides son mucho menos comunes. En este estudio, examinamos la presencia de individuos neopoliploides entre poblaciones tetraploides y hexaploides del shadscale en sitios naturales para investigar los medios por los cuales se pueden formar los octoploides y para encontrar una posible explicación para el predominio de los octoploides sobre los hexaploides. Tabulamos la frecuencia de individuos aparentemente neopoliploides entre adultos y su descendencia retrocruzada, así como la presencia de neopoliploides entre semillas de las plantas femeninas de algunas de las mismas poblaciones. Los neopoliploides encontrados en estos muestreos fueron casi exclusivamente los que se esperarían de la unión de un gameto no reducido y un gameto reducido. Observamos plantas 6x presunta mente neopoliploides en poblaciones tetraploides con una frecuencia del 0.20%, o del 1.91% en las semillas. Entre las poblaciones examinadas de hexaploides, no encontramos adultos 9x, aunque los observamos con una frecuencia del 0.52% en las semillas. Obtuvimos evidencia de la operación de algunos procesos autoploides postulados para explicar la generación de octoploides, pero estos procesos no operan a frecuencias comparables a aquellas para la producción de hexaploides. Aunque es preciso investigar el tema mediante métodos genéticos, la causa más probable para la escasez de los hexaploides en entornos naturales parece ser que algunas razas tetraploides son diploidizadas, produciendo irregularidades meióticas en sus neohexaploides, los cuales no logran formar razas nuevas debido a su infertilidad o inviabilidad.     

Carbon isotope discrimination in the C 4 shrub Atriplex confertifolia along a salinity gradient

Carbon isotope discrimination (Δ) was measured for leaves of Atriplex confertifolia along a salinity gradient in northern Utah. Over this gradient, the variation of Δ values was high for a C 4 species, and the Δ values were positively correlated with salinity in both years of study. Of the possible explanations for this patter, the Δ results are consistent with the notion that salinity induces an increase in the bundle sheath leakiness of these C 4 plants.     

Temperature and high salinity effects in germinating dimorphic seeds of Atriplex rosea

Atriplex rosea L. (Chenopodiaceae; tumbling orach), an annual herb, is a widely established weedy species of disturbed sites in all counties of Utah. Seeds of Atriplex rosea were collected from a salt marsh in Faust, Utah, and are dimorphic, light brown, and 2-2.5 mm wide, or black and 1-2 mm wide. Seed germination responses of the black and brown seeds were studied over a range of salinity and temperature. Both brown and black seeds germinated at 1000 mM NaCl, and the optimal temperature for germination of both types was 20°-30°C. Variation in temperature, however, affected germination of black seeds more than brown seeds. At lower thermoperiod only 40%-50% black seeds germinated in nonsaline control, and germination was almost completely inhibited with the inclusion of salinity. However, all brown seeds germinated in control at temperatures above 5°-15°C, and inhibition caused by salinity was comparatively lower. Brown seeds had a higher germination rate than black seeds at all temperature and salinity treatments. The highest rate of germination of both seeds occurred at the temperature regime of 5°-15°C. Recovery of germination for black seeds when transferred to distilled water after being in various salinity treatments for 20 days was quite variable. Recovery decreased with increase in salinity at lower temperature regimes, increased with salinity at optimal thermoperiod, and had no effect at 20°-30°C. Brown seeds recovered poorly from salinity at all thermoperiods except 5°-15°C, where recovery decreased with an increase in salinity. Brown seeds are adapted to germination in the early part of the growing season, whereas black seeds are capable of surviving harsher conditions and can germinate in later time periods. Characteristics of the dimorphic seeds increase chances for survival in the harsh saline desert environment.     

Differential phytosociological interactions involving male and female Atriplex bonnevillensis

Wind-pollinated dioecious plants often exhibit spatial segregation of the sexes. This partial niche separation has most often been explored using abiotic niche axes. However, if the sexes are truly separated in space, then they are apt to encounter different plant species that may heavily affect growth and reproduction. Also, to the extent that their niches differ, the sexes may respond differently to the same co-occurring species. Here we examine interspecific interactions that influence male and female reproductive potential in Atriplex bonnevillensis. Using Emlen’s interaction assessment, a technique which assesses species interactions based on cover classes, we show that Salsola species compete significantly with females but not males, while Halogeton glomeratus competes with males but not females. The effect of competition only became apparent when we corrected for site-specific fertility. These results imply that differential competition must be considered when studying dioecious plants that display spatial segregation of the sexes.     

Names and types in perennial Atriplex Linnaeus (Chenopodiaceae) in North America selectively exclusive of Mexico

Cited are names and combinations within the woody species of Atriplex as they occur in North America. Types and their repositories are included for all taxa except those for which that information could not be located. New nomenclatural proposals include Atriplex gardneri var. aptera (A. Nelson) Welsh & Crompton, comb. nov.; A. garrettii var. navajoensis (C. A. Hanson) Welsh & Crompton, comb. nov.; Atriplex acanthocarpa var. coahuilensis (Henrickson) Welsh & Crompton, comb. nov. A lectotype is designated for A. breweri S. Watson.     

Isozymes of an autopolyploid shrub, Atriplex canescens (Chenopodiaceae)

Diploid, tetraploid, and hexaploid populations of Atriplex canescens ( x =9) were examined for 18 isozyme systems. Of 24 interpretable loci, only one locus (Per 1 ) was polymorphic. Another locus (Per 2 ) showed a dosage effect. Genetic distance values, D, ranged from near 0 to 0.05, which are in the normal range for local species races. Results from clonal ramets gave identical results. The data and analyses support an essentially autopolyploid origin for the polyploid populations examined.     

Forage quality of rillscale ( Atriplex suckleyi ) grown on amended bentonite mine spoil

At peak standing crop, rillscale ( Atriplex suckleyi ) foliage grown on amended bentonite mine spoil contained adequate digestible energy, crude protein, and all mineral elements except phosphorus necessary for cattle, sheep, antelope, and deer. Amendments (sawdust, NPK, gypsum) generally did not affect forage quality. Iron, manganese, aluminum, sodium, and potassium concentrations were high and may have adversely affected forage quality. Forage utility would be limited to a few months during the growing season.     

Number and condition of seeds in ponderosa pine cones in central Arizona

Ponderosa pine cones from 10 areas in Arizona were collected prior to natural seed dispersal and dissected to determine the number of sound, hollow, and insect-damaged seeds in each cone. Total and sound seed yields per cone did not vary significantly among areas but did vary significantly among trees within each area. Numbers of hollow and Megastigmus -infested (Hymenoptera: Torymidae) seeds varied significantly among areas and trees within areas. Numbers of sound seed increased significantly with increasing cone length but did not change with increasing numbers of cones per cluster. The percentages of Megastigmus -infested seed did not change significantly with increasing cone length or number of cones per cluster.     

Pilfering of stored seeds and the relative costs of scatter-hoarding versus larder-hoarding in yellow pine chipmunks

Yellow pine chipmunks ( Tamias amoenus ) scatter-hoard food during summer and autumn but must form a larder as a winter food source before winter begins. Yellow pine chipmunks do not larder-hoard large quantities of food during the summer, apparently because a summer larder could not be defended from pilferers. We tested the assumption that the rate of pilferage from an unguarded larder would be significantly greater than the rate of pilferage from surface caches (which are also unguarded by yellow pine chipmunks) during the summer and autumn. Buried plastic buckets were used as artificial nests containing larders of 1000 sunflower seeds or 200 Jeffrey pine ( Pinus jeffreyi ) seeds. The pilferage of larder contents was monitored daily and compared to pilferage of surface caches. Animals (yellow pine chipmunks and deer mice, Peromyscus maniculatus ) removed sunflower seeds from caches much faster than from larders, but caches of Jeffrey pine seeds disappeared much more slowly than pine seeds in larders. Further, animals removed pine seeds from larders more quickly than they did sunflower seeds from larders. The difference between seed species was probably because sunflower seeds have much stronger odors, which rodents readily detect, and because chipmunks prefer pine seeds over sunflower seeds. Yellow pine chipmunks must spend a considerable portion of their time foraging for seeds and may not be able to defend a large larder during summer.     

Production, losses, and germination of Ceanothus fendleri seeds in an Arizona ponderosa pine forest

I quantified seed production and ovule losses for Ceanothus fendleri Gray (Fendler ceanothus) plants protected from large ungulate herbivores in a ponderosa pine ( Pinus ponderosa [Laws]) forest of northern Arizona. I also tested seed germination responses to cold stratification and heat treatments in the laboratory. Fruit production on fecund stems ranged from 7.4 to 38.2 fruits per stem, which equated to 22.2-118.2 potential seeds based on 3 ovules per fruit. Stems that produced fruit were significantly large relative to their expected sizes. Predispersal ovule losses ranged from 70.7% to 91.4% across the 2 years studied. A chalcidoid seed parasite ( Eurytoma squamosa Bugbee) consumed 11%-28% of the total number of seeds produced. Postdispersal seed predation varied from 0% to 24% and was significantly affected by forest floor substrate in 1 study year. Cumulative ovule losses were estimated to be 71%-92%. Cold stratification did not significantly affect seed germination, and exposure to 90°C resulted in the highest germination percentage. Both dormant and nondormant seeds suggested a bet-hedging life history strategy. This study provides basic ecological information important for management of ponderosa pine forest and nursery production of C. fendleri .     

Lagomorphs and the dispersal of seeds into communities dominated by exotic annual weeds

Large areas of western rangeland are presently dominated by alien annual weeds such as Bromus tectorum (cheatgrass). These communities resist succession to perennial communities primarily because the annuals are competitively superior to establishing perennial seedlings and they promote fires that favor weeds over perennials. Succession may be further slowed, however, by low rates of seed dispersal into annual grasslands. We investigated the role of lagomorphs ( Sylvilagus nuttalli , Nuttall's cottontail; S. audubonii , desert cottontail; and Lepus californicus , black-tailed jackrabbit) in seed dispersal across an ecotone: 50 m into woodland, border, and 20 m, 50 m, and 100 m into grassland. We searched pellets for juniper seeds visually and for any other species through germination from crushed pellets after cold, moist stratification. Pellets were not even distributed across transects, but there was no trend with respect to position of transect. Juniperus osteosperma (Utah juniper) was the most abundant seed. Both the number of juniper seeds and the proportion of pellets with juniper seeds decreased steadily from a high in woodland to absence at 100 m into grassland. Only 2 dicot seedlings emerged from pellets, 1 Salsola pestifer and 1 unknown that died prior to identification. Consequently, there was little seed movement into grassland; 72% of all seeds were collected from either woodland or border transects. Lagomorphs apparently do not effectively replenish the native perennial seed pool of cheatgrass-dominated disturbances at Dugway.     

The braconid parasitoids (Hymenoptera: Braconidae) associated with seeds of Dipterocarpaceae in Malaysia

Nineteen species of Braconidae have been reared from seeds of Dipterocarpaceae (mainly Shorea species) in West Malaysia. Eleven new species (Brachistinae: Schizoprymnus mesocaudus sp. nov.; Schizoprymnus torreadoroides sp. nov.; Braconinae: Bracon nitidisoma sp. nov.; Bracon rugulifer sp. nov.; Syntomernus shoreatus sp. nov.; Spermatobracon triangulornatus sp. nov.; Cheloninae: Ascogaster cornifera sp. nov.; Ascogaster lamellifera sp. nov.; Phanerotoma divergens sp. nov.; Microgastrinae: Apanteles hemiaurantius sp. nov.; Apanteles striatus sp. nov.) and one new genus (Spermatobracon gen. nov.; type species: Spermatobracon triangulornatus gen. nov. and sp. nov.) are described and illustrated.     

Dispersal of sagebrush-steppe seeds by the western harvester ant ( Pogonomyrmex occidentalis )

Like many seed-harvesting ants, the western harvester ant ( Pogonomyrmex occidentalis ) can act as both a predator and a seed disperser. Dispersal results when seeds are dropped en route to the nest, are left in nest granaries when colonies die or are abandoned, or are removed from granaries and discarded in nest middens. This study examined the density and species identity of seeds discarded in harvester ant nest middens and compared them with those found in nearby soils. Nineteen species of seeds were recovered from middens, compared with 13 species in 5-m reference areas and 9 species in adjacent disk areas. Total density of sound seeds was nearly 3 times higher in middens than at 5 m from the nest and nearly 50 times higher than in disk soils. Moreover, 4 of the 6 most common species overall were significantly more abundant in middens. One species, Munro globemallow ( Sphaeralcea munroana ), was recovered from nearly 50% of middens but was not found in the other 2 areas. These findings suggest that the western harvester ant is a potentially important disperser of some sagebrush-steppe plant species.     

Comparing the effects of granivorous rodents on persistence of Indian ricegrass ( Oryzopsis hymenoides ) seeds in mixed and monospecific seed patches

In desert environments seeds are often heterogeneously distributed in small patches that vary in number of seed species and in seed density. Because seed harvest by rodents is often density dependent (a larger proportion of seeds is removed from high-density seed patches than from low-density patches), the proportion of residual or post-harvest seeds should be greater in low-density patches. In addition, seed preference can affect harvest. We tested whether the residual proportion of a highly preferred seed (Indian ricegrass, Oryzopsis hymenoides ) was less when in a seed patch with a 2nd species (mixed-species patch) than when in a monospecific seed patch. We predicted that the increased overall seed density due to the presence of 2 species in a patch would result in a lower residual proportion of ricegrass seeds in the mixed-species seed patches than in the monospecific patches. As predicted, the residual proportions of Indian ricegrass seeds were less each time ricegrass was paired with one of 6 other species in mixed-species patches. Similarly, the residual proportion of each of those 6 species was less when paired with ricegrass than when in a monospecific patch. We speculate on the potential implications of these results for the population dynamics of plant species and the physical structure of plant communities.     

Desiccation sensitivity and heat tolerance of Prunus ilicifolia seeds dispersed by American black bears ( Ursus americanus )

Carnivore consumption of fruit is a principle means by which many fleshy-fruited plant species achieve long-distance seed dispersal. We examined carnivore dispersal of hollyleaf cherry ( Prunus ilicifolia ) seeds, specifically assessing the survival, desiccation sensitivity, and germination of seeds found in bear scats. Studies were conducted both in the laboratory and in 2 burn areas in Los Padres National Forest, California. Bear scats containing P. ilicifolia seeds were collected in burned and unburned chaparral. We counted seeds in each scat and noted whether endocarps had tooth punctures or rattled audibly when shaken. For comparative germination trials, we also collected fruits and seeds directly from mature shrubs. In the laboratory, following a cold-moist stratification period, seeds were assessed for germinability. In the field, we compared desiccation rates and germinability of seeds from bear scats and freshly collected seeds. We compared rates of moisture loss and germination for seeds subjected to several different conditions, including 25 °C (room temperature), 30 °C and 65 °C (in the lab), and placement on exposed soil in a burn area (in the field), where midday temperatures were approximately 45 °C but likely much higher by late afternoon. Prunus ilicifolia seeds collected from bear scats were largely undamaged; the vast majority of these seeds germinated. In some cases, germination rate was higher for seeds from scats than for seeds from intact fruits. Several results indicate that desiccation is an important cause of reduced germinability. First, seeds that rattled audibly germinated poorly; and the louder the rattle, the lower the germination percentage. Second, seeds (both fresh and those from bear scats) placed in the field under protective screens had greatly reduced levels of germination (an 84% decline) after only 7 days. Third, seeds dried in the lab, even at relatively moderate temperatures, showed a decline in germination with seed moisture loss. The addition of high temperatures accelerated this decline in germination. We discuss the relevance of heat and desiccation sensitivity of seeds dispersed by bears to successful seed germination in burned and late-seral mesic and xeric chaparral. El consumo de fruta por carnívoros es uno de los principales medios por los que muchas especies de plantas de frutos carnosos logran dispersar sus semillas a través largas distancias. Examinamos la dispersión por carnívoros de las semillas del islay ( Prunus ilicifolia ), un arbusto del chaparral, evaluando concretamente la supervivencia, la sensibilidad a la desecación y la germinación de semillas encontradas en las heces de oso. Estos estudios se llevaron a cabo tanto en el laboratorio como en dos áreas quemadas en Los Padres National Forest, California. Se colectaron heces de oso que contenían semillas de P. ilicifolia en el chaparral quemado y en el no quemado. Contamos las semillas en cada bola fecal y notamos si los endocarpios tenían marcas de dientes y si vibraban audiblemente al agitarse. Para llevar a cabo pruebas comparativas de germinación, también colectamos frutos y semillas directamente de los arbustos maduros. En el laboratorio, después de un período de estratificación fría-húmeda, evaluamos la capacidad de germinación de las semillas. En el campo, comparamos las tasas de desecación y de germinación de semillas de las heces de oso con aquellas de semillas colectadas directamente de los arbustos. Se compararon las tasas de pérdida de humedad y germinación de las semillas sujetas a diversas condiciones, entre ellas 25 °C (temperatura ambiente), 30 °C y 65 °C en el laboratorio, y en el campo sobre suelo expuesto en un área quemada, donde la temperatura al mediodía era ~45 °C pero probablemente mucho más alta por la tarde. En general, las semillas de P. ilicifolia colectadas en las heces de oso no estaban dañadas; la gran mayoría de estas semillas germinaron; en algunos casos la tasa de germinación fue más alta que la de frutos intactos. Varios resultados indicaron que la desecación es una causa importante de la reducción en la capacidad de germinación. Primero, las semillas que vibraban audiblemente germinaron poco, y cuanto más fuerte el sonido, menor la germinación. Segundo, las semillas (tanto las frescas como las colectadas de heces de oso) colocadas en el campo bajo una malla protectora exhibieron niveles de germinación muy reducidos (una disminución del 84%) después de sólo siete días. Tercero, la germinación de las semillas desecadas en el laboratorio, incluso a temperaturas relativamente moderadas, disminuyó con la pérdida de humedad de las semillas. Temperaturas más altas aceleraron esta disminución en la germinación. Discutimos la importancia de la sensibilidad al calor y a la desecación en las semillas dispersadas por los osos para la germinación exitosa en el chaparral mésico y árido, tanto en el chaparral recién quemado como en el que está en las etapas finales de sucesión.     

Seed-feeding beetles of the weevil tribe Mecysolobini (Insecta: Coleoptera: Curculionidae) developing in seeds of trees in the Dipterocarpaceae

Species comprising two monophyletic groups of Alcidodes sensu lato (crassus group and dipterocarpi group) feed on fruits of the ecologically and economically important tropical timber family Dipterocarpaceae. Fifteen Alcidodes species are newly described from several thousand specimens of seed-feeding beetles reared from south-east Asian dipterocarp hosts (Dipterocarpoidae) and four other species are revised. Morphological keys are provided for identifying these 19 taxa. Weevil associations are provided for 70 species of Dipterocarpaceae in five genera (16 Dipterocarpus spp., four Dryobalanops spp., six Hopea spp., 39 Shorea spp. and five Vatica spp.). These records relate primarily to specimens reared from seeds in Borneo and in Peninsular Malaysia, but all known previous host records are also included (from India, Andaman Islands, Thailand, Laos, Vietnam and the Philippines) and assessed. These validated and documented host associations refute general assertions that (1) seed-eating beetles are host-specific in the tropics and (2) seed-eating beetles found on dipterocarp hosts are broad generalists across species in several genera of mast-fruiting Dipterocarpaceae. Most Alcidodes species exploit seeds from several dipterocarp species. Some Alcidodes species occur across broad geographic regions with several host species from a dipterocarp genus, other species have been found only on Bornean Hopea or Dipterocarpus species. Others, such as the two previously undescribed species feeding on Vatica species, may be restricted to separate sections of the host genus. Within an extensively sampled and diverse interspecific mast-fruiting dipterocarp community, no Alcidodes species was observed to feed on sympatric species across genera within the Dipterocarpaceae. Additional host records and specimens of Alcidodes spp. throughout the range of the Dipterocarpaceae in south and southeast Asia are required to further define these clades and the tribe Mecysolobini in general as well as to resolve the complex host associations and distributions documented.