A mixed pollination system in Penstemon pseudospectabilis M. E. Jones (Scrophulariaceae)

A population of Penstemon pseudospectabilis M. E. Jones (Scrophulariaceae) on Cave Creek in Cochise County, Arizona, was used in an experimental test of reproductive fitness with three caging treatments: all flying pollinators excluded, hummingbirds excluded, and no exclusion. Twenty plants were chosen and three shoots on each used in the experiment. The flowers were 25.6 (s.d. = 1.5) mm long, the end diameter was 8.5 (s.d. = 1.0) mm, the tube opening was 6.5 (s.d. = 1.0) mm, and the greatest diameter, 75% distal from the receptacle, was 10.1 (s.d. = 0.8) mm (N= 59). Floral nectar contained 11.7% (s.d. = 2.9%) fructose, 13.8% (s.d. = 2.7%) glucose, and 74.5% (s.d. = 5.4%) sucrose (N = 74). There was some evidence, significant only for fructose, that nectar-sugar composition varies between morning and evening. Larger floral dimensions were correlated with lower sucrose and higher hexoses. Casual observation showed Xylocopa sp., small bees, flies, and hummingbirds to be visitors. There was no sign of nectar robbing. Five percent of flowers set seed with all pollen vectors excluded, 44% with hummingbirds excluded, and 63%with no exclusion. Seed set per fruit was 2 with all excluded, 23 with hummingbirds excluded, and 46 with no exclusion. Mean seed set on pollinated flowers was 60, with a range of 2 to 192. Multiple linear regression showed the fraction of fruit setting seed when hummingbirds were excluded to be related to larger flower diameters and shorter flowers. With no pollinator exclusion, fruits setting seed were related to larger diameters and nectar fructose. For seeds per fruit, multiple regression gave similar, but less clear, results. We conclude that P. pseudospectabilis is pollinated by both bees and hummingbirds, with other pollinators not to be excluded as possible contributors. We found no hard evidence of selective forces currently at work.     

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 .     

Breeding biology and flower visitors of the rare White River penstemon, Penstemon scariosus var. albifluvis (Scrophulariaceae)

We studied the breeding system and flower visitors of White River penstemon, a rare endemic from the Uintah Basin of eastern Utah and western Colorado. Bagging treatments and hand-pollination treatments showed that Penstemon scariosus var. albifluvis has a mixed mating system: while some seeds and fruits are produced through selfpollination (both autogamy and geitonogamy), significantly more are produced when flowers are cross-pollinated. The primary flower visitors, and likely pollinators, were several species of native twig- and ground-nesting bees in the families Apidae, Halictidae, and Megachilidae. We found no differences in fruit or seed production between open-pollinated controls and hand-outcrossed flowers, suggesting that pollinator visits were sufficient to maximize female reproductive success. Management plans to conserve White River penstemon must recognize that full reproductive success of this rare plant taxon relies on a suite of pollinating bees, and that the species richness and abundances of bee visitors should be maintained.     

Reproductive biology, hybridization and flower visitors of rare Sclerocactus taxa in Utah's Uintah Basin

We studied the mating system and flower visitors of 2 threatened species of Sclerocactus (Cactaceae) in the Uintah Basin of eastern Utah—an area undergoing rapid energy development. We found that both S. wetlandicus and S. brevispinus are predominantly outcrossed and are essentially self-incompatible. A third presumptive taxon (undescribed; here called S. wetlandicus -var1) is fully self-compatible but cannot produce seeds unless the flowers are visited by pollinators. We found tentative evidence for pollinator limitation of fruit set in S. brevispinus but not in the other taxa. Pollinators are largely species of native ground-nesting bees in the subfamily Halictinae. These bees are indiscriminate collectors of pollen and nectar from Sclerocactus flowers and other plant species. The well-being of Sclerocactus populations requires that land managers ensure the preservation of halictid bee habitat, which includes both nesting sites and a diversity of floral resources. These bees are also undoubtedly the pollen vectors responsible for the production of hybrids in areas where S. wetlandicus and S. brevispinus meet. We experimentally confirmed that reciprocal crosses between S. wetlandicus and S. brevispinus are indistinguishable from each other or from pure crosses in fruits/flower, seeds/fruit and seed weights. We suggest that hybridization preceded energy development, that it attests to the ongoing process of evolution in the Sclerocactus group, and that it should not be suppressed by anthropogenic activity.     

Effect of soil microtopography on seed bank distribution in the shrub-steppe

The relationship between soil surface cryptogamic crusts and seed banks was investigated in the shrubsteppe in the Lower Columbia Basin. Seventy-four percent of the seeds in a disturbed bunchgrass community were found in crevices bordering cryptogamic crust polygons. In a sagebrush/bunchgrass community, 89% of the seeds were found in crevices. In a disturbed bunchgrass community, Bromus tectorum seeds were found in both the seed bank and aboveground vegetation communities. Bromus tectorum seeds were located in the seed bank of a sagebrush/bunchgrass community, although it had a minor presence in the aboveground community. Seeds of Artemisia tridentata Nutt. were not found in either the bunchgrass or sagebrush/bunchgrass communities. The high number of seeds found in crevices bordering the cryptogamic crust suggests that crevices play a role in determining the small-scale distributional pattern of shrub-steppe plants at the Fitzner-Eberhardt Arid Lands Ecology Reserve.     

Pollination experiments in the Mimulus cardinalis--M. lewisii complex

Experimental sets of Mimulus cardinalis and M. lewisii plants were (1) exposed to pollinators and (2) shielded from pollinators at study sties in Red Butte Canyon and Big Cottonwood Canyon, Wasatch Mountains, Utah. The exposed plants produced 1,535 seedlings and the shielded plants only 1. Clearly, seed production is dependent upon cross-pollination. A few syrphid flies were observed visiting the flowers but no hummingbirds or bumble bees, although the latter two have been reported as the main pollinators of M. cardinalis and M. lewisii , respectively. No interspecific hybrids were produced even though the species are fully interfertile, indicating that pollinators are faithful to their species or that different parts of their bodies pick up and carry pollen to the two different species.     

Remarkable waxing, waning, and wandering of populations of Mimulus guttatus : an unexpected example of global warming

Experimental sets of Mimulus cardinalis and M. lewisii plants were (1) exposed to pollinators and (2) shielded from pollinators at study sties in Red Butte Canyon and Big Cottonwood Canyon, Wasatch Mountains, Utah. The exposed plants produced 1,535 seedlings and the shielded plants only 1. Clearly, seed production is dependent upon cross-pollination. A few syrphid flies were observed visiting the flowers but no hummingbirds or bumble bees, although the latter two have been reported as the main pollinators of M. cardinalis and M. lewisii , respectively. No interspecific hybrids were produced even though the species are fully interfertile, indicating that pollinators are faithful to their species or that different parts of their bodies pick up and carry pollen to the two different species.     

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.     

Pollination needs of arrowleaf balsamroot, Balsamorhiza sagittata (Heliantheae: Asteraceae)

Arrowleaf balsamroot, Balsamorhiza sagittata (Pursh) Nutt, is a common, sometimes dominant, longlived forb that flowers early in spring from the foothills to upper-montane areas of the northern Rocky Mountains and Intermountain West. Public land managers desire its seed for rangeland rehabilitation. Through manual pollination field trials, the species was found to have a mixed pollination system. It is primarily xenogamous (46% of ovules yielded plump achenes) but partially self-compatible (31% of achenes were plump). Unvisited flower heads formed virtually no mature achenes; only plump achenes contained seeds with endosperm. Freely visited flower heads in 2 populations produced as many achenes as manual outcross pollinations of flower heads, suggesting that seed production was not pollinator limited. Two species of Osmia bees rely mostly on Balsamorhiza and its close relative, Wyethia , for pollen. At least 165 females per hectare will need to be stocked to achieve thorough flower visitation in cultivated seed production fields.     

Insect-mediated pollination in slickspot peppergrass, Lepidium papilliferum L. (Brassicaceae), and its implications for population viability

Field experiments on the pollination biology of slickspot peppergrass, Lepidium papilliferum L. (Brassicaceae), a rare species endemic to microsites in sagebrush-steppe habitat in southwestern Idaho, were conducted at 2 sites from May to July 2001. Site KB contained over 10,000 plants, whereas site WG contained less than 150 plants. Insect exclusion experiments revealed that seed production in L. papilliferum is dependent on insect-mediated pollination; median percent seed set dropped from 70% to 2% when insects were excluded from flowers. A total of 25 insect families from 5 orders visited L. papilliferum flowers: 24 families at KB and 11 families at WG. Only 9 families contained more than trace amounts of pollen on their bodies: Hymenoptera-Anthophoridae, Apidae, Colletidae, Halictidae, Sphecidae, Vespidae; Coleoptera-Cerambycidae, Dermestidae; Diptera-Syrphidae. Insects from these families are likely responsible for pollination of L. papilliferum , although some may be of only minor significance due to their infrequent visits to flowers. Two of the 4 most common and pollen-laden insects found at KB, honey bees ( Apis mellifera ) and colletid bees, were absent or rare at WG. Three other pollen-carrying families present at KB, Sphecidae, Vespidae, and Halictidae, were not found at WG. We raise several possible explanations for this disparity in pollinator communities between sites and discuss the importance of pollinator diversity to the long term viability of L. papilliferum .     

Variance and replenishment of nectar in wild and greenhouse populations of Mimulus

We compared nectar production in wild populations and greenhouse-grown populations of the monkey flower species of section Erythranthe of the genus Mimulus . Nectar was sampled from over 1000 flowers. For each flower the volume of nectar was measured with a calibrated micropipette and the percentage of sugar with a hand refractometer. Percentage of sugar varied little from flower to flower in both field and greenhouse studies, but volume varied markedly from flower to flower in field studies and even more in greenhouse studies. This high variance in nectar volumes appears to be intrinsic. The amount of nectar in greenhouse populations tended to increase with time in the absence of pollinators. The amount of nectar in field populations tended to remain the same to pollinator withdrawals. The latter conclusion was corroborated by sampling nectar at 2-h intervals all day and comparing the total volume produced by a flower to the volume of nectar produced in control flowers sampled only at the end of the day.     

A life history study of the Snake River plains endemic Lepidium papilliferum (Brassicaceae)

Lepidium papilliferum is an ephemeral species that occupies ""slick spot"" microhabitats in the matrix of sagebrush steppe vegetation of the southwestern Snake River plains, Idaho, USA. We related population demographic data collected from 1993 to 1996 to on-site precipitation data on the Orchard Training Area west of Boise. We also carried out field seed-retrieval and in situ seed bank studies. We found that L. papilliferum has a dual life history strategy. A fraction of each cohort sets seed as summer annuals, while the remaining plants remain vegetative and potentially biennial. Surviving biennials flower and set seed along with the annual cohort of the following year. The switch to flowering as an annual appears to be based on threshold rosette size. Probability of survival to flowering was much lower for biennials than for annuals of the same cohort, but surviving biennials sometimes had enhanced seed production. The summerdry environment of the Snake River plains combined with the slick spot habitat has apparently selected for a primarily summer annual life cycle for this species. Seeds were highly dormant at dispersal and were not responsive to dormancybreaking cues. Those from a given cohort of L. papilliferum remained viable in the soil for at least 11 years. This persistent seed bank provides a buffer against extinction in sequences of years when seed production is low or absent. Estimated seed bank size varied from near zero for a heavily disturbed site that formerly supported the species to 18 viable seeds · dm -2 for an extant population in high-quality habitat. Management for population preservation for L. papilliferum should focus on protecting the seed bank from destruction caused by livestock trampling and other anthropogenic disturbances.     

Cone and seed production in Pinus ponderosa : a review

Factors associated with seed cone production in Pinus ponderosa were reviewed to identify broad patterns and potential effectiveness of restoration activities. Cone and seed production are quite variable, with differences between (1) years, (2) sites, and (3) individual trees. Between-year, population-wide crop failures suggest large-scale triggers for cone and seed production, perhaps high temperatures and dry weather. Stem diameter is the most important determinant for cone production at the tree level, with other factors such as genetic disposition, moisture, soil nutrients, and insect pests and disease playing a smaller role. Some extrinsic factors affect growth rate, indirectly affecting cone production. For example, less competition and lower stand densities result in P. ponderosa trees that increase in diameter more quickly, possibly because of more light, and produce seeds earlier. This literature suggests that restoration activities, especially thinning, will result in trees better able to produce larger seed crops. The effect of prescribed fire is less clear, with contradictory effects depending on site conditions, burn severity, and nutrient status of the site.     

Geobotany of the Niobrara chalk barrens in Colorado: a study of edaphic endemism

The chalk barrens of the Niobrara Formation in the Arkansas River valley of Colorado exhibit a high degree of plant endemism and rarity. We examined their geochemistry and structure as well as the reproductive ecology of their signature species Mirabilis rotundifolius to assess factors influencing endemism. While no single component consistently identified the barren soils, the natural locations that support Mirablilis can be identified using elemental profiles from a cluster analysis of soil extracts. Because seeds germinate and plants proliferate in transplant gardens with non-calcic soils, bedrock and soil chemistry do not appear to be determinant components of endemism. Mirabilis rotundifoilus utilizes diverse pollinators but is capable of autogamy when insects are excluded; seed predation by harvester ants may be a limiting factor for population spread outside the barrens habitat. Mirabilis rotundifolius is a disturbance-tolerant species, part of a barrens cohort that shares traits of suffrutescence, woody caudices, or multibranched rhizomes penetrating the chalk strata. Its presence may be enhanced when disturbance inhibits the presence of other species that compete for limited water resources. We conclude that endemism is dictated not by unusual soil chemistry requirements but rather by structural components of the barrens. In a water-limited environment, shallow soils and thin chalk strata coupled with a functionally adaptive growth habit allow M. rotundifolius to survive when other species are excluded by disturbance or the physiochemical aspects of the barrens.     

Comparative floral biology of Penstemon eatonii and Penstemon cyananthus in central Utah: a preliminary study

A comparison of the floral visitors of two closely related plant species, Penstemon cyananthus and P. eatonii suggests that flower shape and color may affect the number and type of pollinators, and the ability of the plant to set fruit. Penstemon cyananthus, which is most attractive to hymenopteran visitors, has a blue flower, large in diameter, that is positioned as a convenient “landing pad.” Although many types of insects visit the flower, the transport of pollen directly to flowers of another individual of the same species is somewhat inefficient, since fruiting success is low (66.7 percent). The tubular red flowers of P. eatonii are narrow and droop downward from the stem. The nectar is accessible to a specific and well-adapted visitor, the hummingbird. This less promiscuous, bird-pollinated species sets fruit more successfully (82.4 percent) than P. cyananthus.     

Temporal distribution,seed damage and notes on the natural history of Acanthoscelides quadridentatus and Acanthoscelides winderi (Coleoptera: Chrysomelidae: Bruchinae) on their host plant,Mimosa setosa var. paludosa (Fabaceae: Mimosoideae), in the Brazilian Cerrado

The seed beetles, Acanthoscelides quadridentatus and Acanthoscelides winderi are here recorded for the first time feeding on seeds of Mimosa setosa var. paludosa in the Brazilian Cerrado. Our main aims were to describe the temporal distribution, seed damage, and notes on the natural history of these two species on their host plant. We hypothesised that: (a) healthy seeds from infested fruits would have worse germination rate than healthy seeds from noninfested fruits, and (b) females of seed beetles would lay more eggs on large fruits. We made field observations and an experimental field study with the presence of seed beetles versus their exclusion on plants. Results revealed that seed beetles are synchronised with fruiting, with a temporal partitioning in occurrence. Attacked seeds did not germinate, whereas healthy seeds from infested fruits had worse germination rate than healthy seeds from noninfested fruits. Females of seed beetles laid more eggs on large fruits. These results suggest that seed beetles avoid competition through a temporal partitioning curcial for their coexistence, and select large fruits to oviposit as these fruits probably provide more food resource for their offspring. Furthermore, plants might perceive seed beetles’ damage and then reduce resource allocation on infested fruits.     

Desert rodents reduce seedling recruitment of Salsola paulsenii

Heteromyid rodents in the deserts of North America have been shown to harvest large quantities of seeds of both native and introduced plants from soil seed banks, but rarely has the impact of this seed removal been demonstrated experimentally. I used a series of fenced plots (some of which excluded rodents) to demonstrate that heteromyids at a western Nevada study site can measurably reduce seed banks and subsequent seedling establishment of Salsola paulsenii , an introduced invasive weed that has become a significant problem over much of the desert Southwest. The frequency of S. paulsenii seedlings in both 2004 and 2005 was significantly greater around the interior perimeters of plots that permitted access by rodents than in plots that excluded rodents. Density of S. paulsenii seedlings was significantly greater inside than outside rodent exclusion plots, but there was no such difference in seedling density inside versus outside plots that permitted rodent access. Salsola paulsenii has such a conspicuous presence in many desert environments that the effect of rodents in reducing its abundance may not be visually apparent; however, rodents may still ameliorate competitive effects of this weed on coexisting plants. Heteromyid rodents disperse seeds through caching, and they also consume them. Caching may enhance establishment of native plant seedlings, but is unlikely to benefit exotics such as S. paulsenii .     

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.