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.     

Reproduction and demography of Townsendia aprica (Asteraceae), a rare endemic of the Southern Utah Plateau

Townsendia aprica (Asteraceae: Astereae), a rare pulvinate perennial of the Southern Utah Plateau, was listed as threatened under the U.S. Endangered Species Act in 1985. Here we report on the reproductive biology and pollination of this little-known species and provide an estimate, for a single site-year, of size-specific reproductive effort. Last Chance townsendia appears a short-lived perennial that begins reproducing in its 2nd year (1.5-2.0 cm diameter). Maximum reproductive effort is attained with the 2.5-4.0 cm diameter size class: 38% of these plants produced 84% of the flower heads. Few plants survived past the 4-cm size class. The species is primarily self-incompatible: neither autogamous nor geitonogamous breeding system treatments produced a significant number of achenes. Unlike some populations of some congeners, the Ivie Creek population was not apomictic. Outcrossing is the primary means of reproduction and native solitary bees are the most important pollinators. Paramount are several species in the genus Osmia , and the ground-nesting species Synhalonia fulvitarsis , which nests among the T. aprica plants. Synhalonia fulvitarsis also visits a contemporaneous blooming phlox ( P. austromontana ), which may facilitate pollination of the rare townsendia. The Townsendia-Phlox-Synhalonia interaction may represent another example of why we must consider communities rather than individual species in our conservation efforts.     

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.     

Pollinator sharing by three sympatric milkvetches, including the endangered species Astragalus montii

Insects visiting flowers of the endangered Heliotrope milkvetch, Astragalus montii , were compared with those visiting two common sympatric congeners, A. kentrophyta and A. miser , on three sites on the Wasatch Plateau of central Utah for 2 yr. We recorded 27+ species of bees, most of which were uncommon, visiting the three species. All three species were primarily visited by native bees of the genera Osmia (15 species) and/or Bombus (4 species). Most Osmia species visited the three species of Astragalus indiscriminantly; bumblebees preferred A. miser and avoided A. montii . Our hypothesis that A. montii flowers would receive fewer total bee visits and be visited by fewer bee species than their common congeners was rejected: A. montii was intermediate to the two common species in its attractiveness to bees. Also rejected was our hypothesis that the greater similarity between A. montii and A. kentrophyta in flower size, flower morphology, and microhabitat would be associated with greater similarity of flower visitors than either had with A. miser . The data suggest that, rather than competing with each other for pollinators, the three species of Astragalus facilitate each other's visitation rates.     

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.     

On the relative importance of floral color, shape, and nectar rewards in attracting pollinators to Mimulus

Pollinator preferences were observed for the six species of section Erythranthe of the genus Mimulus using greenhouse-grown plants placed in a meadow in the Red Butte Canyon Natural Area, Salt Lake County, Utah. The principal pollinators were hummingbirds and bumble bees. Hummingbirds preferred the species with the most reflexed tubular flowers regardless of color, whereas bumblebees preferred pink. lavender, or yellow flowers to red flowers regardless of shape. Results for the six species were confirmed by observations of F 2 hybrid recombinant plants selected such that flower color could be held constant and flower shape varied and vice versa.     

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.     

Patterns of insect flower visitation in Lavandula buchii Webb (Lamiaceae), an endemic shrub of Tenerife (Canary Islands)

The foraging ecology of insect visitors of the endemic lavender Lavandula buchii (Lamiaceae) was studied in a relict population of the plant in north-west Tenerife (Canary Islands). Timed observations were used to describe this pollination system in terms of the structure of the visitor assemblage, foraging patterns and efficiencies of the main insect species. The visitor pool was dominated by anthophorid solitary bees. Furthermore, it was characterized by a high level of endemicity and a small number of species compared with continental relatives of Lavandula. The main visitors, two anthophorids and a bombyliid fly, clearly differed in their foraging behaviour, i.e. visitation frequency, visit duration, rate of flower probing, and in anatomical traits (body size and proboscis length). In these terms, the bees were by far more efficient foragers than the fly. These results are discussed in the context of insect-flower interactions in insular pollination systems.     

Seed production in Gentiana newberryi (Gentianaceae)

Experimental manipulations and observations in one population of Gentiana newberryi Gray flowers over 2 years showed significant variation in seed production relative to pollinator and soil water availability. When pollinators were rare, there was a significant relationship between number of bees present and number of mature seeds produced, and supplemental hand cross-pollination (xenogamy) did improve seed set in Gentiana newberryi Gray. When pollinators were abundant, supplemental hand cross-pollination did not increase seed set. Self-fertilized seeds (autogamy) germinated at the same rate as cross-pollinated seeds. Seed production in unvisited flowers is probably limited anatomically and is not influenced by the type of fertilization. There was a significant relationship between soil moisture and flower size in G. newberryi , with larger flowers found in wetter areas.     

Is the occluded corolla of Antirrhinum bee-specialized?

This paper provides the first results of pollinator fauna associated to the reproductive biology of Antirrhinum charidemi, Antirrhinum graniticum and Antirrhinum braun-blanquetii. Censuses of over 16 hours spanning the flowering phenology of each species were drawn up. As previously hypothesized, the characteristic occluded (personate) corolla of Antirrhinum was exclusively pollinated by bees, but by a higher number (four) of bee families than predicted. Seven bee species (Bombus hortorum, Anthidium manicatum, Chalicodoma lefebvrei, Anthidium sticticum, Anthophora dispar, Xylocopa violacea and Anthidium cingulatum) account for over 90% of flower visits (2098). The flower visitation index varied between species (0.0126?0.0320), reaching relatively medium values compared to Mediterranean plants. In contrast, reproductive success was found to be high when estimated as both fruit (45.4–84.2%) and seed (75.4–98.4%) sets, suggesting that these bee guilds are effective pollinators of the three self-incompatible Antirrhinum species. Specialization of the personate flower of Antirrhinum is therefore interpreted for bees with both large bodies and high visitation indices.     

Reproductive phenology and flower visitors' guild of Canistropsis microps (Bromeliaceae) in an Atlantic Rainforest of southeastern Brazil

The reproductive phenology and guild of the floral visitors of Canistropsis microps in an Atlantic Forest area in Ilha Grande, RJ were studied for 4 years. The rates of flowering, fruiting and mortality were related to air temperature, rainfall and photoperiod. To register the flower visitors, observations were carried out for 5 days in each reproductive season. The flowering and fruiting occurred in the rainy season and were positively related to the studied climatic variables, whereas the highest mortality rates occurred in the dry season and were negatively related to the climatic variables. The guild of floral visitors of C. microps was composed of Hymenoptera, characterizing the species as a melittophilous plant. These results indicate that the rainy season presents more favourable conditions for investment in sexual reproduction, besides being the period in which potential pollinators have larger population densities and a longer period of activity.     

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.     

Interactions between the sexually deceptive orchid Spiculaea ciliata and its wasp pollinator Thynnoturneria sp. (Hymenoptera: Thynninae)

Males of the thynnine wasp Thynnoturneria sp. attempt to mate with female decoys in the flowers of the elbow orchid Spiculaea ciliata. Experimentally shifted orchids usually attract male wasps quickly, often within 2 minutes of presentation of the ‘bait’ orchids in appropriate habitat. Although the orchid effectively exploits the scramble competition mating system of the wasp, the insect is not totally at the mercy of the deceptive orchid. Fewer than half of all arriving males contact the column of the orchid flower, as required for orchid pollination. Moreover, the number of deceived visitors falls sharply over a short period and the number of wasp visitors does not rebound with the replacement of one bait orchid by another at that location. These observations suggest that patrolling wasp pollinators can discriminate to some extent between orchid decoys and female wasps, especially by learning to avoid particular locations that are associated with unrewarding flower decoys.     

Pollinators of Astragalus monoensis Barneby (Fabaceae): new host records; potential impact of sheep grazing

Important bee species inhabiting the study area are listed, including those observed and collected foraging on Astragalus monoensis , a California rare plant. The significance of each species as a potential pollinator is assessed, based on frequency of occurrence in collecting, observed and published host plant records, and morphology. Three pollinator categories are proposed: observed and/or collected on the plant, probable visitors, and possible visitors. New host plant records for these species are listed. Current sheep grazing practices in the A. monoensis habitat endanger pollinators in four ways: (1) destruction of potential nest sites, (2) destruction of existing nests and contents, (3) direct trampling of adult bees, and (4) removal of food resources. Exposure of the major bee species to each of these factors is assessed utilizing experimental data and published information.         

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.     

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.     

Time-lapse photography reveals the occurrence of unexpected bee-pollination in Calanthe izuinsularis,an endangered orchid endemic to the Izu archipelago

Calanthe izuinsularis is a rare, beautiful and fragrant orchid endemic to the Izu archipelago. Although it is known that mainland populations of closely related Calanthe species are pollinated by medium- to large-sized bees, it is likely that C. izuinsularis has been forced to alter its floral biology to attract alternative pollinators, as large-sized bees are rarely found on the Izu Islands. Indeed, the unusual floral characteristics of C. izuinsularis, which produces pale flowers that emit a strong and fragrant scent during the night, are considered the adaptation to a moth-pollinator syndrome. As expected, our time-lapse photography using a digital camera revealed that the inflorescences of C. izuinsularis were most frequently visited by nocturnal moths such as Noctuidae, Geometridae and Crambidae, which accounted for 50% of all insect visits (in terms of both the number of frames with captured visitors and the number of times visited). However, our study could not provide evidence of a pollinator shift towards moths because none of the moth species was observed with pollinaria. On the other hand, our study found that the pollinaria of C. izuinsularis could become attached to the mesothorax of the small-sized sweat bee Lasioglossum occidens. Therefore, our study suggests that C. izuinsularis is still somewhat dependent on bee pollination, while the orchid has changed its floral colouration and scent. The occurrence of bee-pollination in C. izuinsularis would provide an explanation for how the natural hybridisation between C. aristulifera, C. discolor and C. izuinsularis can occur.     

Morphology of oil-collecting pilosity of female Rediviva bees (Hymenoptera: Apoidea: Melittidae) reflects host plant use

The oil-collecting hair morphology and pilosity on female foretarsi for all 26 species of the southern African endemic bee genus Rediviva was studied and related to host plant use based on about 2700 flower visitation records. Four different types of tarsal pilosity and six principal hair types are identified. Rediviva bees collect floral oil from 12 plant genera representing the families Iridaceae, Orchidaceae, Scrophulariaceae and Stilbaceae, with Diascia being the most frequently used host. Differences in foretarsal pilosity reflect the availability of floral oil and elaiophore morphology of the principal host flowers. Rediviva species with short female forelegs exhibit a greater diversity of tarsal oil-collecting pilosity than long-legged species. This contradicts an earlier hypothesis that the pilosity of species with shorter legs might be less complex than in Rediviva with elongate forelegs.     

A fish survey of the White River, Nevada

In spring and summer 1991 and 1992, we surveyed fishes of the White River system, Nye and White Pine Counties, Nevada, to determine the status of natives. There are 5 known native fishes to the White River: Lepidomeda albivallis (White River spinedace), Crenichthys baileyi albivallis (Preston White River springfish), Crenichthys baileyi thermophilus (Moorman White River springfish), Catostomus clarki intermedius (White River desert sucker), and Rhinichthys osculus ssp. (White River speckled dace). All 5 had declined in range. Lepidomeda albivallis had experienced the greatest decline, with less than 50 remaining, and these were restricted to a 70-m stream reach. Rhinichthys osculus spp. was most widespread, found in 18 spring systems. Cottus bairdi (mottled sculpin) was collected for the 1st time from the White River system, where it was probably native. Protective measures should be implemented to conserve all native White River fishes to include C. bairdi.