PHOTOSYNTHESIS AND CARBON ALLOCATION IN TIPULARIA DISCOLOR (ORCHIDACEAE), A WINTERGREEN UNDERSTORY HERB

Seasonal patterns of photosynthesis and carbon allocation were determined for Tipularia discolor, a summer-deciduous wintergreen orchid of the southeastern United States, to assess the effects of environmental conditions and leaf age on carbon acquisition and allocation patterns. There was no shift in the optimum temperature for photosynthesis (T_opt) on a seasonal basis and T_opt (≈26 C) was at least 10 C higher than daily maximum air temperature during most of the growing season. Lack of photosynthetic adjustment in Tipularia to seasonal fluctuations in temperature and light suggested that the photosynthetic characteristics of this wintergreen were more similar to those of spring ephemerals than to those of evergreens and summer-active herbs. The decline in photosynthetic capacity during the winter growing season for Tipularia, largely due to leaf age effects, gradually reduced net photosynthetic rates in the field despite more favorable light and temperature conditions. Photosynthesis in the field was primarily limited by environmental conditions in early- and mid-season and by photosynthetic capacity in late-season. A ¹⁴CO₂ labelling experiment demonstrated that patterns of carbon allocation to vegetative structures were affected by the season of photosynthetic carbon fixation, whereas reproductive structures received 21% of the recovered labelled carbon regardless of the period of labelling. Carbon acquired and stored during all periods of the growing season was used to produce new vegetative and reproductive structures.     

THE MODE OF POLLINATION IN HABENARIA OBTUSATA (ORCHIDACEAE)

The role of mosquitoes and moths as pollinators of Habenaria obtusata was studied in northern Wisconsin during the summer of 1969. In each population studied, meter-square quadrants were marked and the number of plants, flowers per inflorescence, rate and time of pollinia removed, and capsule set recorded. Carbon dioxide and modified New Jersey Light Traps were set during the flowering period to capture pollinia-bearing insects. The study showed that female mosquitoes of the genus Aedes (particularly A. communis) and two species of Geometrid moths of the genus Xanthorhoe are important pollinators of H. obtusata. Pollination occurred during the last few days of flowering with an average of 18.0% of the pollinia removed and with 14.2% capsule set.     

THE POLLINATION ECOLOGY OF ZENOBIA (ERICACEAE)

Zenobia is a shrub endemic to the coastal plain of the Carolinas and southeastern Virginia. The sole species, Zenobia pulverulenta, appears to be weakly self-compatible and partially self-pollinating. Fruit set was reduced, but not eliminated by excluding insect floral visitors. Insect visitors were collected and their pollen loads examined for Zenobia and foreign pollen. Analysis of these pollen loads and observations of insect-foraging behavior indicate that workers of three species of bumblebee (Bombus bimaculatus, B. griseocollis and B. impatiens) are. the principal pollinators of Zenobia. However, pollinator density and composition appear to be strongly influenced by the habitats in which Zenobia grows. Pollen and nectar serve as the primary rewards for pollinators.     

THE POLLINATION ECOLOGY OF SOLANUM ROSTRATUM (SOLANACEAE)

The pollination mechanisms and pollen vectors of Solarium rostratum have been examined by greenhouse experiments and field studies. Although the capacity for autogamy exists in this weedy annual, it rarely occurs because of two factors: (1) the morphology of the flower and (2) the foraging behavior of the various species of Bombus, the primary pollen vector in the regions studied. The percentages of geitonogamy and xenogamy are dependent on the flight pattern of the bees and the number of open flowers on a plant.     

POLLINATION BIOLOGY OF LISTERA CORDATA (ORCHIDACEAE)

Pollination biology of northern California populations of Listera cordata (L.) R. Br. is presented. The flowers are allogamous, self-compatible, and protandrous. At anthesis, access to the stigma is blocked by the flap-like rostellum. When an insect contacts the touch-sensitive trigger hairs of the rostellum, the pollinia are suddenly released and become cemented to the insect's body. One to two days after the pollinia are removed, the rostellum rises and exposes the receptive stigma. Visitors are attracted to the flowers by their foul odor and minute quantities of nectar presented on the surface of the labellum. Pollination efficiency is high with 61–78% of flowers from several localities setting fruit. The most important and abundant pollinators of L. cordata at our study sites are fungus gnats (Sciaridae and Mycetophylidae). Other less frequently encountered pollinators are members of these families: Braconidae, Ichneumonidae, and Tipulidae. At present, there is no compelling evidence that the floral characteristics of L. cordata have evolved as a result of an evolutionary interaction with any specific group of insects. Nor is there any indication that the flowers are sapromyophilous, despite their putrid odor.     

THE POLLINATION ECOLOGY OF DELPHINIUM TRICORNE (RANUNCUL ACEAE)

The pollination ecology of Delphinium tricorne, a spring-ephemeral herb of eastern deciduous forests, was studied at eight sites in the Ohio River basin from West Virginia to southern Illinois. The plant was found to be obligately dependent for its pollination on queen bumblebees (Bombus Latr. spp.) and hummingbirds, to which the flowers are closely adapted in form, color, function, and blooming phenology. Other bees, including Anthophora ursina, Osmia bucephala, Ptilothrix bombiformis, and Xylocopa virginica; Lepidoptera, including Amphion nessus, Danaus plexippus, Epargyreus clams, Erynnis juvenalis, Hemaris thysbe, Papilio glaucus, P. philenor, P. troilus, Poanes zabulon, Vanessa atalanta, and V. cardui; and the beefly Bombylius major foraged for nectar and/or pollen on the flowers without pollinating them. Techniques employed in the study included insect exclosures to test fertility of plants in the absence of pollinators, analysis of floral colors by reflectance spectrophotometry and ultraviolet photography, analysis of pollinator behavior by cinematography and close-range stereophotography, collection and identification of visitors to flowers, and identification of pollen types carried by foragers on the flowers.     

THE POLLINATION ECOLOGY OF DICENTRA CUCULLARIA

In northeastern Iowa and southwestern Wisconsin the flowers of Dicentra cuculiarla were found to be pollinated almost exclusively by Bombus bimaculatus nectar-foraging queens, which were phenologically synchronized in their emergence from hibernation with the flower's anthesis. Cinematographic and stereophotographic evidence indicated that pollen transfer was effected by the ventral side of the insect's head and anterior thorax contacting essential flower parts and to a lesser degree by the front and middle legs contacting pollen-laden edges of the inner petals. Lepidoptera, Diptera, and small Hymenoptera occasionally encountered on the flowers were ineffective in pollination. Abundant Apis mellifera pollen-foraging workers regularly effected pollination, but being an introduced species it exhibits no naturally developed pollination adaptation to the flower. Nectar spur perforation by B. affinis nectar-foraging queens did not affect plant fertility, and this behavior was related only in part to forager tongue length. Nectar-foraging behavior of B. bimaculalus queens on the flowers was correlated with the phenological development of the annual insect colonies.     

POLLINATION ECOLOGY OF PYRRHOPAPPUS CAROLINIANUS (COMPOSITAE)

Pyrrhopappus carolinianus and Hemihalictus lustrans constitute a mutualistic association: the early morning flowering of Pyrrhopappus provides the matinal bee with a nearly exclusive pollen source, although other plants must be visited for nectar. Female Hemihalictus, the primary pollen vector, tear open the anthers and remove the pollen before it is available to other bees. The foraging behavior of the bee insures cross-pollination. The flight pattern of the bees generally restricts the pollen dispersal range. If cross-pollination fails, then autogamy results from twisting of the styles that brings the stigmas in contact with the pollen presented on the styles of other florets. Schinia mitis is an important predator of the capitulae of P. carolinianus.     

THE POLLINATION ECOLOGY OF ASTRAGALUS CIBARIUS AND ASTRAGALUS UTAHENSIS (LEGUMINOSAE)

Astragalus cibarius and A. utahensis are common perennial species of a widespread legume genus. The pollination of Astragalus has been briefly discussed in the literature, but little work has been done on species in the intermountain West. This study was conducted from 1970–1973 in Utah with mixed and single species populations. The flowers of both species were homogamous and papilionaceous, but the species were different as to color, size, and ultraviolet reflectance. Astragalus cibarius usually flowered 10 days ahead of A. utahensis, but both species flowered earlier than most other plants in the community. Bagging experiments indicated both species were strongly allogamous. Exclosure studies indicated both species relied on insects as pollen vectors. Of the 44 insect species which were observed visiting flowers, only 14 carried Astragalus pollen, and the pollinator fauna varied between study sites. Pollen quantities and distributions on Diptera and Coleoptera indicated a poor potential for pollination. Floral structure, pollen distribution and quantity, and behavior implied that large bees of the families Apidae and Anthophoridae were the primary pollinators. These bees visited only one species of Astragalus when the plants occurred in mixed populations; this constancy may have been related to relative flower abundance. Non-pollinating floral foragers affect other phases of Astragalus life history.     

STUDIES OF THE MONOTROPOIDEAE (ERICACEAE). FLORAL NECTARIES: ANATOMY AND FUNCTION IN POLLINATION ECOLOGY

The morphological and vascular characteristics of the nectaries of the species of the Monotropoideae (Ericaceae) reflect the morphology and declination of the flowers. These, in turn, are related to the pollination systems of the species. The nectaries of members of the Monotropoideae exhibit a range in form from slender, elongate projections, to short, stout ones, or to low ridges between the staminal bases.     

THE DEVELOPMENT OF THE FEMALE GAMETOPHYTE OF EPIPACTIS (ORCHIDACEAE) AND ITS INFERENCE FOR REPRODUCTIVE ECOLOGY

The development of the female gametophytes of Epipactis atrorubens, E. helleborine, and E. palustris have been investigated using confocal scanning laser microscopy. These species have T-tetrads and eight-nucleate female gametophytes in common. The development of the female gametophyte is monosporic in accordance with the Polygonum type. Furthermore, the outer integument grows slowly and reaches to the middle of the inner integument, when the female gametophyte is mature. In the first two species, T-tetrads develop prior to anthesis, which may be correlated with autogamy. The development of the female gametophyte of E. palustris appears to be similar in plants from different localities.     

THE POLLINATION ECOLOGY OF AQUILEGIA ELEGANTULA AND A. CAERULEA (RANUNCULACEAE) IN COLORADO

Aquilegia elegantula Greene and A. caerulea James occur in montane and subalpine habitats in the southern Rocky Mountains of western North America. The red and yellow flowers of A. elegantula are nodding, odorless, protogynous, and secrete a concentrated (44%) sucrose nectar in the floral spurs. Seed set in flowers under pollinator exclosures was 12% while seed set in open-pollinated flowers was 65%. The flowers of A. elegantula are pollinated primarily by the Broad-tailed Hummingbird (Selasphorus platycercus [Swainson]) and by at least three species of pollen-foraging bumblebees, of which Bombus occidentalis Greene is the most common. The blue and white flowers of A. caerulea are erect, mildly fragrant, protandrous, and secrete a 26% sucrose nectar. Seed set in caged flowers in the field averaged 39%. in uncaged flowers 54%. The most important pollinators of A. caerulea are the crepuscular hawkmoth, Hyles (=Celerio) lineata (Fabricius) and ten species of pollen-foraging Bombus. The most abundant bumblebee species, B. occidentalis, is also a frequent nectar thief. Differences in pollination systems alone probably do not constitute an effective anti-hydridization mechanism between A. elegantula and A. caerulea, but do serve to reinforce differences in habitat and flowering time that distinguish the two species.     

BREEDING SYSTEM AND POLLINATION ECOLOGY OF TRIENTALIS BOREALIS (PRIMULACEAE)

Selfing in Trientalis borealis is reduced by spatial separation of stigma and anthers during anther dehiscence and by internal self-incompatibility. Artificial self-pollination resulted in low levels of fruit set (2.0%). Crosses made within patches of plants yielded variable (0–72.7%), but generally low fruit set (x̄ = 21.4%). This may be caused by patches often consisting of a single genet. In contrast crosses made among patches resulted in uniformly high levels of fruit set (x̄ = 84.3%). Pollinator activity was higher at open sites than at heavily shaded sites and was correlated with fruit set. Pollinator activity, however, showed no correlation with seed set. We conclude that the primary pollinators, Halictid and Andrenid bees, play an important role in the breeding system of the plant by promoting outcrossing but that availability or efficiency of pollinators may limit seed set.     

POLLINATION BIOLOGY OF PLATANTHERA STRICTA (ORCHIDACEAE) IN OLYMPIC NATIONAL PARK,WASHINGTON

Platanthera striata Lindley is entomophilous and can produce seed via facultative self-pollination and intraracemic and interracemic pollination. Capsule production is pollinator-limited and seed set may be pollen-limited. In experimental plants capsules produced via self- and intraracemic pollination contained fewer seeds with normally developed embryos than did capsules produced via interracemic pollination. The inflorescence of Platanthera stricta is fragrant and is attractive to a wide array of anthophilous insects. It is pollinated by a diverse assemblage of short-tongued insects. The primary pollinators are Eustroma fasciata B. and McD. (Lepidoptera: Geometridae), Bombus flavifrons Cresson and B. melanopygus Nylander (Hymenoptera: Apidae), an undescribed species of Greya (Lepidoptera: Prodixidae), and several species of Empis, Rhamphomyia, and Anthepiscopus longipalpis Melander (Diptera: Empididae). Small amounts of glucose are present on the raceme. The extrafloral glucose may retain small pollinators on the inflorescence until they locate the floral spur aperture.     

COMMUNITY STUDIES IN POLLINATION ECOLOGY IN THE HIGH TEMPERATE ANDES OF CENTRAL CHILE. I. POLLINATION MECHANISMS AND ALTITUDINAL VARIATION

Pollination mechanisms and pollinators are reported for a total of 137 species (75% of the non-abiotically pollinated flora) as they occur at three altitudinal levels (subandean scrub: 2,200–2,600 m; cushion-plant zone: 2,700–3,100 m; subnival feldfield: 3,200–3,600 m) in the Andean (alpine) zone on the Cordon del Cepo (33°17'S) in central Chile as part of community oriented research in reproductive biology in the high temperate Andes of South America. Only around 4% of the species studied failed to be visited by potential pollinators. Hymenopterans (principally bees) are important pollinators of 50% of the biotically pollinated flora, butterflies of 24% and flies of 46%. Other vectors include beetles, moths, and hummingbirds. An estimated 17% of the flora is anemophilous. Bee species-richness, specialist feeding, and melittophily reach maxima in the subandean scrub; thereafter, bees diminish rapidly in number, with bees pollinating only 13% of the subnival flora as contrasted with 68% of the subandean flora. Although fly and butterfly species-richness also decline with increasing altitude, the proportions of species pollinated by these vectors actually increases. High-altitude populations of melittophilous species with broad altitudinal ranges are invariably serviced by fewer bee species as compared with lower populations. The rich bee fauna at the lower end of the Andean zone in central Chile appears to have resulted from upward colonization from that of the subtending lowland Mediterranean sclerophyllous woodland vegetation. Altitudinal variation in pollination spectra is discussed in relation to contrasting life history characteristics and different modes of thermoregulation in the insect groups involved.     

POLLINATION ECOLOGY OF LABIATAE IN A PHRYGANIC (EAST MEDITERRANEAN) ECOSYSTEM

This study was conducted in a phryganic (East Mediterranean) ecosystem at Daphni, near Athens, Greece. The Labiatae, represented by ten species belonging to nine genera, dominate in this ecosystem type. They flower from February to July. Both flowering time and nectar quantity are related to the species ability to tolerate intense water stress. Labiatae are visited by 201 insect species. Of these, 43 are exclusively supported by the family and 37 are monotropous. Solitary bees (mainly Anthophoridae, Megachilidae, Halictidae) constitute 47.3% of pollinators. The family is important in hosting specialized bees (15 species) in phrygana, particularly late in the flowering season. Labiatae species form two equally represented groups in this system; namely, the late winter-early spring (early) flowering, visited by relatively few pollinator species, and the late spring-summer (late) flowering species, visited by numerous pollinators. This temporal distinction is accompanied by different pollination profiles that include duration of anthesis, reward to pollinators, floral attractiveness, and flower character differentiation. All of these attributes are maximized in the early flowering period. This strategy suggests a mechanism for resource partitioning at a time when the pollinator resource is limited and competition for the services of pollinators is expected to be intense. Contrary to the current theory concerning cornucopian species, the copiously rewarding flowers of Labiatae in phrygana are not those abundantly serviced by pollinators.