The effects of plant density upon pollination success, reproductive effort and fruit parasitism in Cistus ladanifer L. (Cistaceae)

Human activity often causes a decline in the local density of plant populations. Below some critical lower density, populations may suffer a progressive decline in reproductive success because of the difficulties associated with finding suitable mates. Therefore, to conserve endangered plant species it is necessary to understand in greater detail how changes in population density affect different determinants of plant reproductive success. We simultaneously recorded individual plant pollination success, reproductive effort and fruit parasitism in three populations of Cistus ladanifer L. in eastern Portugal. Pollination success declined significantly as distance to the nearest conspecific increased (p<0.001). However, reproductive effort and fruit parasitism showed the opposite pattern (both p<0.001). On average, plants farther than 1 m from their nearest conspecific suffered from three times more fruit parasitism compared to plants closer to a conspecific. Overall, net female reproductive output decreased as nearest neighbor distance rose (p<0.001). Thus, isolated plants were able to compensate only partially for reduced pollination success through increased reproductive effort. We conclude that management plans for plant populations should recognize that reproductive success is the accumulated result of several different processes, which may each respond to plant density in different ways.     

Soluble reactive phosphorus (SRP) transport and retention in tropical, rain forest streams draining a volcanic landscape in Costa Rica: in situ SRP amendment to streams and laboratory studies

Soluble reactive phosphorus (SRP) transport/retention was determined in two rain forest streams (Salto, Pantano) draining La Selva Biological Station, Costa Rica. There, SRP levels can be naturally high due to groundwater enriched by geothermal activity within the surfically dormant volcanic landscape, and subsequently discharged at ambient temperature. Combined field and laboratory approaches simulated high but natural geothermal SRP input with the objective of estimating the magnitude of amended SRP retention within high and low SRP settings and determining the underlying mechanisms of SRP retention. First, we examined short-term SRP retention/transport using combined SRP-conservative tracer additions at high natural in situ concentrations. Second, we attempted to observe a DIN response during SRP amendment as an indicator of biological uptake. Third, we determined SRP release/retention using laboratory sediment assays under control and biologically inhibited conditions. Short-term in situ tracer-SRP additions indicated retention in both naturally high and low SRP reaches. Retention of added SRP mass in Upper Salto (low SRP) was 17% (7.5 mg-P m⁻² h⁻¹), and 20% (10.9 mg-P m⁻² h⁻¹) in Lower Salto (high SRP). No DIN response in either nitrate or ammonium was observed. Laboratory assays using fresh Lower Salto sediments indicated SRP release (15.4 ± 5.9 μg-P g dry wt.⁻¹ h⁻¹), when incubated in filter sterilized Salto water at ambient P concentration, but retention when incubated in filter sterilized river water amended to 2.0 mg SRP l⁻¹ (233.2 ± 5.8 μg-P g dry wt.⁻¹ h⁻¹). SRP uptake/release was similar in both control- and biocide-treated sediments indicating predominantly abiotic retention. High SRP retention even under biologically saturated conditions, absence of a DIN response to amendment, patterns of desorption following amendment, and similar patterns of retention and release under control and biologically inhibited conditions all indicated predominantly abiotic P flux.     

Interspecific Territoriality by a Rufous-Tailed Hummingbird (Amazilia tzacatl): Effects of Intruder Size and Resource Value1

I studied two aspects of interspecific territoriality in a Costa Rican nectarivorous bird, the rufous-tailed hummingbird (Amazilia tzacatl). First, I examined variation in the response of the territory holder to four species of intruding hummingbirds by quantifying the proportion of intruders chased from the territory. This measure of territory defense varied significantly among species of intruders and was negatively related to the intruding species’body mass, possibly due to potential costs associated with becoming involved in escalated contests with larger individuals. Second, I tested for an effect of resource manipulation on territory defense. I increased the resource value of the territory by injecting extant natural flowers with artificial sucrose solution. While the frequency of territorial intrusions did not change, the proportion of intruders chased by the territory holder increased. Apparently, nectar supplementation changed the territory holder's perception of resource value but had little impact on the intruders’perception.     

Pollination biology and the impact of floral display, pollen donors, and distyly on seed production in Arcytophyllum lavarum (Rubiaceae)

In animal-pollinated plants, two factors affecting pollen flow and seed production are changes in floral display and the availability of compatible mates. Changes in floral display may affect the number of pollinator visits and the availability of compatible mates will affect the probability of legitimate pollination and seed production. Distyly is a floral polymorphism where long-styled (pin) and short-styled (thrum) floral morphs occur among different individuals. Distylous plants frequently exhibit self and intra-morph incompatibility. Therefore changes in morph abundance directly affect the arrival of compatible pollen to the stigmas. Floral morph by itself may also affect female reproductive success because floral morphs may display differences in seed production. We explored the effects of floral display, availability of neighboring compatible mates, and floral morph on seed production in the distylous herb ARCYTOPHYLLUM LAVARUM. We found that floral display does not affect the mean number of seeds produced per flower. There is also no effect of the proportion of neighboring legitimate pollen donors on seed production in pin or thrum flowers. However, floral morphs differed in their female reproductive success and the thrum morph produced more seeds. Hand pollination experiments suggest that differences in seed production between morphs are the result of pollen limitation. Future research will elucidate if the higher seed production in thrum flowers is a consequence of higher availability of pollen donors in the population, or higher efficiency of the pin morph as pollen donor.     

Systemic activity of plant extracts in Cedrela odorata (Meliaceae) seedlings and their biological activity on Hypsipyla grandella (Lepidoptera: Pyralidae) larvae

1 Deterrent and toxic substances known to affect Hypsipyla grandella larvae were bioassayed to determine their possible systemic activity under laboratory conditions in Turrialba, Costa Rica. 2 Leaflets from Spanish cedar Cedrela odorata seedlings grown in vitro on culture media treated with different plant extracts were excised from the seedlings and exposed to H. grandella larvae. 3 Both Azatin EC (derived from neem, Azadirachta indica) and plant extracts (from Quassia amara wood, Ruta chalepensis leaves and Sechium pittieri fruits) were able to translocate within Spanish cedar seedlings and express their biological activity against H. grandella larvae. 4 Rates of leaflet consumption were nil for seedlings treated with the commercial standard (carbofuran) and Azatin EC, both causing larval death. 5 Intermediate levels of consumption were reached with the three plant extracts, all of them causing deterrence, because they precluded feeding activity by larvae when exposed to treated leaflets.     

Variation in floral morphology and plant reproductive success in four Ipomoea species (Convolvulaceae) with contrasting breeding systems

• This study tested the hypothesis that self‐compatibility would be associated with floral traits that facilitate autonomous self‐pollination to ensure reproduction under low pollinator visitation. In a comparison of two pairs of Ipomoea species with contrasting breeding systems, we predicted that self‐compatible (SC) species would have smaller, less variable flowers, reduced herkogamy, lower pollinator visitation and higher reproductive success than their self‐incompatible (SI) congeners.
• We studied sympatric species pairs, I. hederacea (SC)– I. mitchellae (SI) and I. purpurea (SC)–I. indica (SI), in Mexico, over two years. We quantified variation in floral traits and nectar production, documented pollinator visitation, and determined natural fruit and seed set. Hand‐pollination and bagging experiments were conducted to determine potential for autonomous self‐pollination and apomixis.
• Self‐compatible Ipomoea species had smaller flowers and lower nectar production than SI species; however, floral variation and integration did not vary according to breeding system. Bees were primary pollinators of all species, but visitation rates were seven times lower in SC than SI species. SC species had a high capacity for autonomous self‐pollination due to reduced herkogamy at the highest anther levels. Self‐compatible species had two to six times higher fruit set than SI species.
• Results generally support the hypothesis that self‐compatibility and autonomous self‐pollination ensure reproduction under low pollinator visitation. However, high variation in morphological traits of SC Ipomoea species suggests they maintain variation through outcrossing. Furthermore, reduced herkogamy was associated with high potential for autonomous self‐pollination, providing a reproductive advantage that possibly underlies transitions to self‐compatibility in Ipomoea.

     

Geographic variation in reproductive success of Stenocereus thurberi (Cactaceae): Effects of pollination timing and pollinator guild

? Premise of the study: It has been proposed that species of columnar cacti from dry tropical areas depend on bats for their reproduction, whereas species from dry subtropical areas are also pollinated by other species. To test this hypothesis, we examined the effects of pollinator guild and of variation in time and space on the reproductive success of a widespread species. ? Methods: Changes in fruit set, seed set, and pollinator activity through time were recorded in three widely separated populations of Stenocereus thurberi. Breeding system and sources of pollination limitation were determined by controlled pollinator exclusions in each population. ? Key results: Significant differences were found in the timing of activity and in the effectiveness of pollinators among sites. In the northern and central populations, reproductive success depends on bats, whereas in the southern population a combination of pollinators was more effective. No difference between open and hand cross-pollination treatments was found in the northern and central populations, which suggests no pollen limitation. However, significant differences were detected in the southern population, which indicates temporal differences in pollinator abundance or arrival time. ? Conclusions: Local variation in pollinator assemblages and reproductive success could greatly affect the evolution of pollination systems. The pattern of generalist pollination in the southernmost populations and specialized pollination in the central and northern populations contradicts the hypothesis of latitudinal variation. In the absence of nocturnal pollinators, the accumulated nectar can sustain visits by diurnal pollinators, a bet-hedging strategy that increases the chances of fruit set in some populations.     

极小种群野生植物崖柏的生殖物候、传粉及胚胎发育研究

以濒危植物崖柏（Thuja sutchuenensis Franch.）为对象,对其生殖物候、传粉机制进行观察,并采用石蜡切片法对其胚胎发育过程进行研究。结果显示：崖柏于8月分化出大、小孢子叶球,次年3月传粉,为花粉无气囊、具传粉滴、胚珠直立型传粉机制,球果于10月开裂;显微观察发现,传粉期花粉进入珠孔后,贮藏在珠心上方的贮粉室内,同时珠心组织中分化出孢原细胞,进入雌配子体发育阶段,5月中旬,花粉管开始萌发,6月初完成受精,进入胚胎发育阶段,10月初,胚胎发育成熟。研究表明崖柏从大、小孢子叶球形成至种子成熟的整个发育过程中均存在败育,而胚珠败育及雌配子体游离核时期至幼胚发育期间的败育是其生殖障碍的主要原因。本研究获得了崖柏生殖生物学的基础资料,为其人工繁育和制定保护策略提供了重要依据。     

Effects of floral herbivory on foraging behaviour of bumblebees and female reproductive success in Pedicularis gruina (Orobanchaceae)

The effects of floral herbivores on floral traits may result in alterations in pollinator foraging behaviour and subsequently influence plant reproductive success. Fed-upon plants may have evolved mechanisms to compensate for herbivore-related decreased fecundity. We conducted a series of field experiments to determine the relative contribution of floral herbivores and pollinators to female reproductive success in an alpine herb, Pedicularis gruina, in two natural populations over two consecutive years. Experimental manipulations included bagging, hand supplemental, geitonogamous pollination, and simulated floral herbivory. Bumblebees not only avoided damaged flowers and plants but also decreased successive visits of flowers in damaged plants, and the latter may reduce the level of geitonogamy. Although seed set per fruit within damaged plants was higher than that in intact plants, total seed number in damaged plants was less than that in intact plants, since floral herbivory-mediated pollinator limitation led to a sharp reduction of fruit set. Overall, the results suggest that resource reallocation within inflorescences of damaged plants may partially compensate for a reduction in seed production. Additionally, a novel finding was the decrease in successive within-plant bumblebee visits following floral herbivory. This may increase seed quantity and quality of P. gruina since self-compatible species exhibit inbreeding depression. The patterns of compensation of herbivory and its consequences reported in this study give an insight into the combined effects of interactions between floral herbivory and pollination on plant reproductive fitness.     

On the low reproductive success of two Cyrtopodium species (Orchidaceae: Cyrtopodiinae): The relative roles of biotic and abiotic pollination

Rewardless orchid species are pollinated by deception and have reduced reproductive success. Those species that present self-compatibility followed by alternative mechanisms of autogamy may compensate for such episodic allogamy events. This is the case for rain-assisted self-pollination, a rare mechanism in Orchidaceae. In this study, we investigated the reproductive biology of two terrestrial species: Cyrtopodium hatschbachii and C. paludicolum. We studied their reproductive systems, examined pollinators and male and female successes, and ascertained if rain promotes self-pollination. Both species are allogamous but self-compatible. However, natural fruit set is low, not exceeding 1.40% during 4 years of observation. This may result from the reduced frequency of pollinator visitation followed by a high pollinator specificity, because each species was pollinated by a single specialist bee. Although spontaneous selfing did not occur, we verified fruits formed by rain action. The evaporation of the water droplet that accumulates on stigma helps the pollinarium to slip back, promoting selfing. This occurred in approximately 10% of C. hatschbachii fruits, but in C. paludicolum it was recorded only once during three observation years. This indicates that the phenomenon happens at random and may vary temporally. Nevertheless, rain may confer additional fruit set, especially under circumstances of pollinator limitation. Overall, we confirmed deception as a strategy for this understudied genus. In addition, we report a further two cases of rain-assisted self-pollination for Cyrtopodium, for species from the center of diversity of the genus, indicating that it may be more widespread than previously thought.     

Effects of nectar robbing on pollinator behavior and plant reproductive success of Pitcairnia angustifolia (Bromeliaceae)

Nectar robbers may have direct and indirect effects on plant reproductive success but the presence of nectar robbing is not proof of negative fitness effects. We combined census data and field experiments to disentangle the complex effects of nectar robbing on nectar production rates, pollinator behavior, pollen export, and female reproductive success of Pitcairnia angustifolia. Under natural conditions flowers were visited by four different animal species including a robber‐like pollinator and a secondary robber. Natural levels of nectar robbing ranged from 40 to 100%. Natural variation in nectar robbing was not associated with fruit set in any year whereas seed set was weakly positively associated for 1 year only. Artificial nectar robbing did not increase nectar production or concentration, did not affect the behavior of long‐billed hummingbirds, and when faced with artificially robbed flowers, these visitors behaved as secondary nectar robbers. The number of stigmas within a patch that received pollen dye analogs and the average distance traveled by these analogs were not significantly different between robbing treatments (robbed flowers versus unrobbed flowers), but the maximum distance traveled by these pollen analogs was higher when nectar robbing was not prevented. Overall, the proportion of robbed flowers on an inflorescence had a neutral effect to a weak positive effect on the reproduction of individual plants (i.e. positive association between nectar robbing and fruit set in 2002) even when it clearly changed the behavior of its most efficient pollinator potentially increasing the frequency of nectar robbing within a plant.     

Effects of pollen quality and quantity on pollen limitation in Crataegus monogyna (Rosaceae) in NW Spain

Pollen limitation occurs when plants produce less fruits and/or seeds than they would with adequate pollen receipt. If the addition of cross-pollen to stigmas increases fruit/seed production, it is interpreted as an evidence of pollen limitation. Much of the limitation may be associated with the quality rather than quantity of pollen; however, most studies do not discriminate between the two, which may lead to misinterpretation of the results. We studied the effects of quality and quantity of pollen on the reproduction of a northern Spanish population of Crataegus monogyna. The treatments included self- and cross-pollination, and supplementation to open and bagged flowers. The response variables considered were number of pollen grains per stigma, pollen tubes per style, and initial and final fruit set. In the Cantabrian range, C. monogyna requires insect pollinators to set fruit and is partially self-incompatible. We found that the number of pollen tubes did not differ between cross- and self-pollination treatments; however, self-pollinated flowers set less fruits than flowers that received pure cross-pollen or were supplemented with both cross- and self-pollen. The experimental design allowed us to infer qualitative rather than quantitative pollen limitation. Comparison of the number of pollen grains and tubes, and initial and final fruit set among pollination treatments suggested post-zygotic embryo selection against selfed progeny.     

Beyond the pollination syndrome: nectar ecology and the role of diurnal and nocturnal pollinators in the reproductive success of Inga sessilis (Fabaceae)

Inga species present brush‐type flower morphology allowing them to be visited by distinct groups of pollinators. Nectar features in relation to the main pollinators have seldom been studied in this genus. To test the hypothesis of floral adaptation to both diurnal and nocturnal pollinators, we studied the pollination ecology of Inga sessilis, with emphasis on the nectar secretion patterns, effects of sequential removals on nectar production, sugar composition and the role of diurnal and nocturnal pollinators in its reproductive success. Inga sessilis is self‐incompatible and pollinated by hummingbirds, hawkmoths and bats. Fruit set under natural conditions is very low despite the fact that most stigmas receive polyads with sufficient pollen to fertilise all ovules in a flower. Nectar secretion starts in the bud stage and flowers continually secreting nectar for a period of 8 h. Flowers actively reabsorbed the nectar a few hours before senescence. Sugar production increased after nectar removal, especially when flowers were drained during the night. Nectar sugar composition changed over flower life span, from sucrose‐dominant (just after flower opening, when hummingbirds were the main visitors) to hexose‐rich (throughout the night, when bats and hawkmoths were the main visitors). Diurnal pollinators contributed less than nocturnal ones to fruit production, but the former were more constant and reliable visitors through time. Our results indicate I. sessilis has floral adaptations, beyond the morphology, that encompass both diurnal and nocturnal pollinator requirements, suggesting a complementary and mixed pollination system.     

Efficient pollination and high reproductive success in two Brazilian Spiranthinae orchids: Insights on the evolutionary history of pollination within the Pelexia clade

The pollination process and breeding system of Pachygenium bonariense and Pachygenium pteryganthum were documented for native populations from Rio Grande do Sul, Brazil. The flowers secrete nectar (on average with 65% to 67% of the concentration of total sugars), which is concealed into an incurved spur. Both species are pollinated by native, large Apidae bees, and the pollinaria adhere under the ventral surface of their labrum while bees try to reach the nectar. P. bonariense is pollinated by native bumblebees Bombus (Fervidobombus) morio and P. pteryganthum by females of oil-gathering Centris (Melacentris) sp. Both orchid species are self-compatible but pollinator-dependent (i.e., unable to set fruit and seed in the absence of pollinators). They also displayed high natural reproductive success (from 22% to 90%) and Nilsson's male efficiency factors ranging from 2.2 to 3.6, suggesting that, during our observations, more than one flower was pollinated for each flower acting as the pollen donor. To enrich the discussion, additional unpublished data on the pollination and breeding system of other orchid species of the Pelexia clade (the Spiranthinae group containing the studied species) are also presented and briefly illustrated. A phylogenetic framework of the clade is used to ascertain well-supported evolutionary scenarios for pollination-related floral traits (viscidium, protandry and a group of pollinators). The characteristic “dorsally adhesive” viscidium can be considered a synapomorphy for the subclade formed by the genera Cyclopogon, Veyretia, Sarcoglottis, Pachygenium, Pelexia and Brachystele, with a reversal to the plesiomorphic condition (ventrally adhesive) at the latter genus. Pollination by bees is likely the rule within the subclade and may be regarded as plesiomorphic for the whole Spiranthinae subtribe.     

Systematic revision of Neotropical Achalcus and a related new genus (Diptera: Dolichopodidae, Achalcinae) with comments on their phylogeny, ecology and zoogeography

A revision of the species of the subfamily Achalcinae of the Neotropical region is presented. A new genus, Australachalcus gen. nov. , is erected based on the presence of six dorsocentral bristles and synapomorphies in the hypopygium. Eight new Achalcus species ( bilineatus , brevicornis , costaricensis , cyanocephalus , maculipennis , micromorphoides , niger , tibialis ) and eight new Australachalcus species ( acornis , browni , cummingi , incisicornis , pseudorobustus , robustus , setosus , variabilis ) are described, and Achalcus albipalpus Parent, A. brevinervis Van Duzee, A. longicornis Van Duzee and Enlinia edwardsae (Van Duzee) are newly referred to Australachalcus and redescribed. A key to males of all species and females of eight species is provided. One Palaearctic species, Achalcus melanotrichus Mik, and nine New Zealand Achalcus species ( chaetifemoratus , luteipes , medius , minor , minusculus , minutus , nigroscutatus , relictus , separatus ) described by Parent are also transferred to Australachalcus . In Neotropical Achalcinae, three Achalcus and more than four different Australachalcus species groups can be distinguished, whereas all Holarctic Achalcus species belong to the single A. flavicollis species group. In the Neotropics, Australachalcus is nearly entirely confined to Chile, whereas Achalcus is recorded from Costa Rica, Venezuela, Ecuador, Colombia, Peru and Chile, with no or little overlap in distribution between the separate species groups. Beyond Chile, most achalcine species treated here were collected at higher altitudes. Most Chilean species are only active during the southern summer, whereas in Costa Rica achalcine representatives are encountered throughout the year.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 27–73.     

Microhabitat Selection by Antlion Larvae, Myrmeleon Crudelis: Effect of Soil Particle Size on Pit-Trap Design and Prey Capture

Selective pressure for choosing an adequate habitat should be strong in semisedentary animals because they have limited mobility once established. I examined microhabitat preferences and the adaptive value of these preferences in the antlion larva Myrmeloen crudelis, a semisedentary insect that digs pit traps in soils to capture small arthropods. I tested the habitat preferences of M. crudelis between two soil types in a tropical dry forest of Costa Rica. Specifically, I compared the soil particle composition size within and outside antlion aggregations and manipulated the availability of fine- and coarse-grained soil to assess how differences in soil grain size affect the design and performance of larval traps. Adjacent to antlion pits the soil was composed of a greater proportion of fine-grained particles (2 mm) than soil 1 m away from the pits. A set of experiments demonstrated that (1) in the presence of equal availability of fine- and coarse-grained soils, all larvae built their pits in fine-grained soil; (2) the larvae required less time to start and finish traps in fine-grained soil; (3) the larvae constructed larger and deeper pits in fine-grained soil; and (4) prey capture increased greatly in fine-grained traps compared with coarse-grained traps. Antlion larvae respond to variations in the proportion of fine particles in the soil, suggesting that antlion aggregations result from an active microhabitat selection. The preference for fine-grained soils is adaptive since pits constructed in such substrate are functional for longer periods and much more successful in trapping prey than pits in coarse-grained soil. Sit-and-wait predators that use sessile traps are spatially constrained to track prey abundance. Therefore, the ability to detect and select microhabitats with better conditions that enhance capture success may be under strong selection for this type of organism.     

The pollination seesaw of Rhynchospora cephalotes (L.) Vahl (Cyperaceae): Influence of plant location on the role of wind and insects as pollen vectors

Ambophily (wind and insect pollination) has been reported for some genera of the typically wind-pollinated family Cyperaceae, including the genus Rhynchospora. The significance of wind and insect pollination can vary, depending on local microclimatic conditions. Rhynchospora cephalotes is an ambophilous species that can grow under different environmental conditions, either along forest edges or inside forest fragments. This study, therefore, tests the hypothesis that (a) there is greater contribution by wind than insects to fruit set of the individuals at the forest edge and (b) there is greater contribution by insects than wind to fruit set of the individuals inside the forest. Field work was carried out in a fragment of Atlantic Forest in northeastern Brazil. We quantified the number of visits by insects, wind velocity and the reproductive success (fruit set) of R. cephalotes provided by wind and insects through exclusion experiments performed at the edge of and inside the forest. We observed a greater number of visits per day by pollinating bees to individuals inside the forest (36.83 ± 7.46) than to individuals at the edge (16.66 ± 6.53). The wind speed was significantly higher at the edge (1.71 ± 0.46 m/s) than inside the forest (0.97 ± 0.18 m/s). Bees and wind are both pollen vectors of R. cephalotes, but bees were the pollen vector that contributed most to fruit set (63.3%) for individuals inside the forest, whereas wind was the primary vector for individuals at the forest edge (76.6%). This seesaw in importance of each pollen vector in the two different environments guarantees high fruit set in R. cephalotes under different microclimatic conditions.