Taxonomic and population differentiation of mitochondrial diversity inPinus banksiana andPinus conforta

We have studied two mitochondrial DNA polymorphisms in 741 individuals from 16 allopatric populations ofPinus banksiana Lamb. andPinus contorta Dougl. Restriction fragments of both polymorphisms distinguished the two species qualitatively, except in aP. Banksiana population whose ancestors were involved in hybridization withP. contorta.COXI-associated restriction fragments were monomorphic within species, whileCOXII-associated restriction fragments were highly variable inP. contorta (H_es=0.68). Population differentiation was substantial inP. contorta (F_st=0.31 among subspecies; mean F_st=0.66 within subspecies) and consistent with predictions for maternally inherited markers. Plant mitochondrial markers appear to be useful for the investigation of seed migration routes, hybridization and introgression, breeding zone designation, and the development of germ plasm conservation sampling strategies.     

Study onPlatanus hispanica Miller pollen content in the air of Seville, southern Spain

The work was carried out using a Cour trap that sampled the air of the city for 8 consecutive years (1987–1994). The pollen ofPlatanus hispanica is the fourth most abundant in the air of Seville (a mean of 11.05% of the total pollen collected). The variation throughout the years in the sum of weekly concentrations ofPlatanus hispanica pollen presents a certain biennial rhythm, in which years of high and low collection of pollen alternate. The starting day of the main pollination period (MPP) is negatively related with the mean of the mean temperatures for February (r=0.73,r ²=0.53,P=0.0398) and is earlier (at the beginning of March) when the mean temperature for February is high, and vice versa. The pattern of pollen variation inPlatanus hispanica remains constant through the years—pollen appears abruptly in high weekly concentrations (> 150 grains/m³) in March (sporadically at the beginning of April), with a week of maximum pollen emission (WMPE) in which more than 50% of the annual pollen is collected (in 6 of the 8 years), and a main pollination period (MPP) of 2 or 3 weeks (except in 1989 when it was 5 weeks). In every year (except 1989), weekly mean temperatures increased during the MPP, the duration of which depends on mean temperature and mean rainfall: mean temperatures > 16°C and absence of rainfall shorten the MPP, while lower temperatures and presence of rainfall lengthen it. The meteorological conditions most often found during the WMPE are mean temperatures > 15°C and rainfall absent or almost so.     

Mycorrhiza-like interaction by Morchella with species of the Pinaceae in pure culture synthesis

 Isolates from two species of Morchella were tested for ability to form mycorrhizae in pure culture synthesis with Arbutus menziesii, Larix occidentalis, Pinus contorta, Pinus ponderosa, andPseudotsuga menziesii. Ectomycorrhizal structures (mantle and Hartig net) formed with the four species of the Pinaceae but not with A. menziesii. Results are compared to previous studies on morel mycorrhizae and discussed in an ecological context. Accepted: 23 October 1999     

Comparative capitular morphology and anatomy of Coreopsis L. and bidens L. (compositae), including a review of generic boundaries

The capitular and floral morphology and anatomy ofBidens L. andCoreopsis L. were studied. All the North American species ofCoreopsis were studied. Selected species ofBidens from North and South America andCoreopsis from South America were included. The results were compared with previous observations on African species ofBidens (incl.Coreopsis). Emphasis was given to character states of the ray florets, paleae, stylearm apices, outer phyllaries, achenes, and pollen grains. Some of the character states are unique features ofCoreopsis, e.g., globular and elongately conical receptacles, deltoid outer phyllaries, truncate and indistinctly 3–5-dentate, 3–4-lobed ray florets, narrowly spathulate paleae, subulate paleae with linear-filiform upper half, hairy and apically 3-cleft paleae, truncate, convex or shallowly conical stylearm apices with the sweeping hairs limited to the area above the stigmatic surfaces and the orbicular to circular achenes. The cylindric setaceous pappus bristles so commonly encountered inBidens are unknown inCoreopsis. The pappus bristles inCoreopsis are paleaceous but similar, though thicker ones are also found in African species ofBidens (incl.Coreopsis) with winged achenes. Twin-celled hairs (setulae) with differing degrees of wall thickness are found on the achenes ofCoreopsis sect.Pseudoagarista (Mexico and South America),Coreopsis sect.Pugiopappus (California), AfricanBidens with winged achenes (e.g.,B. prestinaria, B. macroptera) and some North AmericanBidens (e.g.,B. aristosa). Similar sclerotic parenchyma make up the achenial wings of species in both genera. These may be interpreted as homologous structures, indicating the underlying similarity of these taxa and their derivation from a common ancestral stock.     

Peculiarities of sexual reproduction inFagus crenata forests in relation to annual production of reproductive organs

Annual production rates of reproductive organs inFagus crenata forests in the lower area of the species' range were studied using 10 litter traps in 1980–1986. The production rates of dispersed pollen were estimated by multiplying the number of fallen male inflorescences per ha per year by the mean amount of pollen per inflorescence before anthesis. Large annual fluctuations in the production rates of male and female inflorescences were recognized, whereas their annual trends were synchronized with each other. Pollen production rates were within the range 1.0–6900 (mean: 1630)×10⁹ha⁻¹ yr⁻¹, the maximum/minimum ratio attaining 7000.F. crenata was the lowest producer of pollen among seven tree species studied: the number of pollen grains equivalent to a single ovule was in the range 6.0–14×10⁴. Furthermore, the mean dry weight of a single pollen grain (3.77×10⁻⁵mg) was higher than for wind-pollinated species. Three factors seemed to cause the low seed fertility ofF. crenata. The dry-matter production rate in the best seed year reached 3252 kg ha⁻¹ yr⁻¹, of which pollen accounted for 259 kg ha⁻¹ yr⁻¹. Unproductive years with less than 10% of the maximum production occurred four times in a 7-yr period. In such years there were fewer male and female inflorescences, and more fruit dropped as a result of insect damage. Lower nut dissemination would play an important role in suppressing any increase in nut predators, and fewer flowers would be produced to avoid wastage of photosynthates in a cool-temperate climate.     

Antibacterial effects of knotwood extractives on paper mill bacteria

Hydrophilic knotwood extracts from 18 wood species were assessed in disc diffusion and liquid culture tests for antibacterial effects against three species of paper mill bacteria. The Pinus sylvestris, P. resinosa, P. contorta, and P. banksiana extracts decreased or inhibited bacterial growth. The susceptibility order was P. sylvestris > P. resinosa > P. contorta > P. banksiana, correlating with the concentrations of pinosylvin and pinosylvin monomethyl ether in these wood species. Also, Pseudotsuga menziesii and Thuja occidentalis extracts had a small inhibitory effect. The Gram-positive Bacillus coagulans was more susceptible to the extracts than the Gram-negative Burkholderia multivorans and Alcaligenes xylosoxydans. The main components in the Pinus knotwood extracts were pinosylvin monomethyl ether and pinosylvin, suggesting these to be the active components. Therefore, pure pinosylvin, pinosylvin monomethyl ether, and dihydro-pinosylvin monomethyl ether were also tested. All compounds showed antibacterial effects. However, higher concentrations were needed for these pure compounds than for the knotwood extracts. Pinosylvin had stronger antibacterial effects than pinosylvin monomethyl ether. This work shows that knotwood extracts, especially from Pinus species, have a potential for use as natural biocides in papermaking.     

Effects of past logging and grazing on understory plant communities in a montane Colorado forest

Throughout Pinus ponderosa–Pseudotsuga menziesii forests of the southern Colorado Front Range, USA, intense logging and domestic grazing began at the time of Euro-American settlement in the late 1800s and continued until the early 1900s. We investigated the long-term impacts of these settlement-era activities on understory plant communities by comparing understory composition at a historically logged and grazed site to that of an environmentally similar site which was protected from past use. We found that species richness and cover within functional groups rarely differed between sites in either upland or riparian areas. Multivariate analyses revealed little difference in species composition between sites on uplands, though compositional differences were apparent in riparian zones. Our findings suggest that settlement-era logging and grazing have had only minor long-term impacts on understories of upland Front Range P. ponderosa–P. menziesii forests, though they have had a greater long-term influence on riparian understories, where these activities were likely the most intense. This article was written and prepared by US Government employees on official time, and therefore it is in the public domain and not subject to copyright.     

Altitude trends in conifer leaf morphology and stable carbon isotope composition

The natural ratio of stable carbon isotopes (δ¹³C) was compared to leaf structural and chemical characteristics in evergreen conifers in the north-central Rockies, United States. We sought a general model that would explain variation in δ¹³C across altitudinal gradients. Because variation in δ¹³C is attributed to the shifts between supply and demand for carbon dioxide within the leaf, we measured structural and chemical variables related to supply and demand. We measured stomatal density, which is related to CO₂ supply to the chloroplasts, and leaf nitrogen content, which is related to CO₂ demand. Leaf mass per area was measured as an intermediate between supply and demand. Models were tested on four evergreen conifers: Pseudotsuga menziesii, Abies lasiocarpa, Picea engelmannii, and Pinus contorta, which were sampled across 1800 m of altitude. We found significant variation among species in the rate of δ¹³C increase with altitude, ranging from 0.91‰ km^–1 for A. lasiocarpa to 2.68‰ km^–1 for Pinus contorta. Leaf structure and chemistry also varied with altitude: stomatal density decreased, leaf mass per area increased, but leaf nitrogen content (per unit area) was constant. The regressions on altitude were particularly robust in Pinus contorta. Variables were derived to describe the balance between supply and demand; these variables were stomata per gram of nitrogen and stomata per gram of leaf mass. Both derived variables should be positively related to internal CO₂ supply and thus negatively related to δ¹³C. As expected, both derived variables were negatively correlated with δ¹³C. In fact, the regression on stomatal density per gram was the best fit in the study (r ²=0.72, P<0.0001); however, the relationships were species specific. The only general relationship observed was between δ¹³C and LMA: δ¹³C (‰)=–32.972+ 0.0173×LMA (r ²=0.45, P<0.0001). We conclude that species specificity of the isotopic shift indicates that evergreen conifers demonstrate varying degrees of functional plasticity across environmental gradients, while the observed convergence of δ¹³C with LMA suggests that internal resistance may be the key to understanding inter-specific isotopic variation across altitude. Received: 1 June 1999 / Accepted: 2 November 1999     

Development of a simple molecular marker specific for detecting the self-compatible<Emphasis Type="Italic">S4′</Emphasis> haplotype in sweet cherry (<Emphasis Type="Italic">Prunus avium</Emphasis> L.)

In sweet cherry (Prunus avium L.), theS4′ haplotype, characterized by a self-incompatibility (SI) defect in pollen, is self-compatible and is derived from the self-incompatibleS4 haplotype by x-ray mutagenesis.SFBs (S haplotype-specific F-box protein genes) have been found to associate with pollen determinant of SI. This report identified theSFB4′ of the self-compatibleS4′ haplotype. The alignment of the sequences ofSFB4′ andSFB4 by the BLAST program revealed a 4-bp deletion inSFB4′, which is TTTA. The sequence polymorphism generated by the TTTA deletion inSFB4′ was exploited to develop a simple molecular marker specific for detecting theS4′ but not theS4 haplotype. The simple marker specific to theS4′ haplotype can be visualized directly on an agarose gel, so it can be immediately applied to a marker-assistant cherry-breeding program. Thus, this work provides a practical molecular marker for cherry breeding. Principal author. An erratum to this article is available at .     

Water flux and canopy conductance of natural versus planted forests in Patagonia, South America

In NW Patagonia, South America, natural shrublands and mixed forests of short Nothofagus antarctica (G. Forst.) Oerst. trees are currently being replaced by plantations with Pseudotsuga menziesii (Mirb) Franco. This land use change is controversial because the region is prone to drought, and replacement of native vegetation by planted forests may increase vegetation water use. The goal of this study was to examine the physiological differences, especially the response of water flux and canopy conductance to microclimate, that lead to greater water use by exotic trees compared to native trees. Meteorological variables and sapflow density of P. menziesii and four native woody species were measured in the growing season 2005–2006. Canopy conductance (gc) was estimated for both the exotic (monoculture) and native (multi-species) systems, including the individual contributions of each species of the native forest. Sapflow density, stand-level transpiration and gc were related to leaf-to-air vapor pressure difference (VPD). All native species had different magnitudes and diurnal patterns of sapflow density compared to P. menziesii, which could be explained by the different gc responses to VPD. Stomatal sensitivity to VPD suggested that all native species have a stronger stomatal control of leaf water potential and transpiration due to hydraulic limitations compared to P. menziesii. In conclusion, differences in water use between a P. menziesii plantation and a contiguous native mixed forest of similar basal area could be explained by different gc responses to VPD between species (higher sensitivity in the native species), in addition to particular characteristics of the native forest structure.     

Pollen evolution and systematics inAnnonaceae with special reference to the disulcate Australian endemic genera

All genera ofAnnonaceae endemic in Australia (Ancana, Fitzalania, Haplostichanthus) show almost exactly the same type of disulcate (disulculate) pollen with intact exine extending over the sulci. Tetrad stages inHaplostichanthus andAncana reveal a latudinal subequatiorial orientation of the two sulci at the proximal hemisphere. Sometimes they fuse into a ±zonosulcate aperture.Fissistigma pollen grains are ±globose and have a flattened pole with a central elevation and a concentric groove, covered by a somewhat reduced exine. This palynological characters give further support for separating the generaAncana andFissistigma. Germination was observed inHaplostichanthus where the pollen tube emerges at one of the two sulci and inFissistigma where the flattened part breaks up during germination. The aperture types described here are obviously transitional stages between aperturate and inaperturate pollen grains and are discussed in regard to pollen evolution.     

Trade-offs between the persistence of foliage and productivity in two<Emphasis Type="Italic"> Pinus</Emphasis> species

We investigated interspecific variation in leaf lifespan (persistence) and consequent differences in leaf biochemistry, anatomy, morphology, patterns of whole-tree carbon allocation and stand productivity. We tested the hypothesis that a species with short-lived foliage, Pinus radiata D. Don (mean leaf lifespan 2.5 years), grows faster than P. pinaster Ait., a species with more persistent foliage (leaf lifespan 5.6 years), and that the faster growth rate of P. radiata is associated with a greater allocation of nitrogen and carbon to photosynthetic tissues across a range of scales. In fully sunlit foliage, the proportion of leaf N in the major photosynthetic enzyme Rubisco (ribulose-1, 5-bisphosphate carboxylase) was greater in P. radiata than in P. pinaster, and, in mid-canopy foliage, the proportion of leaf N in thylakoid proteins was greater in P. radiata. A lesser proportion of needle cross-sectional area was occupied by structural tissue in P. radiata compared to P. pinaster. Foliage mass in stands of P. radiata was 9.7 t ha^–1 compared with 18.2 t ha^–1 in P. pinaster while leaf area index of both species was similar at 4.6 m² m^–2, owing to the compensating effect of differences in specific leaf area. Hence trade-offs between persistence and productivity were apparent as interspecific differences in patterns of whole-tree carbon allocation, needle morphology, anatomy and biochemistry. However, these interspecific differences did not translate into differences at the stand scale since rates of biomass accumulation were similar in both species (P. radiata 6.9±0.9 kg year^–1 tree^–1; P. pinaster 7.4±0.9 kg year^–1 tree^–1). The similarities in performance at larger scales suggest that leaf area index (and radiation interception) determines growth and productivity. Received: 13 July 1999 / Accepted: 31 January 2000     

Zur Skulptur der Pollen-Exine bei drei Centrospermen (Gisekia,Limeum, Hectorella), bei Gyrostemonaceen und Rhabdodendraceen

The exine-sculpturing of pollen fromGisekia africana (Gisekiaceae),Limeum argute-carinatum (Molluginaceae) andHectorella caespitosa (Hectorellaceae) with supratectate spinulae and an anulopunctate tectum is in accordance with the manyCentrospermae investigated so far with the SEM. Pollen ofRhabdodendron macrophyllum and three species fromGyrostemonaceae do not exhibit these surface details. WhileRhabdodendron with a finely reticulate exine probably fits intoRutaceae, the unique columellaless exine ofGyrostemonaceae has no direct counterpart, neither inCentrospermae nor inCapparales, the two orders to which this family was allied. The genera ofGyrostemonaceae can be distinguished by different arrangements of minute pollen surface details.

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Zur Skulptur der Pollen-Exine bei drei Centrospermen (Gisekia, Limeum, Hectorella), bei Gyrostemonaceen und Rhabdodendraceen

     

The Effect of Temperature on Reproduction in FivePrimulaSpecies

Primula vulgarisHuds.,P. verisL.,P. frondosaJanka, and threepopulations ofP. farinosaL. were legitimately and illegitimatelypollinated, and the self-fertileP. scoticaselfed and cross-pollinatedand then subjected to uniform temperature conditions of 6, 15or 26 °C for 4 d before gynoecia were examined for pollengermination and pollen tube growth, or plants progressed toseed set at 15 °C, after which seeds were weighed, germinated,and seedlings grown on. The temperature responses of pollengermination and pollen tube growth were not always congruent,and varied between species, populations, and often between morphs(pin and thrum) in the distylous species. Nevertheless, optimaltemperature responses tended to be lower for vernal species(P. vulgarisandP. veris) and for subarcticP. scoticathan forlater flowering montane species. However, no relationship wasfound between pollen temperature response, and fertility. Thegreatest seed set occurred after legitimate pollination at 15°C in most cases; a flowering temperature of 26 °C tendedto impede seed set, except forP. scoticaand the low altitudepopulation ofP. farinosa. InP. veris, P. frondosaand the highaltitude population ofP. farinosa,some illegitimate pollen germinationand pollen tube growth occurred at 26 °C, but this did notlead to increased within-morph seed set in these self-incompatiblespecies at this relatively high temperature. Temperature atflowering frequently affected average seed weight, and inP.verisand two populations ofP. farinosathis attribute may havebeen influenced by seed number, the average seed weight of few-seededcapsules tending to be greater than for many-seeded capsules.A high seed weight might mitigate the disadvantageous effectsof low fecundity resulting from interactions with floweringtemperature. However, inP. vulgarisandP. scoticainteractionsbetween flowering temperature and seed weight may have other,undetermined, causes. The seed of four species germinated leastwell in standard conditions when set following a flowering temperatureof 6 °C, which tends to support the hypothesis that temperatureat flowering can affect seed physiology; in contrast the seedof the two upland populations ofP. farinosagerminated leastwell after flowering at 26 °C. We conclude that much morework is needed on interactions between temperature and reproductiveefficiency, but that preliminary indications suggest that aglobal increase in temperature at flowering might adverselyaffect the quantity and quality of seed set in some species.Copyright1998 Annals of Botany Company Distyly, pollen tube,Primula farinosa, Primula frondosa, Primula scotica, Primula veris, Primula vulgaris,reproduction, seed set, temperature.     

Recovery of aSasa tsuboiana population after mass flowering and death

The recovery process of aSasa tsuboiana population after a mass flowering and death in 1977 was investigated by 15 years of observation in the Hira Mountains, Kinki district, western Japan. Seed production was high (6600–13 800 seeds m⁻² inSasa plots and 3900 seeds m⁻² in a forest plot) but emergent seedling density was low (14–21 seedlings m⁻²), probably because of seed predation byMicrotus montebelli occurring between seed shedding and the next spring. The seedling density had decreased further by the next year and theS. tsuboiana population recovered from only a limited number of seedlings. In spite of such a low initial density, theS. tsuboiana population was able to regenerate successfully and attained the previous full stand height in 7–16 years.Miscantbus sinensis invaded and delayed the recovery ofS. tsuboiana in one plot, butS. tsuboiana became dominant as it caught up with the height ofM. sinensis. Seedling growth patterns, such as frequent tillering, the onset of rhizome extension in the early stage of seedling growth and frequent culm production from rhizomes, played important roles in the successful regeneration ofS. tsuboiana.     

Stamen movements in flowers ofOpuntia (Cactaceae) favour oligolectic pollinators

Opuntia brunneogemmia andO. viridirubra occur sympatrically in the Serra do Sudeste, Rio Grande do Sul, Brazil. Their flowers have 450–600 thigmonastic stamens and provide large amounts of pollen and nectar for bees. Bees of 41 species were registered at the flowers ofO. brunneogemmia and 30 at the flowers ofO. viridirubra. Females of three oligolectic species are the only effective pollinators:Ptilothrix fructifera (Anthophoridae),Lithurgus rufiventris (Megachilidae), andCephalocolletes rugata (Colletidae). During their visits inOpuntia-flowers, bees touch the filaments and stimulate the movement of the stamens to the centre of the flower. At the end of this movement, the anthers are densely packed around the style. As a consequence the pollen is presented in an easily accessible upper layer of anthers and various, nearly inaccessible lower layers. The lower layers contain about 80% of the pollen reward. Only females of the three oligolectic pollinators exploit the pollen from the lower layers and reach the nectar furrow. Therefore, through their stamen movements,Opuntia flowers hide most of their pollen from flower visitors but favour effectively pollinating, oligolectic bees.     

Persistence of<Emphasis Type="Italic">Pseudotsuga menziesii</Emphasis> (Douglas-fir) in temperate coniferous forests of the Pacific Northwest Coast,USA

Old-growthPseudotsuga-Tsuga forests of the Pacific Northwest Coast of North America are characterized by the presence of large, old trees ofPseudotsuga menziesii var.menziesii (Douglas-fir). Colonizing soon after a stand-replacing disturbance,P. menziesii persists in these forests, coexisting for centuries with the late-successional species.P. menziesii survives by maintaining emergent status in the uppermost part of the forest canopy, above the crowns of competing late-successional species. After reaching maximum tree height and crown size,P. menziesii maintains shoots and foliage of the established crown by epicormic shoot production. In this review, we propose that attaining emergent status in the upper canopy combined with the process of crown maintenance contributes to the persistence ofP. menziesii into later stages of succession, making this species a long-lived pioneer that between infrequent disturbances can coexist with late-successional species for centuries.     

Population structure in two species of the reef goby<Emphasis Type="Italic"> Gnatholepis</Emphasis> (Teleostei: Perciformes) among four South Pacific island groups

Individuals of two species in the reef goby genus Gnatholepis ( G. anjerensis and G. scapulostigma) were sampled from across the South Pacific, including the Society Islands, the Tuamotu Archipelago, the Cook Islands, and Fiji. The two species examined differ in habitat preference, with one (G. anjerensis) preferring shallow, lagoon habitats, and the other (G. scapulostigma) usually occupying habitats outside the lagoons. Partial sequence of the mitochondrial ND2 gene (451–483 base pairs) was determined for 267 individuals and analyzed for demographic structure, population structuring, and correlation of population structure with geographic distance between populations. Both species exhibit demography characteristic of population expansion in the Pleistocene, approximately 88,000 years ago for G. anjerensis and 237,000 years ago for G. scapulostigma. Comparisons of _st indicate more structuring among populations of G. anjerensis than G. scapulostigma, although only G. scapulostigma shows a significant correlation between geographic distance and population divergence. It is postulated that the population expansions in Gnatholepis species are related to Pleistocene fluctuations in sea level, and that the ecology of G. anjerensis populations may contribute to their greater population structuring.Communicated by P.F. Sale     

Notes on the pollination mechanisms ofMoraea inclinata andM. brevistyla (Iridaceae)

Individual flowers ofMoraea inclinata are nectariferous and last about six hours. They appear to be pollinated largely by bees in the familyHalictidae (Lasioglossum spp.,Nomia spp.,Zonalictus) and to a lesser extent by bees in the familyAnthophoridae (Amegilla). The mechanism of bee-pollination inM. inclinata is the Iris type; i.e., each flower consists of three pollination units (an outer tepal, a partly exserted anther, and the opposed style branch which terminates in a pair of petal-like crests). Bees rarely visit more than one pollination unit per flower. Transferral of pollen to the bee is passive and nototribic although all bees collected on the flowers were female and 55% of the bees carried pollen loads with 2–5 pollen taxa in their scopae.Moraea brevistyla flowers are nectariferous but lack scent and last two days. They are visited infrequently by bees and only one femaleLasioglossum spec. carried the pollen ofM. brevistyla. Unlike flowers ofM. inclinata those ofM. brevistyla deposit pollen only on the head and thorax. Bee-mediated autogamy in both species is avoided due to the erratic foraging patterns of the bees and the flexibility of each stigma lobe as the bee backs out of the flower. Approximately 2–4 flowers in the inflorescences of both species (6–8 flowers/infloresence) develop into capsules.     

Pollen morphology and its taxonomic significance of the monotropoideae (Ericaceae)

A palynological survey, including LM, SEM and TEM is presented for eight genera, nine species and 76 samples of the Monotropoideae which is composed of 10 genera and 13 species. On the basis of the aperture number and shape, the following six pollen types are recognized: 1) 3-colp(oroid) ate—Allotropa, 2) 2-colpor(oid)ate—Monotropsis andHemitomes, 3) 4-colpor(oid)ate—Pterospora, Sarcodes andPleuricospora, 4) 2- and 3-colpor(oid)ate—Monotropa hypopitys, 5) 3-colporate and-porate —Monotropa uniflora, 6) 3- to 4- porate—Monotropastrum humile. Relationships among taxa within the Monotropoideae are illustrated on the palynological characters including the aperture type, exine sculpture and structure. Both pollen grains with two and four apertures have evolved independently from pollen grains with three apertures, according to the infraspecific variation of aperture numbers and the usual occurrence of three-aperturate pollen grains in the Ericaceae. 3-colp(oroid)ate pollen ofAllotropa is the most primitive and occupies an isolated position in the subfamily, on the other hand 3- to 4-porate pollen ofMonotropastrum humile is most advanced in the subfamily, probably even in the Ericaceae. A reduced sexine inMonotropastrum humile appears to be a specialized pollen character. Infraspecific geographical difference in palynological characters is revealed in two species ofMonotropa for the first time.M. hypopitys is basically characterized by pollen grains with two apertures in the New World vs. three apertures in the Old World.M. uniflora is basically characterized by 3-porate pollen in the New World vs. 3-colporate in the Old World. In both species pollen grains with more primitive characters usually occur in the Old World.