Dust seed production and dispersal in Swedish Pyroleae species

Dust seeds are the smallest seeds in angiosperms weighing just about a few micrograms. These seeds are characteristic of most orchids, and several studies have been performed on seed features, fecundity and dispersal of orchid dust seeds. In this study we examine seed features, seed production and seed dispersal in another plant group with dust seeds, the Pyroleae (Monotropoideae, Ericaceae), focusing on six species: Pyrola chlorantha, P. minor, P. rotundifolia, Chimaphila umbellata, Moneses uniflora and Orthilia secunda. Seed production per capsule among these species was in the range between ca 1000 and 7800, and seed production per capsule bearing shoot was in the range between ca 7000 and 60 000. Combining our results with published information on pollen‐ovule ratios suggests that these Pyroleae species have a generally efficient pollination system. The most fecund species was P. minor, the only species among the investigated that is probably largely self‐pollinating. The investigated Pyroleae species have a seed production comparable to the less fecund orchid species. We studied seed dispersal in the field in one of the species, P. chlorantha. Despite the extremely small and potentially buoyant seeds, the vast majority of seeds are deposited close to the seed source, within a few meters. Further studies on the recruitment ecology of the investigated Pyroleae species are currently under way.     

Studies in the morphology and systematics of berberidaceae

The floral vasculature in three allied genera,Plagiorhegma, Jeffersoria andAchyls is investigated, and the results are compared with those ofEpimedium andVancouveria which are related closely toPlagiorhegma andJeffersonia. The vasculature in the receptacle ofPlagiorhegma andJeffersonia is similar, but that ofAchlys is much simpler. Slightly different trace patterns are observed in the sepals ofPlagiorhegma andJeffersonia. InJeffersonia, the 3-trace condition leaving 2 or 3 gaps is most frequently observed, but inPlagiorhegma traces of a double nature leaving a single gap are more frequent. The traces to the innermost sepals, petals and stamens are usually of a double nature leaving a single gap in both genera. Regular division and fusion are not observed in the receptacular stele. The vascular differentiation between sepals and petals is more advanced inPlagiorhegma andJeffersonia than inEpimedium andVancouveria. InAchlys, the traces are all staminal and single throughout their course. Two parts recognized in the pistils ofPlagiorhegma, Jeffersonia andAchlys are traversed by independent vasculature. The comparisons of pistil morphology including vasculature ofPlagiorhegma, Jeffersonia, Achlys, Epimedium andVancouveria lead to the interpretation that the pistils are based on the same morphological plan. The probable evolutionary trend in pistil is then suggested in these five genera.     

Splash seed dispersal by raindrops

Splash seed dispersal by raindrops was investigated for plants in southern Japan. Nine families, 10 genera and 19 species were confirmed as raindrop-dispersed plants. The 10 genera were Gentiana, Gratiola, Chrysosplenium, Mazus, Mitella, Ophiorrhiza, Sagina, Sedum, Trigonotis and Veronica. The method of splash rain dispersal in these species was clarified. Raindrop-dispersed species were all small herbaceous plants with a vertical pedicel and an apically opening fresh capsule when the seeds mature. Open capsules were cup-shaped or boat-shaped and can accommodate raindrops easily. The raindrops splashed the seeds from the capsule. In general, the seeds weighed very little, but they were heavier than powder or dust seeds dispersed by wind. A strong negative correlation was found between seed weight and the number of seeds per capsule. In the case of Trigonotis brevipes (Maxim.) Maxim., raindrops were received into the cup-shaped calyx-tube and dispersed the fruitlets. Some species, such as Gentiana thunbergii (G. Don) Griseb., Gentiana zollingeri Fawcett and Ophiorrhiza japonica Blume, had hydroscopic movement capsules that opened widely only when wet. Raindrop-dispersed plants were found in various habitats. For example, some plants grew together on rocks along the mountain torrents where splash water could easily be caught. The results of the laboratory and field experiments indicated that the dispersal distance of seeds by raindrops was 1m or less. For small herbaceous plants, splash dispersal by rain might be an effective and advantageous method of seed dispersal because dispersal is not affected by plant height.     

REPRODUCTIVE ECOLOGY OF JEFFERSONIA DIPHYLLA (BERBERIDACEAE)

The reproductive ecology of Jeffersonia diphylla (L.) Pers. (Berberidaceae) was investigated by studying its breeding system, ovule production, seed set, seed dispersal by ants and seed predation by rodents. This species flowers early in the spring and is facultatively autogamous. In a typical year fruit and seed set is high (90%), however, freezing temperatures from late spring frosts in 1983 and 1985 resulted in low fruit set (7% and 20%, respectively), and reduced seed set in those flowers that produced fruit. No differences in seed set between selfed and outcrossed flowers were observed over a two-yr period (1983–84). Ovule number per capsule increased with plant size as measured by leaf number. Seed set and seed wt were unaffected by leaf number unless leaves were removed after flowering was initiated. Jeffersonia diphylla is myrmecochorous. Ants removed seeds faster when seeds were placed in areas where J. diphylla plants were absent, suggesting that dispersal within J. diphylla populations is ant limited. Moreover, fresh (1 day old) seeds were removed by ants faster than 3 day old seeds. Seed predation by rodents prior to dehiscence from capsules is heavy in large populations (85–90%), and apparently negligible in small populations. Predation of seeds that are released from capsules is heavy (approx. 66%), particularly at night. Overall, seed predators consume about 96% of the seed crop in well established populations, but probably much less in small young populations. Hence, seedling recruitment is likely to be higher in small populations, whereas ramet production from rhizomes is the primary mode of propagation in large ones. The evolution of autogamy, early flowering, and myrmecochory are discussed in light of the results of this study.     

FRUITS AND SEEDS OF TRIBE GORDONIEAE (THEACEAE) FROM THE EOCENE OF NORTH AMERICA

A new genus and species Gordoniopsis polysperma and two new species of Gordonia (Gordonieae, Camellioideae, Theaceae) are described based on fossil fruit and seed remains. These specimens are part of a large flora consisting of various plant organs from the middle Eocene Claiborne Formation in western Kentucky and Tennessee. Gordoniopsis is a five-valved loculicidally dehiscent capsule similar to capsules of Gordonia but differing in having unwinged seeds and a greater number of seeds per locule. The two Gordonia species are among the earliest unequivocal records of the genus and two of only four fossil Gordonia species known with in situ seeds. Two extinct genera, Gordoniopsis and Andrewsiocarpon, and the extant genus Gordonia in the tribe Gordonieae are known from the middle Eocene Claiborne flora, suggesting an early radiation within the tribe. Based on a survey of Recent fruits and seeds we concur with Keng's proposal to merge Laplacea with Gordonia.     

Fitness of natural willow hybrids in a pioneer mosaic hybrid zone

Hybrid fitness is an important parameter to predict the evolutionary consequences of a hybridization event and to characterize hybrid zones. We studied fitness parameters of F₁ and later‐generation hybrids between the lowland species Salix purpurea and the alpine S. helvetica that have recently emerged during colonization of an alpine glacier forefield. Fruit production (number of capsules per catkin and fruit set) did not differ between hybrids and parents, but the number of seeds per capsule of F₁ hybrids was slightly lower than that of later‐generation hybrids and of the parents. Germination rates and seedling growth were tested on three substrates (pH 4.5, 7.0, and 8.0). Germination rates of seeds collected from F₁ hybrids were lower on acid and neutral substrates, but equal at pH 8.0 compared to all other groups, while the seeds from later‐generation hybrids performed as well as the parents on all three substrates. In seedling growth, the colonizer S. purpurea performed better than all other taxa on all three substrates, while hybrids resembled the subalpine species S. helvetica. Results suggest that endogenous selection acts against F₁ hybrids, but favors fitter genotypes in later‐generation hybrids. Exogenous selection via soil pH appears to be weak during seedling establishment. The pioneer vegetation on the glacier forefield may offer sufficient niche space for hybrid seedlings. Owing to the relatively high fitness of the hybrids and the scattered distribution of hybrids and parental individuals on the glacier forefield, this hybrid zone can be assigned to a mosaic model, probably facilitating gene flow and introgression between the parental species. As establishment of the hybrid zone appears to be linked to a colonization process, we propose to call it a pioneer mosaic hybrid zone.     

THE LEVEL OF AGAMOSPERMY IN A NEBRASKA POPULATION OF SPIRANTHES CERNUA (ORCHIDACEAE)

This study investigated reproduction in a prairie population of Spiranthes cernua in eastern Nebraska. Spiranthes cernua reproduces both sexually and asexually, through adventitious embryony. Agamospermous seeds are monoembryonic, polyembryonic, have extruded or free embryos, or lack embryos. These kinds of seeds are extremely rare in Spiranthes vernalis, a close relative that relies exclusively on sexual reproduction. Sexually derived seeds are monoembryonic or lack embryos. One objective was to quantify the amount of sexual reproduction in a natural population. In 1985 and 1986, maximum estimates of sexual seed production per capsule were 19.2% and 33.6%, respectively. Thus, every capsule examined contained a high proportion of agamospermous seeds. In addition, attempts were made to manipulate the reproductive system with experimental crosses performed in the field. The treatments examined agamospermy, autogamy, and outcrossing. In all treatments the majority of seeds in a capsule were agamospermous.     

SEED SET AND SEED PREDATION IN PEDICULARIS FURBISHIAE,A RARE ENDEMIC OF THE ST. JOHN RIVER,MAINE

We measured seed set and losses due to mammal and insect predation during 1983 in seven northern Maine populations of Furbish's Lousewort (Pedicularis furbishiae S. Wats.; Scrophulariaceae), a perennial herb endemic to the banks of the St. John River. Only about a quarter of initiated flowers developed a capsule with a full complement (mean = 25) of seeds. Secondary (lateral) inflorescences matured a smaller proportion of capsules than primary (terminal) inflorescences. About 58% of the initiated flower primordia were aborted or did not develop mature seeds by early September; this was probably not due to pollinator limitation. Several agents could have reduced seed production. Although spittlebugs infested between 3 and 30% (average 20%) of developing flower scapes by population, they did not depress the number of inflorescences initiated or the number of capsules matured. In contrast, 74% of capsules containing a lepidopteran seed predator (the plume moth Amblyptilia pica [Walsingham], newly discovered as a seed predator on P. furbishiae) had no seeds (mean = 3 seeds). Seed predation affected 39% of the potentially maturable capsules. In 6–30% of all capsules preyed upon, the lepidopteran was parasitized by an ichneumonoid wasp larva; these capsules matured about 7 seeds. Herbivory by rabbits, rodents, and deer also reduced seed set, eliminating 98% of the scapes at one site, and averaging 40% for other populations. Mammals and insects together removed at least 62% of maturable seeds. Populations varied widely in predation and seed set. Predispersal seed loss in P. furbishiae, coupled with slow seedling growth and occasional catastrophic mortality from ice scour and erosion, could limit population growth in this rare endemic.     

A Histological Study of the Effect of Feeding bySmicronyxspp. (Coleoptera: Curculionidae) Larvae on Seed Production byStriga hermonthica(Scrophulariaceae)

Experiments were conducted in Burkina Faso to determine the effect of feeding bySmicronyxspp. (Coleoptera: Curculionidae) larvae on the development of fruit capsule and seed production inStriga hermonthica(Scrophulariaceae). In treated plotsSmicronyxadults were released in the cages while in untreated plots (= control) noSmicronyxwere released. TheStrigacapsules were collected, fixed, and prepared for histological examination. UninfestedStrigacapsules developed normally. In infested capsules, three types of galls were distinguished: they reflected the development of capsule and the seeds. There was a synchrony between seed development and larval development in the same capsule. In most instances, one egg or larva was found per gall.Smicronyxlarvae completely destroyedStrigaseeds in the developing capsule by two mechanisms: (1) by directly eating most of the seeds and (2) by eating the placenta that normally feed the seeds.     

POLLEN MORPHOLOGY AND THE RELATIONSHIPS OF CIRCAEASTER,OF KINGDONIA,AND OF SARGENTODOXA TO THE RANUNCULALES

The pollen of three monotypic genera, Circaeaster, Kingdonia, and Sargentodoxa has been examined by light and scanning electron microscopy and in the case of the last genus, also by transmission electron microscopy. The type of tectum found in Circaeaster and Kingdonia, derivations of a compound layer of striae, has a restricted distribution in the Order Ranunculales. Of 64 genera examined in this order only six had species with a similar tectum. They include Achlys, Epimedium, Jeffersonia, and Vancouveria of the Berberidaceae s.l., the controversial Hydrastis, and Trollius of the Ranunculaceae. Circaeaster and Kingdonia have been considered as related since both have rare and primitive vegetative characteristics, the most notable being open dichotomous leaf venation. They are probably best treated as a ditypic family, Circaeasteraceae. The pollen of Sargentodoxa, especially the structure of the exine, closely resembles that of the Lardizabalaceae. However, the fruits of Sargentodoxa have been considered to be distinct from those of the Lardizabalaceae, suggesting that it be treated as a separate, but closely allied, family.     

Studies on the paradox of seedling rarity in Vaccinium myrtillus L. in NE Scotland

Summary

On the moors studied Vaccinium myrtillus produced many berries containing highly viable seed, yet seedlings were rare. Offtake of berries by birds or mammalian herbivores was apparently minor, most berries simply falling from the bushes when ripe. Fallen berries quickly disappeared, probably being removed by small rodents. Berries were fed experimentally to captive field voles and capercaillies, and it was found that only c. 1% of the seeds survived. For the voles, most of these viable seeds probably resulted from contamination of the droppings with partially-eaten berries or discarded extracted seeds, processes which must contribute to dispersal in the wild. The soil seed-bank was shown to be small, and the viability of buried seed declined from c. 90% to c. 20% in 3 years, due to decay in the soil H horizon and premature germination in the litter layer. Fewer seedlings established from experimental sowings on grass and Vaccinium myrtillus turf than on bare peat. We conclude that Vaccinium myrtillus has a seed-dispersal strategy which secures some long-distance movement by frugivores at the expense of much lost seed, and that the small seed-bank and low numbers of seedlings result from poor defence of the seeds and weak competitive ability of the seedlings; other attributes of Vaccinium myrtillus, e.g. efficient vegetative reproduction, compensate for these losses and enable it to be a successful species.     

INBREEDING DEPRESSION IN MORPHOLOGICAL AND PHYSIOLOGICAL TRAITS OF SCHIEDEA LYDGATEI (CARYOPHYLLACEAE) IN TWO ENVIRONMENTS

We compared inbreeding depression in hermaphroditic Schiedea lydgatei and its gynodioecious sister species, S. salicaria, to infer the level of inbreeding depression in their common ancestor. With measurements of selfing rates, this information can be used to assess the importance of inbreeding depression in the evolution of breeding systems in S. lydgatei and S. salicaria. Morphological and physiological characters related to fitness were compared for inbred and outcrossed S. lydgatei in high- and low-fertilizer environments in the greenhouse. Seed mass, number of seeds per capsule, germination, survival, biomass, number of flowers, and age at first flowering were compared for inbred versus outcrossed progeny. We also measured inbreeding depression in maximal rates of photosynthetic carbon assimilation and stomatal conductance to water vapor, traits that affect fitness through their influence on plant carbon balance and water-use efficiency (ratio of carbon gain to water loss). All traits except number of seeds per capsule in parents and survival showed inbreeding depression, with the magnitude depending on family and environment. High inbreeding depression is likely in the ancestor of S. lydgatei and S. salicaria, indicating that, with sufficiently high selfing rates, females could spread in populations. Hermaphroditism in S. lydgatei is probably favored by low selfing rates. In contrast, the evolution of gynodioecy in S. salicaria apparently has been favored by relatively high selfing rates in combination with high inbreeding depression.     

Rotala pseudojuniperina sp. nov. (Lythraceae) from India

A new species, Rotala pseudojuniperina, is described and illustrated. It grows in a seasonal pond on a high altitude lateritic plateau ecosystem of the northern Western Ghats, India. It resembles the African species R. juniperina in having trimerous flowers, but differs in longer sepal appendages, ovoid capsule and ellipsoid seeds. Amongst the Indian species, R. pseudojuniperina shows affinities towards R. densiflora but differs in trimerous flowers, ovoid capsule and longer and ellipsoid seeds.     

FRUIT TRAITS,AND SEED CONSUMPTION BY RODENTS,OF CRESCENTIA ALATA (BIGNONIACEAE) IN SANTA ROSA NATIONAL PARK,COSTA RICA

The pulp in the large hard fruits of Crescentia alata (Bignoniaceae: jicaro) are eaten by horses, surrogate Pleistocene dispersal agents, and the seeds pass through the horse to germinate in dung. I hypothesize that the distribution of jicaro has probably changed dramatically with the introduction of the horse. Two samples of ripe jicaro fruits weighed an average of 210 and 390 g, and contained an average of about 400 and 600 seeds. There was strong variation within and between two trees in the number of seeds per fruit. Mature live seeds weighed an average of 36 and 40 mg, but varied strongly in weight between and within fruits. The seeds were found to be highly edible to a small terrestrial native rodent (Liomys salvini, Heteromyidae) and this rodent avidly harvests the seeds from horse dung but cannot open the hard fruits by itself. The interaction between horses, jicaro fruits, and Liomys mice is probably representative of that which used to occur between Pleistocene megafauna and a number of contemporary Central American trees.     

Vivipary in Ophiorrhiza mungos L. – a rare phenomenon in angiosperms

Vivipary, the precocious germination of seeds within the parent plant, is a specialised feature of evolutionary and biological importance that ensures survival of a plant. Reports on vivipary in angiosperms are rare, accounting for <0.1% of flowering plants. Here, we report a remarkable case of occurrence of vivipary in Ophiorrhiza mungos. A study was conducted to collect information on the morphology of the capsules that support vivipary, environmental factors that induce vivipary, survival mode and the survival of viviparous seedlings. The hydroscopic movement of the cup‐shaped capsules of O. mungos was found to help in viviparous germination during the rainy season. Of the total seeds in a capsule, 70% showed viviparous germination. The seedlings remaining inside the capsule attain a height of 0.98 ± 0.4 cm and reach the ground when the capsule falls. On the ground, seedlings obtain easy anchorage to the substratum since they have already germinated. Vivipary appears to be an adaptation of O. mungos to the rainy season for ensuring viable offspring. This suggests that vivipary in this species might be artificially induced by continuous spraying with water to rescue seeds in all seasons for use in large‐scale propagation to meet increasing market demand and conservation of this valuable anticancer medicinal herb.     

What are the consequences of ant–seed interactions on the abundance of two dry-fruited shrubs in a Mediterranean scrub?

Strong interactions between dry-fruited shrubs and seed-harvesting ants are expected in early successional scrubs, where both groups have a major presence. We have analysed the implications of the seed characteristics of two dry-fruited shrub species (Coronilla minima and Dorycnium pentaphyllum) on seed predation and dispersal mediated by harvester ants and the consequences of these processes on spatio-temporal patterns of plant abundance in a heterogeneous environment. We found that large C. minima seeds were collected much more (39%) than small D. pentaphyllum seeds (2%). However, not all of the removed seeds of these plant species were consumed, and 12.8% of the seeds were lost along the trails, which increased dispersal distances compared with abiotic dispersal alone. Seed dropping occurred among all microhabitats of the two plant species, but especially in open microhabitats, which are the most suitable ones for plant establishment. The two plant species increased their presence in the study area during the study period: C. minima in open microhabitats and D. pentaphyllum in high vegetation. The large size of C. minima seeds probably limited the primary seed dispersal of this species, but may have allowed strong interaction with ants. Thus, seed dispersal by ants resulted in C. minima seeds reaching more suitable microhabitats by means of increasing dispersal distance and redistribution among microhabitats. In contrast, the smaller size of D. pentaphyllum seeds arguably allows abiotic seed dispersal over longer distances and colonization of all types of microhabitats, although it probably also limits their interaction with ants and, consequently, their redistribution in suitable microhabitats. We suggest that dyszoochory could contribute to the success of plant species with different seed characteristics in scrub habitats where seeds are abundantly collected by seed-harvesting ants.     

小叶兜兰的果实生长和雌配子体发育

为了解濒危兰科植物小叶兜兰(Paphiopedilum barbigerum Tang et Wang)胚珠和雌配子体的发育过程,采用常规石蜡切片技术对其果实的生长动态进行了研究。结果表明,授粉后60~75 d的蒴果内种子数量迅速增加,到授粉后120 d时种子充满整个蒴果。授粉后40 d的胎座上分化形成多数由1层表皮细胞包被1列细胞的胚珠原基;授粉后60 d时位于胎座指状结构末端处紧靠表皮细胞下方的孢原细胞分化为大孢子母细胞。之后,大孢子母细胞经过减数分裂和有丝分裂最终形成成熟胚囊;授粉后135 d胚囊发育成熟,附着在胎座上的种子个体分化明显。小叶兜兰胚囊的发育类型为双孢子葱型,胚珠为倒生胚珠,薄珠心,单珠被,成熟胚囊为8核。这为小叶兜兰的生殖生物学及繁殖体系的建立提供理论依据。     

APHANOSPERMA,A NEW GENUS OF ACANTHACEAE FROM MEXICO WITH UNUSUAL DIASPORES

Recent collections and studies of an unusual species of Acanthaceae from the dry forests of northwestern Mexico reveal it to be generically distinct. Aphanosperma comprises a single species, A. sinaloensis, and is included in the Ruellioideae: Justicieae: Odontoneminae near Carlowrightia. At least four characteristics of the capsule appear to be unique in the family, including progressive reflexing during maturation and permanent retention of the seeds in the capsule valves which thus become the diaspores. The species is shown to be self-compatible and fully autogamous.     

Mixed pollen load and late‐acting self‐incompatibility flexibility in Adenocalymma peregrinum (Miers) L.G. Lohmann (Bignonieae: Bignoniaceae)

• Mixed cross and self‐pollen load on the stigma (mixed pollination) of species with late‐acting self‐incompatibility system (LSI) can lead to self‐fertilized seed production. This “cryptic self‐fertility” may allow selfed seedling development in species otherwise largely self‐sterile. Our aims were to check if mixed pollinations would lead to fruit set in LSI Adenocalymma peregrinum, and test for evidence of early‐acting inbreeding depression in putative selfed seeds from mixed pollinations.
• Experimental pollinations were carried out in a natural population. Fruit and seed set from self‐, cross and mixed pollinations were analysed. Further germination tests were carried out for the seeds obtained from treatments.
• Our results confirm self‐incompatibility, and fruit set from cross‐pollinations was three‐fold that from mixed pollinations. This low fruit set in mixed pollinations is most likely due to a greater number of self‐ than cross‐fertilized ovules, which promotes LSI action and pistil abortion. Likewise, higher percentage of empty seeds in surviving fruits from mixed pollinations compared with cross‐pollinations is probably due to ovule discounting caused by self‐fertilization. Moreover, germinability of seeds with developed embryos was lower in fruits from mixed than from cross‐pollinations, and the non‐viable seeds from mixed pollinations showed one‐third of the mass of those from cross‐pollinations.
• The great number of empty seeds, lower germinability, lower mass of non‐viable seeds, and higher variation in seed mass distribution in mixed pollinations, strongly suggests early‐acing inbreeding depression in putative selfed seeds. In this sense, LSI and inbreeding depression acting together probably constrain self‐fertilized seedling establishment in A. peregrinum.

     

Nectarless flowers with deep corolla tubes in Pedicularis: does long pistil length provide an arena for male competition?

Pollinator‐mediated selection does not seem to have a direct influence on the evolution of a long corolla tube in a nectarless flower. We hypothesized that the long pistil length of the nectarless flower with a deep corolla tube provided an opportunity for male competition. Pedicularis siphonantha, a nectarless and partially self‐incompatible lousewort with substantial variation in corolla tube length, was used to test the hypothesis. We tested whether and how corolla tube length affected seed production per capsule and seed germination rate with different pollination treatments. Flowers were hand‐pollinated with pollen from one self donor and one outcross donor and mixed pollen grains consisting of equal amounts from the two donor types, respectively. Additionally, seeds from open‐pollinated flowers with different corolla tube lengths were collected separately for measurement of germination rate. Pollination treatment and corolla tube length significantly affected number of seeds per capsule. Moreover, a significant positive relationship between seeds per capsule and corolla tube length was found when mixed hand pollination was conducted. Seeds of self hand‐pollinated flowers had a lower germination rate than those from outcross‐pollinated flowers. Under open pollination, seeds from flowers with longer corolla tubes tended to have higher germination rate. In P. siphonantha, outcross pollen may have a higher probability of contributing to the next generation when transferred to flowers with longer corolla tubes. The pistil length, therefore, should be seen as a female choice mechanism, which provides an arena for male‐to‐male competition. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 526–532.