Hygrochastic capsule dehiscence supports safe site strategies in New Zealand alpine Veronica (Plantaginaceae)

Background and Aims

Hygrochasy is a capsule-opening mechanism predominantly associated with plants in arid habitats, where it facilitates spatially and temporally restricted dispersal. Recently, hygrochastic capsules were described in detail for the first time in alpine Veronica in New Zealand. The aim of the present study was to investigate whether hygrochastic capsules are an adaptation of alpine Veronica to achieve directed dispersal to safe sites. We expect that by limiting dispersal to rainfall events, distances travelled by seeds are short and confine them to small habitat patches where both seedlings and adults have a greater chance of survival.

Methods

Dispersal distances of five hygrochastic Veronica were measured under laboratory and field conditions and the seed shadow was analysed. Habitat patch size of hygrochastic Veronica and related non-hygrochastic species were estimated and compared.

Key Results

Dispersal distances achieved by dispersal with raindrops did not exceed 1 m but weather conditions could influence the even distribution of seeds around the parent plant. Compared with related Veronica species, hygrochastic Veronica mostly grow in small, restricted habitat patches surrounded by distinctly different habitats. These habitat patches provide safe sites for seeds due to their microtopography and occurrence of adult cushion plants. Non-hygrochastic Veronica can be predominantly found in large habitats without clearly defined borders and can be spread over long distances along rivers.

Conclusions

The results suggest that hygrochasy is a very effective mechanism of restricting seed dispersal to rainfall events and ensuring short-distance dispersal within a small habitat patch. It appears that it is an adaptation for directed dispersal to safe sites that only exist within the parent habitat.     

Faunal influences on New Zealand seed dispersal characteristics

A range of distinctive dispersal features have been recognised within the New Zealand flora, and a wide range of fauna are involved in the dispersal of seed in New Zealand, either by consuming fruit or seed, or as transporters of adhesive seed. In this study the composition of New Zealand’s zoochorous fauna (except insects) was examined using both trait matching within environmental, morphological and behavioural variables, and compared to the trait pattern of the groups of plant species they disperse. The importance of the different dispersal groups to the plant species they disperse varies with habitat, landform, region of New Zealand, foraging behaviour, and morphology. Over half of New Zealand’s vertebrate fauna are involved in fruit dispersal, though only 6% are considered frugivorous—the remainder include varying quantities of insects and other plant material in their diets. Flighted species are over-represented in wooded environments and higher strata and flightless species predominate in low alpine and grassland habitats. The frugivore-fruiting plant interaction group shows some indications of ecological generalism as frugivorous species consume a range of fruit sizes across all vegetation strata and fruit-bearing plants have lower species diversity and occupy a wide range of habitats. Granivores are over-represented in wetland habitats and the eastern South Island. The importance of species which unintentionally disperse adhesive seed depends on whether they are volant (higher importance in coastal environments) or flightless (higher importance dry grasslands and in low alpine areas). A subgroup of birds, such as the ratite Apteryx spp. and the now extinct Dinornithiform moa, with loose feathers (“velcro” species) are over-represented in lower vegetation strata and this matches the zone where many attachment-dispersed plant species present their seed.     

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.     

Ombrohydrochory: Rain-operated seed dispersal in plants – With special regard to jet-action dispersal in Aizoaceae

The present review describes the ombrohydrochoric dispersal syndrome in plants, i.e. seed expulsion by raindrops. There are two different ombrohydrochoric dispersal modes – dispersal by rain wash and by ballistic forces. Both have been reported from the understory of tropical and temperate forests, from wetlands and from deserts, and from numerous families and genera. A special form of ombrohydrochoric dispersal is the jet-action rain-operated seed dispersal mechanism which is restricted to the semi-desert ice plants, Aizoaceae, one of the major families of the angiosperms. Within this family, 98% of the species possess hygrochastic capsules with an ombrohydrochoric seed dispersal mechanism which in part are also responsible for the remarkable speciation burst and radiation. The highly complex capsules open when wet, and the seeds are expelled by a ‘jet action’ with the kinetic energy of raindrops. The halves of the covering membranes of a locule form a nozzle near the centre of the capsule which serves as a jet. Drops of water falling on the distal opening (after the locule has been filled with water) result in an explosive expulsion of water droplets and seeds through that jet. More seeds are dispersed further away from the capsule than in those capsule types without such a jet mechanism.     

SEED PRODUCTION AND DISPERSAL IN DALECHAMPIA (EUPHORBIACEAE): DIVERGENT PATTERNS AND ECOLOGICAL CONSEQUENCES

Patterns of capsule development and methods of seed dispersal are described and compared for 16 neotropical species of Dalechampia. In 15 species all the capsules in a single inflorescence develop simultaneously. In one species, D. spathulata, the capsules develop sequentially. I suggest that the difference between D. spathulata and the other species is the result of the fact that D. spathulata occurs in a low-light, rainforest environment, whereas the other species grow in high-light environments. Sequential development of capsules appears to reduce the loss of seeds to seed predators in species that occur in light-limited environments. The seeds of all 16 species of Dalechampia are dispersed by explosive dehiscence of the capsules. Within the genus there are several different adaptations that increase the average dispersal distance for the seed crop.     

Temporal and spatial patterns of seed dispersal in two Cistus species (Cistaceae)

Cistus species are obligate seeding, early colonizers that follow disturbance, particularly fire, in Mediterranean ecosystems. We studied seed release, seed dispersal and soil seed populations in stands of Cistus ladanifer and C. libanotis. Seed release started in mid- to late summer (C. ladanifer) or in early autumn (C libanotis), and continued for a very extended period: 8-10 months in C. ladanifer, and for a mean of 16 months in C. libanotis. The xerochastic capsules of both species released seeds by successive dehiscence of the locules. All capsules begin to dehisce simultaneously at the start of the seed release period, but in C. libanotis capsule fragmentation replaced dehiscence early in the seed release period. In plants of both species, seed shadows were characterized by a peak of density beneath the plant canopy and a very short tail of much lower densities, indicating that seeds are concentrated beneath mother plants when dispersed. Nevertheless, in late May, at the onset of the fire season, soil seed densities beneath plant canopies were low compared with densities expected from seed shadows, but were apparently high enough to allow recovery of the stands if a disturbance, such as fire, had taken place. Seed-eating Bruchidae in summer and granivorous ants during the seed release period were apparently the main causes of seed losses. Results suggest that in both Cistus species, the staggered seed release could constitute an efficient risk-reducing trait. The plant pool of seeds existing throughout most of the year could be a relevant component of Cistus seed banks.     

Correlates of geographic range size in New Zealand Chionochloa (Poaceae) species

Aim To test for correlations between plant traits and geographic range size. Location: New Zealand. Methods Trait data were derived from comparative experiments, in which plants were grown in pots or in a common garden, that tested for intrinsic differences among the species in traits relating to growth, reproduction and dispersal. Controlled experiments were used to test for differences in responses to drought and waterlogging stress. Geographic range size was measured as the number of 10 km grid squares in the New Zealand region containing at least one occurrence of the species. Results Growth rate, dispersal capacity and environmental tolerance were all positively related to geographic range size. Geographically restricted species tended to have more variable flowering between years. Flowering intensity, reproductive allocation, seed set, diaspore size, and responses to single environmental factors were not related to geographic range size. Main conclusions The differences between range‐restricted and widespread Chionochloa species appear to represent alternative strategies of coping with environmental change in a dynamic landscape. Range‐restricted species are specialized to temporally persistent habitats that are of limited geographic extent. As a consequence, they have evolved traits that conflict with persistence in widespread habitats. The implication for conservation management is that the conservation of rare plants will frequently depend on protection of their habitats. The widespread Chionochloa species possess traits that enable them to disperse to and occupy a greater range of habitats. These traits have allowed some of these species to expand their ranges following environmental changes that favoured an increase in grassland extent.     

The biogeography of the austral, subalpine genus Ourisia (Plantaginaceae) based on molecular phylogenetic evidence: South American origin and dispersal to New Zealand and Tasmania

Molecular phylogenetic analyses of 26 of the 28 species of Ourisia , including eight of ten subspecies and two purported natural hybrids, are presented and used to examine the biogeography of the genus, which is distributed in subalpine to alpine habitats of South America, New Zealand and Tasmania. Gondwanan vicariance, often cited as the cause of this classic austral biogeographical pattern, was rejected by parametric bootstrapping of our combined dataset. Alternatively, various lines of evidence are presented in favour of a South American origin of Ourisia and subsequent dispersal to Australasia. Specifically, the genus likely arose in the Andes of central Chile and spread to southern Chile and Argentina, to the north-central Andes, and finally to Tasmania and New Zealand. The ancestor of the New Zealand species probably first arrived on the South Island, where the New Zealand species of Ourisia are most diverse, and migrated to the North and Stewart Islands. Because the Tasmanian and New Zealand species are sister to one another, the direction of dispersal between these two areas is equivocal. These results agree with other molecular phylogenetic studies that show that past dispersal between southern hemisphere continents has played an important role in the evolutionary history of many high-elevation austral plants. Our data also show that within South America, many of the geographical barriers (with the exception of the Atacama Desert) that have played a role in the evolution of other plant groups have not affected Ourisia species. Within New Zealand, the phylogeny and biogeography of species of Ourisia coincide with the geological history of the country and patterns of other alpine plants. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 479–513.     

Wind dispersal of alpine plant species: A comparison with lowland species

Question: The prominent role of wind dispersal in alpine habitats has been recognized early but has rarely been quantified. The aim of this study is to compare wind dispersal under alpine and lowland conditions and to analyse whether differences are caused by species traits, e.g. terminal velocity of seeds (V_term) or weather conditions. Location and Methods: We characterized wind dispersal potential of > 1100 Central European species using measured V_term To quantify the habitat effect on wind dispersal, we measured meteorological key‐parameters and simulated dispersal distance spectra of nine selected species under typical alpine conditions (foreland of the Scaletta‐glacier, Switzerland) and typical lowland conditions (grassland in Bad Lippspringe, Germany). Results: Lowland species had higher V_term compared to alpine species. However, this difference is absent when only species of species of open habitats are concerned. The meteorological measurements showed that the alpine habitat was mainly characterized by higher frequency and strength of updrafts. The simulations showed that under alpine conditions long distance dispersal occurred much more frequent. Conclusions: More than 50 % of the alpine species have a fair chance to be dispersed by wind over long distances, while this proportion is less than 25 % for species from open habitats in the lowland. The more prominent role of wind dispersal in alpine habitats is mainly a result of differences in environmental conditions, namely more intense vertical turbulence in the alpine habitat, and does not result from prominent differences in plant traits, namely V_term, between alpine and lowland species.     

A new rain-operated seed dispersal mechanism in Bertolonia mosenii (Melastomataceae), a Neotropical rainforest herb

Although widespread among fungi, lichens, liverworts, and mosses, seed dispersal mechanisms operated by rain are unusual among flowering plants. Generally speaking, two mechanisms are involved in seed dispersal by rains: the splash-cup and the springboard. Here we describe a new seed dispersal mechanism operated by rain in a Neotropical rainforest herb Bertolonia mosenii Cogniaux (Melastomataceae). The study was carried out at the lowland Atlantic rainforest, southeastern Brazil. We experimentally demonstrate that rain is necessary to release the seeds from the capsules through what we call "squirt-corner" seed dispersal mechanism: when a raindrop strikes the mature fruit, the water droplet forces the seeds outward to the angles (corners) of the triangular capsule and the seeds are released. As far as we know squirt-corner represents a new rain-operated seed dispersal mechanism, and a novel seed dispersal mode both for Melastomataceae and for flowering plants from Neotropical forests.     

Between-year variation in seed weights across altitudes in the high-alpine plant <Emphasis Type="Italic">Eritrichium nanum</Emphasis>

Seed weight is a prominent life history trait of plants affecting dispersal, establishment, and survival. In alpine environments, the few studies investigating the effect of elevation on seed weight within species have mainly detected a decrease in seed weight with increasing elevation. This relationship is generally attributed to the adverse climate at high elevations. In order to test this hypothesis, we analyzed seed weight variation across altitudes (2,435–3,055 m a.s.l.) in two consecutive years that differed in weather conditions in the high-alpine cushion plant Eritrichium nanum. We found a significant reduction in seed weight with increasing elevation in both years, but in the growing season with more adverse weather conditions, the reduction was more substantial than in the more favorable year. We conclude that alpine plants may be able to produce well-developed seeds at low elevations in almost all years, independent of weather conditions, whereas reproduction through seeds is potentially limited to years of favorable weather at high elevation.     

Seed dispersal systems in the New Zealand flora

Knowledge of the dispersal mechanisms used by plants is important in phylogenetic, ecological, biogeographical, and conservation studies. Here we attempt to assign dispersal mechanisms to the entire flora-2595 plant species of the New Zealand Botanical Region. Anemochory is the most frequent dispersal mechanism, utilised by 79% of species. The next most frequent mechanisms are endozoochory (33%), hydrochory (28%), epizoochory (26%), and ballistic (8%). Polychory is common, particularly in monocotyledonous and dicotyledonous families and hydrochorous, epizoochorous, and ballistically dispersed species. Epizoochory is more common in New Zealand than in other regions, and species using this dispersal mechanism are over-represented among threatened species. Frugivory is less common than previously reported, and is under-represented among threatened species. Some mechanisms are poorly known, and entanglement and capsulivory are dispersal mechanisms apparently unique to New Zealand. Dispersal mechanisms reflective of New Zealand's distinctive assemblages of large flightless birds and reptiles are not apparent. A pattern of reduction in dispersal-related structures is evident in some genera. The mechanisms utilised by some species are ambiguous. Thus there remains a need for further investigation of the dispersal mechanisms utilised by plants in New Zealand.     

Seedling emergence responds to both seed source and recruitment site climates: a climate change experiment combining transplant and gradient approaches

Seedling recruitment allows genetic recombination and production of dispersal units. Both the climate experienced by the source populations (seed source effect) and the weather experienced by the seeds during germination and seedling emergence (recruitment site effects) are important for seedling recruitment. Separating these effects in the field is essential to assess potential climate change impacts on plant population. We combine experimental seed transplant and gradient analyses to separate the effects of seed source and recruitment site temperature and precipitation for the seedling emergence of two alpine/lowland species pairs (Viola biflora/Viola palustris, Veronica alpina/Veronica officinalis). Combining these approaches allows us to compare local responses versus responses along environmental gradients, but also tests for local adaptation and/or pre-conditioning effects (adaptive seedling emergence responses). Veronica officinalis emergence increased with increasing seed source temperature in both the experimental and the gradient approaches, and showed adaptive seedling emergence. Viola biflora, Viola palustris and Veronica alpina emergence decreased with recruitment site temperature in both approaches. Both Violas emergences increased with recruitment site precipitation, in both approaches for the alpine violet, and in the gradient approach for lowland one. Emergence was primarily affected by the environment of the recruitment site, whereas seed source climate and adaptive seedling emergence impacted recruitment in only one of our species. The responses to recruitment site temperatures were negative, whereas the response to seed source temperature was positive. Ignoring the distinctions between these different mechanisms can lead to erroneous conclusions regarding potential climate change impacts on plant recruitment.     

Seed dispersal in Erythronium dens-canis L. (Liliaceae): variation among habitats in a myrmecochorous plant

Erythronium dens-canis is a geophyte which produces a single flower each season. The fruits produce small seeds with relatively large elaiosomes. We performed experiments to investigate primary and secondary seed dispersal mechanisms of this species in different habitats in the western part of the Cantabrian Range in northwest Spain. Sticky traps were used to measure primary dispersal of seeds up to 0.5 m from mother plants. Seed cafeteria experiments were performed in different habitats to examine the role of ants and rodents in secondary seed transport and seed predation. Our results indicate that: (a) primary seed dispersal is positively skewed (99% of seeds fall within 20 cm of the mother plant) and seed dispersal distances vary significantly among plants; (b) secondary dispersal is exclusively by myrmecochory, although the proportion of seeds removed by ants differs significantly among habitats; (c) ant species composition and abundances vary among habitats; and (d) freshly dropped seeds are more likely to be removed than seeds that have begun to dry out. We conclude that secondary dispersal of seeds is greatly influenced by habitat but not by small-scale microhabitat. This revised version was published online in June 2006 with corrections to the Cover Date.     

Ant-mediated expansion of an obligate seeder species during the first years after fire

Most obligate seeder species build up a soil seed bank that is associated with massive seed germination in the year immediately after a fire. These species are also shade‐intolerant and disappear when vegetation cover closes, creating unsuitable conditions for seedling recruitment. The only way for these plants to expand their populations is when habitats suitable for seedling recruitment arise (i.e. in years immediately after a fire). However, short primary seed dispersal of obligate seeders does not allow these plants to colonise the suitable habitats, and these habitats can only be colonised by secondary seed dispersion. We hypothesised that Fumana ericoides, an obligate‐seeding small shrub, not only establishes abundantly in the first year after fire, but also expands its local range in the following years due to secondary dispersal by ants while suitable habitats are still available. We tested this hypothesis using experimental studies and a simulation model of potential population expansion in a recently burned area. Results showed that F. ericoides not only established prolifically in the year immediately after fire, but was also able to recruit new individuals and expand its population in the years following the fire, despite a low germination rate and short primary seed dispersal. Ant‐mediated seed dispersal and availability of suitable habitats were key factors in this phenomenon: ants redistributed seeds in suitable habitats while they were available, which accelerated the expansion of F. ericoides because new plants established far away from the core population.     

Nurse effect of the native cushion plant Azorella monantha on the invasive non-native Taraxacum officinale in the high-Andes of central Chile

Positive interactions among native plant species are common in alpine habitats, particularly those where one species (nurse plant) generates microclimatic conditions that are more benign than the surrounding environment, facilitating the establishment of other species. Nonetheless, these microclimatic conditions could facilitate the establishment of non-native species as well. A conspicuous component of the alien alpine flora of the central Chilean Andes is the perennial herb Taraxacum officinale agg. (dandelion). In contrast to other alien species that are restricted to human-disturbed sites, T. officinale is frequently observed growing within native plant communities dominated by cushion plants. In this study we evaluated if T. officinale is positively associated with the cushion plant Azorella monantha. Via seedling survival experiments and gas-exchange measurements we also assessed the patterns of facilitation between cushions and dandelions, and explore the potential mechanisms of invasion by dandelions. T. officinale grows spatially positively associated with cushions of A. monantha. Survival of seedlings, as well as their net-photosynthetic rates and stomatal conductance, were higher within cushions than in open areas away from them, suggesting that the microclimatic modifications generated by this native cushion facilitates the establishment and performance of a non-native invasive species. Our results, as well as other recent studies, highlight the role of native communities in facilitating rather than constraining non-native plant invasions, particularly in stressful habitats such as alpine environments.     

蚁对植物种子的传播作用

许多种子植物依靠动物传播种子 ,称为动物传播。根据动物类群的不同 ,可分为哺乳类传播 ,鸟传播 ,鱼传播 ,蚁传播等。鸟传播和蚁传播的研究近年取得了很大的进展 ,但国内在这方面研究较缺乏 ,作者已就鸟传播作了综述报道 ,现将蚁传播的研究综述报道如下。1　蚁与植物的相互关系蚁类属膜翅目 (Ｈｙｍｅｎｏｐｔｅｒａ) ,蚁科 (Ｆｏｒｍｉｃｉ ｄａｅ) ,典型的社会性昆虫。多数蚁类是肉食性的 ,以小动物或更小的蚁类为食 ,但也有很多蚁类是植物食性的。在大多数生态系统中均有蚁类分布 ,而且蚁类数量众多 ,在森林生态系统中每 1ｈａ可达 6～10…     

Seed dispersal and colonization ability of plants — Assessment and implications for conservation

Seed dispersal is one of the least understood phases in plant life-cycles, despite being recognized as fundamentally important for understanding species distribution, abundance, population dynamics and life-history variation. For species living in fragmented habitats, dispersal may be crucial for regional persistence. This paper synthesizes studies performed in Swedish semi-natural grasslands and deciduous forests. The main objective was to develop a simple method to assess the dispersal and colonization ability of plants. Dispersal and colonization were considered as a series of steps: seed production, seed transport and seedling recruitment. With the use of verified assumptions on (i) a trade-off between seed production and seed size, (ii) a relatively small predictive power of seed transport for species actual dispersal and colonization, (iii) a positive effect of seed size on recruitment, and (iv) a general prevalence of recruitment limitations by seed availability, a prediction is made that dispersal and colonization ability will be highest among species with intermediate seed sizes in a given community. This prediction is supported by field data. Based on estimated dispersal and colonization ability, and the ability of local populations to persist, a scheme is suggested with four basic categories of plants. Some implications are suggested for conservation, in a landscape scale, of plants belonging to each of these four categories.     

新疆东部天山蝶类多样性及其垂直分布

2006-2008年研究了新疆东部天山蝶类多样性和垂直分布.结果表明:研究区域内共记录蝴蝶7科43属63种,占新疆已记录蝶类种数的24.80％,区系组成主要是古北种,占73％;其次是广布种,占27％,没有发现东洋种.其中蛱蝶科的物种数最多,为11属19种,蚬蝶科的物种数最少,只有1属1种.按海拔将生境分为5个垂直自然带,包括低山灌木草原带、山地森林草原带、亚高山草甸带、高山草甸带、垫状植被带.蝶类物种数和个体数排序为亚高山草甸带＞山地森林草原带＞低山灌木草原带＞高山草甸带＞垫状植被带.采用Shannon-Wiener指数和G-F指数对蝶类物种和科、属的多样性进行了分析评价,结果显示亚高山草甸带的蝶类多样性最为丰富,其次是山地森林草原带和低山灌木草原带,而高山草甸带和垫状植被带的蝶类多样性相对较低,物种和科、属多样性分析结果均一致.蝶类垂直分布明显,物种数和个体数随海拔变化的趋势类似,均为先增加后下降.蝶类区系成分随着海拔升高发生改变,广布种的比例逐渐降低,高山草甸带和垫状植被带只有古北种分布.研究结果显示,生境改变对蝴蝶群落影响明显,保护生境是保护蝴蝶生存的最主要措施.     

SEED GERMINATION TIMING PATTERNS IN INTERMOUNTAIN PENSTEMON (SCROPHULARIACEAE)

Regulation of seed germination timing is an important component of population persistence, especially for short-lived plants in seasonal environments with unpredictable year-to-year weather variation. Seed germination patterns were examined in the laboratory for seeds from 135 populations belonging to 38 species and 13 sections of the genus Penstemon (Scrophulariaceae), from habitats ranging from warm desert to alpine tundra. Seeds of most species were dormant at dispersal and required a period of moist chilling to become germinable. Response to chilling was related to probable chilling duration at the collection site. Populations from habitats with severe winters produced seeds with long chilling requirements, while those from habitats with mild winters produced seeds with short chilling requirements. Populations from midelevation habitats produced seeds with intermediate chilling requirements but with a sizeable fraction whose dormancy was not broken by chilling of any length. Another pattern observed mostly in warm desert populations was little primary dormancy at autumn temperatures combined with induction of a fraction into secondary dormancy by short chilling. Species with a wide habitat range included populations with contrasting germination patterns. Parallel habitat-correlated patterns were observed in different sections. Most species showed germination patterns that combined predictive mechanisms with potential for carryover of a persistent seed bank. Results of common garden experiments suggested that germination differences had a strong genetic basis both among populations and among plants in a population. Adaptive radiation in the genus Penstemon has thus included the evolution of habitat-specific germination timing strategies in multiple lineages within the genus.