The proposal is made that seed scientists need an internationally acceptable hierarchical system of classification for seed dormancy. Further, we suggest that a modified version of the scheme of the Russian seed physiologist Marianna G. Nikolaeva be adopted. The modified system includes three hierarchical layersclass, level and type; thus, a class may contain levels and types, and a level may contain only types. The system includes five classes of dormancy: physiological dormancy (PD), morphological dormancy (MD), morphophysiological dormancy (MPD), physical dormancy (PY) and combinational dormancy (PY + PD). The most extensive classification schemes are for PD, which contains three levels and five types (in the nondeep level), and MPD, which contains eight levels but no types. PY is not subdivided at all but probably should be, for reasons given. Justifications are presented for not including mechanical dormancy or chemical dormancy in the modified scheme. PD (non-deep level) is the most common kind of dormancy, and occurs in gymnosperms (Coniferales, Gnetales) and in all major clades of angiosperms. Since, first, this is the class and level of dormancy in seeds of wild populations of Arabidopsis thaliana and, secondly, Type 1 (to which seeds of A. thaliana belong) is also common, and geographically and phylogenetically widespread, it seems that biochemical, molecular and genetic studies on seed dormancy in this model species might have rather broad application in explaining the basic mechanism(s) of physiological dormancy in seeds.

Physical dormancy (PY) is caused by a water-impermeable seed or fruit coat. It is known, or highly suspected, to occur in nine orders and 15 families of angiosperms (sensu Angiosperm Phylogeny Group 1998), 13 of which are core eudicots. The Zingiberales is the only monocot order, and Cannaceae (Canna) the only monocot family, in which PY is known to occur. Six of the nine orders, and 12 of the 15 families, in which PY occurs are rosids. Furthermore, six of the families belong to the Malvales. The water-impermeable palisade layer(s) of cells are located in the seed coats of 13 of the families, and in the fruit coats of Anacardiaceae and Nelumbonaceae. In all 15 families, a specialized structure is associated with the water-impermeable layer(s). The breaking of PY involves disruption or dislodgment of these structures, which act as environmental 'signal detectors' for germination. Representatives of the nine angiosperm orders in which PY occurs had evolved by the late Cretaceous or early Tertiary (Paleogene). Anatomical evidence for PY in fruits of the extinct species Rhus rooseae (Anacardiaceae, middle Eocene) suggests that PY had evolved by 43 Ma, and probably much earlier. We have constructed a conceptual model for the evolution of PY, and of PY+ physiological dormancy (PD), within Anacardiaceae. The model begins in pre-Eocene times with an ancestral species that has large, pachychalazal, non-dormant (ND), recalcitrant seeds. By the middle Eocene, a derived species with relatively small, partial pachychalazal, orthodox seeds and a water-impermeable endocarp (thus PY) had evolved, and by the Oligocene, PD had been added to the seed (true seed + endocarp) dormancy mechanism. It is suggested that climatic drying (Eocene), followed by climatic cooling (Eocene-Oligocene transition), were the primary selective agents in the development of PY. An evolutionary connection between PY and recalcitrance is suggested by the relatively high concentration of these two character states in the rosids. Phylogenetic data and fossil evidence seem to support the PY AE (PY + PD) evolutionary sequence in Anacardiaceae, which also may have occured in Leguminosae.

Summary 1The ability of seeds to survive desiccation is an important functional trait and is an integral part of plant regeneration ecology. Despite this, the topic has received relatively little attention from ecologists. In this study, we examine the relationships between seed desiccation tolerance and two important aspects of plant regeneration ecology: habitat and dormancy. This is done by comparative analysis of a data set of 886 tree and shrub species from 93 families. 2 The proportion of species displaying desiccation sensitive seeds declines as the habitat becomes drier, and possibly also cooler, although the latter observation requires cautious interpretation. Desiccation sensitivity is most common in moist, relatively aseasonal vegetation zones, but is infrequent in, though not absent from arid and highly seasonal habitats. 3 The highest frequency of desiccation sensitivity occurs in non-pioneer evergreen rain forest trees, although 48% of the species examined have desiccation tolerant seeds. In contrast, all pioneer taxa within the data set have drying tolerant seeds. 4 Desiccation sensitivity is more frequent in seeds that are non-dormant on shedding ( c. 31%), than dormant ( c. 9%). Highest frequencies of drying tolerance occur in seeds with physical or combinational dormancy, at 99% and 100%, respectively. 5 Although there is an association between non-dormancy and desiccation sensitivity in both tropical and temperate zones, the relationship does not appear to be causal. 6 Working from the hypothesis that seed desiccation sensitivity represents a derived state in extant species, we use the results to investigate and discuss possible ecological trade-offs and associated fitness advantages. These may explain the hypothesized repeated loss of this trait. The frequent association between large seed size and desiccation sensitivity is also considered.

SummarySeed dormancy, by controlling the timing of germination, can strongly affect plant survival. The kind of seed dormancy, therefore, can influence both population and species-level processes such as colonization, adaptation, speciation, and extinction.We used a dataset comprising over 14 000 taxa in 318 families across the seed plants to test hypotheses on the evolution of different kinds of seed dormancy and their association with lineage diversification.We found morphophysiological dormancy to be the most likely ancestral state of seed plants, suggesting that physiologically regulated dormancy in response to environmental cues was present at the origin of seed plants. Additionally, we found that physiological dormancy (PD), once disassociated from morphological dormancy, acted as an 'evolutionary hub' from which other dormancy classes evolved, and that it was associated with higher rates of lineage diversification via higher speciation rates.The environmental sensitivity provided by dormancy in general, and by PD in particular, appears to be a key trait in the diversification of seed plants.