The ontogeny of disparity in Cupressaceae seed cones.

Ontogenetic shape change has long been recognized to be important in generating patterns of morphological diversity and may be especially important in plant reproductive structures. We explore how seed cone disparity in Cupressaceae changes over ontogeny by comparing pollination-stage and mature cones. We sampled cones at pollen and seed release and measured cone scales using basic morphometric shape variables. We used multivariate statistical methods, particularly hypervolume overlap calculations, to measure morphospace occupation and disparity. Cone scales at both pollination and maturity exhibit substantial variability, although the disparity is greater at maturity. Mature cone scales are also more clustered in trait space, showing less overlap with other taxa than at pollination. These patterns reflect two growth strategies that generate closed cones over maturation, either through thin laminar scales or relatively thick, peltate scales, resulting in two distinct regions of morphospace occupation. Disparity patterns in Cupressaceae seed cones change over ontogeny, reflecting shifting functional demands that require specific patterns of cone scale growth. The evolution of Cupressaceae reproductive disparity therefore represents selection for trajectories of ontogenetic shape change, a phenomenon that should be widespread across seed plants.

Heavy metal accumulation characteristics and physiological response of Sabina chinensis and Platycladus orientalis to atmospheric pollution.

Heavy metal (Cu, Mn, Zn, Pb, and Cd) concentrations were measured in the leaves of Sabina chinensis and Platycladus orientalis collected from urban, suburban, and rural sites in Tianjin, China. Photosynthetic pigment contents, reactive oxygen species content, malondialdehyde (MDA) content and antioxidant enzyme activity were investigated, providing physiological response parameters. Our comparison of the sites revealed that urbanization significantly influenced the heavy metal concentrations in both plant leaves. At the rural site, both plant leaves exhibited the lowest heavy metal accumulation. The highest Cu, Mn, and Zn concentrations were found in S. chinensis leaves from the urban site; the highest Pb and Cd concentrations were found in P. orientalis leaves from the urban site. These results indicate that the urban site contained larger heavy metal concentrations in the plant leaves that may reflect the anthropogenic emission gradient. It is also found that S. chinensis may be used to monitor airborne heavy metal pollution because it is highly quick response to heavy metals, while P. orientalis may be used for mitigation due to its high resistance. The results of this study can contribute to the development of monitoring and environmental management plans by providing information on sensitive and resistant tree species for city greening in North China.

Herb and conifer roots show similar high sensitivity to water deficit.

Root systems play a major role in supplying the canopy with water, enabling photosynthesis and growth. Yet, much of the dynamic response of root hydraulics and its influence on gas exchange during soil drying and recovery remains uncertain. We examined the decline and recovery of the whole root hydraulic conductance (Kr) and canopy diffusive conductance (gc) during exposure to moderate water stress in two species with contrasting root systems: Tanacetum cinerariifolium (herbaceous Asteraceae) and Callitris rhomboidea (woody conifer). Optical dendrometers were used to record stem water potential at high temporal resolution and enabled non-invasive measurements of Kr calculated from the rapid relaxation kinetics of water potential in hydrating roots. We observed parallel declines in Kr and gc to <20% of unstressed levels during the early stages of water stress in both species. The recovery of Kr after rewatering differed between species. T. cinerariifolium recovered quickly, with 60% of Kr recovered within 2 h, while C. rhomboidea was much slower to return to its original Kr. Recovery of gc followed a similar trend to Kr in both species, with C. rhomboidea slower to recover. Our findings suggest that the pronounced sensitivity of Kr to drought is a common feature among different plant species, but recovery may vary depending on root type and water stress severity. Kr dynamics are proposed to modulate gc response during and following drought.

Effects of vegetation, terrain and soil layer depth on eight soil chemical properties and soil fertility based on hybrid methods at urban forest scale in a typical loess hilly region of China.

Although the spatial mapping and fertility assessment of soil chemical properties (SCPs) are well studied in the Loess Plateau region of China at farmland scale, little is known about spatial mapping the SCPs and their fertility and their influence factors at urban forest scale. The objectives of this study were to (1) compare the performance of two spatial interpolation methods, Ordinary kriging (OK) and regression kriging (RK), and (2) explain the relationships of the vegetation, terrain, and soil layer depth between the eight SCPs and their fertility, and (3) find the limiting factors of soil comprehensive fertility in this study area? The Yan'an urban forest was taken as study case, used hybrid spatial interpolation methods based on OK and RK to mapping eight SCPs and the soil fertility in each soil layer (0-20 cm, 20-40 cm, and 40-60 cm) for 285 soil samples. The results indicated that RK outperformed OK for total nitrogen (TN), available potassium (AK), organic matter (OM) in 0-60 cm profile and available phosphorus (AP) in the 0-20 cm and 40-60 cm soil layers because RK considered the impact of terrain. The terrain factors, comprising the relative terrain position, slope, aspect, and relative elevation significantly affected the SCPs and spatial heterogeneity of fertility, where the vegetation cover types determined the average SCPs to some extent. On average, the six SCPs (except total potassium and AP) and the fertility decreased as the soil layer depth increased. Ten vegetation cover types comprising broadleaved mixed natural forest (BM), cultivated land (CL), economic forest (EF), grassland (GL), Platycladus orientalis natural forest (PON), Platycladus orientalis plantation (POP), Pinus tabuliformis plantation (PT), Quercus wutaishanica natural forest (QW), Robinia pseudoacacia plantation (RP), and Shrubwood (SW) were associated with significant differences in TN, OM, AN, AP, and AK, across the three soil layers. QW, PON, and BM also had higher content of TN, OM, AN, and AK contents than the other vegetation cover types. There were small differences in TK, AK, and pH among the 10 vegetation cover types. We concluded that AN, TN, and OM are the limiting factors of soil comprehensive fertility in this region. These results improve understanding of the spatial mapping, influence and limiting factors of SCPs and their fertility at urban forest scales.

Responses of Intrinsic Water-use Efficiency and Tree Growth to Climate Change in Semi-Arid Areas of North China.

Tree-level intrinsic water-use efficiency (iWUE) is derived from the tree-ring 13C isotope composition (δ13C) and is an important indicator of the adaptability for trees to climate change. However, there is still uncertainty regarding the relationship between long-term forest ecosystem carbon sequestration capacity and iWUE. To determine whether elevated atmospheric CO2 concentration (Ca) increase iWUE and tree growth (basal area increment, BAI), dendrochronological methods and stable isotope analyses were used to examine annual changes in the tree-ring width and carbon isotope composition (δ13C) of Platycladus orientalis in northern China. The iWUE derived from δ13C has increased significantly (p < 0.01). Long-term iWUE trend was largely and positively driven by the elevated atmospheric CO2 concentration and temperature. We observed a general increase in averaged BAI, which had significant positive correlation with iWUE (R2 = 0.3186, p < 0.01). Increases in iWUE indeed translated into enhanced P. orientalis growth in semi-arid areas of northern China. Elevated atmospheric CO2 concentration significantly (p < 0.01) stimulated P. orientalis biomass accumulation when Ca was less than approximately 320 ppm in the early phase; however, this effect was not pronounced when Ca exceeded 320 ppm.

Aridity drove the evolution of extreme embolism resistance and the radiation of conifer genus Callitris.

Xylem vulnerability to embolism is emerging as a major factor in drought-induced tree mortality events across the globe. However, we lack understanding of how and to what extent climate has shaped vascular properties or functions. We investigated the evolution of xylem hydraulic function and diversification patterns in Australia's most successful gymnosperm clade, Callitris, the world's most drought-resistant conifers. For all 23 species in this group, we measured embolism resistance (P50 ), xylem specific hydraulic conductivity (Ks ), wood density, and tracheary element size from natural populations. We investigated whether hydraulic traits variation linked with climate and the diversification of this clade using a time-calibrated phylogeny. Embolism resistance varied widely across the Callitris clade (P50 : -3.8 to -18.8 MPa), and was significantly related to water scarcity, as was tracheid diameter. We found no evidence of a safety-efficiency tradeoff; Ks and wood density were not related to rainfall. Callitris diversification coincides with the onset of aridity in Australia since the early Oligocene. Our results highlight the evolutionary lability of xylem traits with climate, and the leading role of aridity in the diversification of conifers. The uncoupling of safety from other xylem functions allowed Callitris to evolve extreme embolism resistance and diversify into xeric environments.

Environmental controls of evapotranspiration in a mixed plantation in North China.

The mixed plantation plays an important role in the water cycle in the hilly area of North China. To evaluate the effect of afforestation on the water balance in this region, the temporal variation of evapotranspiration (ET) and environmental controls were investigated based on the eddy flux measurement of water vapor in a 31-year-old mixed plantation from 2006 to 2010. During 5 years, annual ET ranged from 513 to 680 mm, with an average of 579 mm. Growing season ET accounted for 72-82 % of annual ET during the 5-year period and its interannual variation was determined by the number of rainy days. In the non-growing and growing seasons, monthly ET was primarily dependent on monthly mean soil water content and monthly mean net radiation, respectively. Annual mean Priestley-Taylor coefficient (α) was 0.64, and the decoupling factor (Ω) was 0.48. High values of α and Ω implied that ET was energy limited in the growing seasons of 2006-2010. The mean annual ratio of ET to precipitation (ET/P) was 1.10. The density of the mixed plantation was around 50 % higher than the optimal value determined by local water capacity, leading to a large ET/P ratio. The dense plantation needs to be thinned to prevent excessive water loss in the hilly area of North China.

Experimental evidence for negative turgor pressure in small leaf cells of Robinia pseudoacacia L versus large cells of Metasequoia glyptostroboides Hu et W.C.Cheng. 1. Evidence from pressure-volume curve analysis of dead tissue.

This paper provides a mini-review of evidence for negative turgor pressure in leaf cells starting with experimental evidence in the late 1950s and ending with biomechanical models published in 2014. In the present study, biomechanical models were used to predict how negative turgor pressure might be manifested in dead tissue, and experiments were conducted to test the predictions. The main findings were as follows: (i) Tissues killed by heating to 60 or 80 °C or by freezing in liquid nitrogen all became equally leaky to cell sap solutes and all seemed to pass freely through the cell walls. (ii) Once cell sap solutes could freely pass the cell walls, the shape of pressure-volume curves was dramatically altered between living and dead cells. (iii) Pressure-volume curves of dead tissue seem to measure negative turgor defined as negative when inside minus outside pressure is negative. (iv) Robinia pseudoacacia leaves with small palisade cells had more negative turgor than Metasequoia glyptostroboides with large cells. (v) The absolute difference in negative turgor between R. pseudoacacia and M. glyptostroboides approached as much as 1.0 MPa in some cases. The differences in the manifestation of negative turgor in living versus dead tissue are discussed.

Experimental evidence for negative turgor pressure in small leaf cells of Robinia pseudoacacia L versus large cells of Metasequoia glyptostroboides Hu et W.C. Cheng. 2. Höfler diagrams below the volume of zero turgor and the theoretical implication for pressure-volume curves of living cells.

The physiological advantages of negative turgor pressure, Pt , in leaf cells are water saving and homeostasis of reactants. This paper advances methods for detecting the occurrence of negative Pt in leaves. Biomechanical models of pressure-volume (PV) curves predict that negative Pt does not change the linearity of PV curve plots of inverse balance pressure, PB , versus relative water loss, but it does predict changes in either the y-intercept or the x-intercept of the plots depending on where cell collapse occurs in the PB domain because of negative Pt . PV curve analysis of Robinia leaves revealed a shift in the x-intercept (x-axis is relative water loss) of PV curves, caused by negative Pt of palisade cells. The low x-intercept of the PV curve was explained by the non-collapse of palisade cells in Robinia in the PB domain. Non-collapse means that Pt smoothly falls from positive to negative values with decreasing cell volume without a dramatic change in slope. The magnitude of negative turgor in non-collapsing living cells was as low as -1.3 MPa and the relative volume of the non-collapsing cell equaled 58% of the total leaf cell volume. This study adds to the growing evidence for negative Pt .

The impact of xylem cavitation on water potential isotherms measured by the pressure chamber technique in Metasequoia glyptostroboides Hu & W.C. Cheng.

Pressure-volume (PV) curve analysis is the most common and accurate way of estimating all components of the water relationships in leaves (water potential isotherms) as summarized in the Höfler diagram. PV curve analysis yields values of osmotic pressure, turgor pressure, and elastic modulus of cell walls as a function of relative water content. It allows the computation of symplasmic/apoplastic water content partitioning. For about 20 years, cavitation in xylem has been postulated as a possible source of error when estimating the above parameters, but, to the best of the authors' knowledge, no one has ever previously quantified its influence. Results in this paper provide independent estimates of osmotic pressure by PV curve analysis and by thermocouple psychrometer measurement. An anatomical evaluation was also used for the first time to compare apoplastic water fraction estimates from PV analysis with anatomical values. Conclusions include: (i) PV curve values of osmotic pressure are underestimated prior to correcting osmotic pressure for water loss by cavitation in Metasequoia glyptostroboides; (ii) psychrometer estimates of osmotic pressure obtained in tissues killed by freezing or heating agreed with PV values before correction for apoplastic water dilution; (iii) after correction for dilution effects, a solute concentration enhancement (0.27MPa or 0.11 osmolal) was revealed. The possible sources of solute enhancement were starch hydrolysis and release of ions from the Donnan free space of needle cell walls.

Survival, growth and vulnerability to drought in fire refuges: implications for the persistence of a fire-sensitive conifer in northern Patagonia.

Fire severity and extent are expected to increase in many regions worldwide due to climate change. Therefore, it is crucial to assess the relative importance of deterministic vs. stochastic factors producing remnant vegetation to understand their function in the persistence of fire-sensitive plants. Vegetation remnants (areas within the landscape that have not burned for a considerable amount of time) may occur stochastically or in more predictable locations (fire refuges) where physical conditions decrease fire severity. Our aim was to determine if remnant forests of the fire-sensitive conifer Austrocedrus chilensis are associated with biophysical attributes that allow persistence in a fire-prone Patagonian landscape. We conducted a multi-scale approach, determining attributes of forest remnants and their surroundings (matrices) through remote sensing and field-based biophysical and functional characteristics, and quantifying how tree survival probability relates to microsite conditions. Trees within remnants displayed abundant fire scars, were twofold older and had threefold larger growth rates than matrix trees. Remnants were associated with high rocky cover and elevated topographical positions. Tree survival increased in hilltops, eastern aspects, and with sparse vegetation. Trees within remnants experienced severe reductions in growth during droughts. Our results suggest that A. chilensis remnants are mainly the result of refuges, where environmental conditions increase fire survival, but also increase susceptibility to drought. A trade-off between fire survival and drought vulnerability may imply that under increasing drought and fire severity, locations that in the past have served as refuges may reduce their ability to allow the persistence of fire-sensitive taxa.

Experimental study of the types of cavitation by air seeding using light microscopy.

Recently, three types of cavitation: (i) expanding gradually; (ii) expanding-exploding, becoming a long-shaped bubble-lengthening by degrees; (iii) suddenly exploding and fully filling the conduit instantly, were proposed. Directed by this theory, experiments were performed using light microscopy to study the natural drying processes of xylem sections of Platycladus orientalis (L.) Franco. Three different phenomena of gas filling process in conduits were captured by replaying recorded videos. The first phenomenon is that a bubble emerging in a conduit expands and elongates gradually to fill the conduit. The second phenomenon is that a bubble emerging in a conduit expands gradually, and then suddenly becomes long-shaped, and extends continuously. The third phenomenon is that a bubble instantly fully fills a conduit. This paper suggests in these experiments that after losing the bulk water of a section, as the water stress of that section became more severe, the water pressures of different conduits of the section were not necessarily the same, and as time went on, the water pressures decreased constantly. Considering some practical factors, the three phenomena captured in our experiment are explained by our theory.

Evolutionary History of Atmospheric CO2 during the Late Cenozoic from Fossilized Metasequoia Needles.

The change in ancient atmospheric CO2 concentrations provides important clues for understanding the relationship between the atmospheric CO2 concentration and global temperature. However, the lack of CO2 evolution curves estimated from a single terrestrial proxy prevents the understanding of climatic and environmental impacts due to variations in data. Thus, based on the stomatal index of fossilized Metasequoia needles, we reconstructed a history of atmospheric CO2 concentrations from middle Miocene to late Early Pleistocene when the climate changed dramatically. According to this research, atmospheric CO2 concentration was stabile around 330-350 ppmv in the middle and late Miocene, then it decreased to 278-284 ppmv during the Late Pliocene and to 277-279 ppmv during the Early Pleistocene, which was almost the same range as in preindustrial time. According to former research, this is a time when global temperature decreased sharply. Our results also indicated that from middle Miocene to Pleistocene, global CO2 level decreased by more than 50 ppmv, which may suggest that CO2 decrease and temperature decrease are coupled.

Conifer species adapt to low-rainfall climates by following one of two divergent pathways.

Water stress is one of the primary selective forces in plant evolution. There are characters often cited as adaptations to water stress, but links between the function of these traits and adaptation to drying climates are tenuous. Here we combine distributional, climatic, and physiological evidence from 42 species of conifers to show that the evolution of drought resistance follows two distinct pathways, both involving the coordinated evolution of tissues regulating water supply (xylem) and water loss (stomatal pores) in leaves. Only species with very efficient stomatal closure, and hence low minimum rates of water loss, inhabit dry habitats, but species diverged in their apparent mechanism for maintaining closed stomata during drought. An ancestral mechanism found in Pinaceae and Araucariaceae species relies on high levels of the hormone abscisic acid (ABA) to close stomata during water stress. A second mechanism, found in the majority of Cupressaceae species, uses leaf desiccation rather than high ABA levels to close stomata during sustained water stress. Species in the latter group were characterized by xylem tissues with extreme resistance to embolism but low levels of foliar ABA after 30 d without water. The combination of low levels of ABA under stress with cavitation-resistant xylem enables these species to prolong stomatal opening during drought, potentially extending their photosynthetic activity between rainfall events. Our data demonstrate a surprising simplicity in the way conifers evolved to cope with water shortage, indicating a critical interaction between xylem and stomatal tissues during the process of evolution to dry climates.

Interspecific differences in whole-plant respiration vs. biomass scaling relationships: a case study using evergreen conifer and angiosperm tree seedlings.

PREMISE OF THE STUDY: Empirical studies and theory indicate that respiration rates (R) of small plants scale nearly isometrically with both leaf biomass (ML) and total plant biomass (MT). These predictions are based on angiosperm species and apply only across a small range of body mass. Whether these relationships hold true for different plants, such as conifers, remains unclear. METHODS: We tested these predictions using the whole-plant maintenance respiration rates and the biomass allocation patterns of the seedlings of two conifer tree species and two angiosperm tree species. Model Type II regression protocols were used to compare the scaling exponents (α) and normalization constants (ß) across all four species and within each of the four species. KEY RESULTS: The data show that the scaling exponents varied among the four species and that all differed significantly from isometry. For conifers, scaling exponents for R vs. MT, and R and ML were numerically smaller than those of the broadleaved angiosperm species. However, across the entire data set, R scaled isometrically with ML and with MT as predicted by the West, Brown, and Enquist (WBE) theory. We also observed higher respiration rates for small conifer seedlings compared to comparably sized angiosperm seedlings. CONCLUSIONS: Our data add credence to the view that the R vs. M scaling relationship differs among species, and that in general, the numerical values of this interspecific scaling relationship will depend on the species pooled in the analysis and on the range of body sizes within the data set.

Abscisic acid mediates a divergence in the drought response of two conifers.

During water stress, stomatal closure occurs as water tension and levels of abscisic acid (ABA) increase in the leaf, but the interaction between these two drivers of stomatal aperture is poorly understood. We investigate the dynamics of water potential, ABA, and stomatal conductance during the imposition of water stress on two drought-tolerant conifer species with contrasting stomatal behavior. Rapid rehydration of excised shoots was used as a means of differentiating the direct influences of ABA and water potential on stomatal closure. Pinus radiata (Pinaceae) was found to exhibit ABA-driven stomatal closure during water stress, resulting in strongly isohydric regulation of water loss. By contrast, stomatal closure in Callitris rhomboidea (Cupressaceae) was initiated by elevated foliar ABA, but sustained water stress saw a marked decline in ABA levels and a shift to water potential-driven stomatal closure. The transition from ABA to water potential as the primary driver of stomatal aperture allowed C. rhomboidea to rapidly recover gas exchange after water-stressed plants were rewatered, and was associated with a strongly anisohydric regulation of water loss. These two contrasting mechanisms of stomatal regulation function in combination with xylem vulnerability to produce highly divergent strategies of water management. Species-specific ABA dynamics are proposed as a central component of drought survival and ecology.

DNA sequence variation of drought-response candidate genes in Austrocedrus chilensis

Background: Austrocedrus chilensis (D. Don) Pic. Ser. et Bizzarri commonly known as Patagonian cypress is a member of the Cupressaceae family, characterized by a high adaptive potential for growing in marginal areas and good timber quality. The species grows over a wide area and under a wide range of rainfall. This study assessed adaptive genetic variation at SNP level in candidate genes involved in response to drought stress. Results: A total of 18 single nucleotide polymorphisms (SNPs) were found among 1,428 bp. Average nucleotide diversity value (π = 0.00312) was similar to those previously reported in other Cupressaceae. The Fst average among genes and populations was 0.163 and the lowest differentiation was observed in continuous and humid populations. A number of neutrality tests were applied to find evidence of positive selection in our candidate gene set, but only AcAQP2 gene in Pedregoso and San Ramón populations revealed significant departures from neutrality with positive values suggesting balancing selection. Conclusions: In this study we report the levels of nucleotide diversity searched in some drought stress candidate genes in Austrocedrus chilensis and the selective factors that may be acting on this species.

Old-growth Platycladus orientalis as a resource for reproductive capacity and genetic diversity.

AIMS: Platycladus orientalis (Cupressaceae) is an old-growth tree species which distributed in the imperial parks and ancient temples in Beijing, China. We aim to (1) examine the genetic diversity and reproductive traits of old-growth and young populations of P. orientalis to ascertain whether the older populations contain a higher genetic diversity, more private alleles and a higher reproductive output compared with younger populations; (2) determine the relationships between the age of the population and the genetic diversity and reproductive traits; and (3) determine whether the imperial parks and ancient temples played an important role in maintaining the reproductive capacity and genetic diversity of Platycladus orientalis. METHODS: Samples from seven young (younger than 100 yrs.) and nine old-growth (older than 300 yrs.) artificial populations were collected. For comparison, three young and two old-growth natural populations were also sampled. Nine microsatellite loci were used to analyze genetic diversity parameters. These parameters were calculated using FSTAT version 2.9.3 and GenAlex v 6.41. IMPORTANT FINDINGS: The old-growth artificial populations of P. orientalis have significantly higher genetic diversity than younger artificial populations and similar levels to those in extant natural populations. The imperial parks and ancient temples, which have protected these old-growth trees for centuries, have played an important role in maintaining the genetic diversity and reproductive capacity of this tree species.

[Measurement accuracy of granier calibration based on transpiration of Platycladus orientalis].

In order to understand the accuracy of Granier' s thermal dissipation method in measuring tree water consumption, a comparative study was made from May to June, 2010. The sap flow density of potted Platycladus orientalis was measured with thermal dissipation probe, which was compared with the whole-plant gravimetric measurement. There were significant linear relationships (R2 > 0.825) between the sap flow velocity in both north and south directions of P. orientalis measured by thermal dissipation probe and the transpiration rate measured by gravimetric method. The average daily sap flux in the north and south directions of P. orientalis were 10.6% and 15.1% lower than the daily average transpiration of P. orientalis, respectively, but the differences were not significant. Therefore, Granier's method had high reliability in the measurement of P. orientalis transpiration at daily scale, though the large temperature fluctuation between day and night could result in a lower daily sap flux than daily transpiration.

Distribution of living Cupressaceae reflects the breakup of Pangea.

Most extant genus-level radiations in gymnosperms are of Oligocene age or younger, reflecting widespread extinction during climate cooling at the Oligocene/Miocene boundary [∼23 million years ago (Ma)]. Recent biogeographic studies have revealed many instances of long-distance dispersal in gymnosperms as well as in angiosperms. Acting together, extinction and long-distance dispersal are likely to erase historical biogeographic signals. Notwithstanding this problem, we show that phylogenetic relationships in the gymnosperm family Cupressaceae (162 species, 32 genera) exhibit patterns expected from the Jurassic/Cretaceous breakup of Pangea. A phylogeny was generated for 122 representatives covering all genera, using up to 10,000 nucleotides of plastid, mitochondrial, and nuclear sequence per species. Relying on 16 fossil calibration points and three molecular dating methods, we show that Cupressaceae originated during the Triassic, when Pangea was intact. Vicariance between the two subfamilies, the Laurasian Cupressoideae and the Gondwanan Callitroideae, occurred around 153 Ma (124-183 Ma), when Gondwana and Laurasia were separating. Three further intercontinental disjunctions involving the Northern and Southern Hemisphere are coincidental with or immediately followed the breakup of Pangea.