In a warming climate, temperature-sensitive plants must move toward colder areas, that is, higher latitude or altitude, by seed dispersal [1]. Considering that the temperature drop with increasing altitude (-0.65°C per 100 m altitude) is one hundred to a thousand times larger than that of the equivalent latitudinal distance [2], vertical seed dispersal is probably a key process for plant escape from warming temperatures. In fact, plant geographical distributions are tracking global warming altitudinally rather than latitudinally, and the extent of tracking is considered to be large in plants with better-dispersed traits (e.g., lighter seeds in wind-dispersed plants) [1]. However, no study has evaluated vertical seed dispersal itself due to technical difficulty or high cost. Here, we show using a stable oxygen isotope that black bears disperse seeds of wild cherry over several hundred meters vertically, and that the dispersal direction is heavily biased towards the mountain tops. Mountain climbing by bears following spring-to-summer plant phenology is likely the cause of this biased seed dispersal. These results suggest that spring- and summer-fruiting plants dispersed by animals may have high potential to escape global warming. Our results also indicate that the direction of vertical seed dispersal can be unexpectedly biased, and highlight the importance of considering seed dispersal direction to understand plant responses to past and future climate change.

Vertical seed dispersal, i.e. seed dispersal towards a higher or lower altitude, is considered a critical process for plant escape from climate change. However, studies exploring vertical seed dispersal are scarce, and thus, its direction, frequency, and mechanisms are little known. In the temperate zone, evaluating vertical seed dispersal of animal-dispersed plants fruiting in autumn and/or winter is essential considering the dominance of such plants in temperate forests. We hypothesized that their seeds are dispersed towards lower altitudes because of the downhill movement of frugivorous animals following the autumn-to-winter phenology of their food plants which proceeds from the mountain tops to the foot in the temperate zone. We evaluated the vertical seed dispersal of the autumn-fruiting wild kiwi, Actinidia arguta, which is dispersed by temperate mammals. We collected dispersed seeds from mammal faeces in the Kanto Mountains of central Japan and estimated the distance of vertical seed dispersal using the oxygen isotope ratios of the dispersed seeds. We found the intensive downhill seed dispersal of wild kiwi by all seed dispersers, except the raccoon dog (bear: mean −393.1 m; marten: −245.3 m; macaque: −98.5 m; and raccoon dog: +4.5 m). Mammals with larger home ranges dispersed seeds longer towards the foot of the mountains. Furthermore, we found that seeds produced at higher altitudes were dispersed a greater distance towards the foot of the mountains. Altitudinal gradients in autumn-to-winter plant phenology and other mountain characteristics, i.e. larger surface areas and more attractive human crops at lower altitudes compared to higher altitudes, were considered drivers of downhill seed dispersal via animal movement. Strong downhill seed dispersal by mammals suggests that populations of autumn-to-winter fruiting plants dispersed by animals may not be able to sufficiently escape from current global warming in the temperate zone.

This study examined the vertical gradient of radar reflectivity below the detected bright band in stratiform regions from the Tropics to the extratropical latitudes using data from the Ku‐band (13.6 GHz) precipitation radar on board the Global Precipitation Measurement (GPM) Core Observatory. Stratiform precipitation profiles with reflectivity decreasing (increasing) from the melting level toward the surface occur frequently in the tropical oceans (mid‐ and high‐latitude oceans). High fractions of downward increasing stratiform pixels are found over the North Pacific Ocean throughout the year and over East Asia except for winter. In contrast, the North American continent and the adjacent North Atlantic Ocean are characterized by low fractions of downward‐increasing pixels during summer. The difference is consistent with the dominant type of convection over East Asia (warm‐type clouds) and over the North American continent (cold‐type clouds). Even in the tropical oceans such as the Atlantic and eastern Pacific intertropical convergence zones, there are some areas with moderate fractions of downward‐increasing stratiform pixels where the warm rain process dominates. The downward reduction of reflectivity in the stratiform region of MCSs is obviously due to evaporation, which is a function of lower‐tropospheric relative humidity. The downward increase of reflectivity in stratiform regions over the midlatitude oceans appears the result of raindrop growth. This is achieved via the collection of cloud droplets while falling through low‐level clouds produced by large‐scale vertical motion in the lower troposphere due to large‐scale convergence associated with synoptic‐scale systems.

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