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The tertiary relict Cercidiphyllum japonicum is an important canopy tree species of riparian forests in Japan, despite typically occurring at low densities. Once mature, canopy individuals are typically 30 cm in diameter at breast height and have high annual seed production. Seedlings tend to germinate on steep slopes with exposed soil, but not in...
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... forests (Ohno 2008), although it is mostly found in low abundance and pure stands are rare. Saplings are not often observed, implying that germination may be limited in forests. The species is long lived, often forming large canopies and multi-stemmed trunks (stools) through sprouting (i.e., producing shoots from the base or from roots) ( Fig. 4.2). It is possible that vegetative reproduction may compensate for low sapling recruitment, allowing C. japonicum to coexist at low density with other canopy ...
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... species, called "Katsura" in Japanese, is a sacred tree with ancient associations to moon, mountain, and water deities. Therefore, many large, multi-stemmed individuals are designated as natural monuments throughout the country (Fig. 4.3). Although C. japonicum is an important species both culturally and ecologically, its life history remains poorly understood. It is thought that populations are maintained on microtopography features created by various disturbances in riparian forests over long periods. However, such old-growth forests are now scarce due to extensive ...
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... japonicum is a tall, straight-stemmed species up to 30 m in height and 1 m in diameter (Fig. 4.1a). Associated canopy species within riparian forests include F. platypoda, P. rhoifolia, Aesculus turbinata, and Acer spp. Many individuals have multi-stemmed trunks, with wide, shoot-producing rootstocks (stools) and expansive individual canopies (Fig. 4.2). Leaves are borne on short and long shoots and are atypical in shape; those produced in spring are heart shaped, and those produced in summer are rhomboid (Fig. ...
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... C. japonicum individuals (12.6 individuals/ha) (Table 4.1) with a main stem !4 cm diameter at breast height (DBH), measured from the main (largest) stem in multi-stemmed individuals. Most individuals were in the canopy layer (n ¼ 47), with nine in the subcanopy layer, and three in the shrub layer. The DBH of main stems ranged from 5 to 153 cm ( Fig. ...
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... individuals were distributed within a V-shaped valley in the study area (Fig. 4.5), which had 30-degree slopes on the valley sides and roughly 12-degree slopes in sedimentary debris flow areas (Chap. 1). Alluvial fan and terrace debris flows in upstream areas contained rich soil with a litter layer, but there was little litter on terrace scarps, new landslide sites, old landslide slopes, and talus in downstream ...
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... valley than in the sedimentary debris flow areas upstream, and individuals were distributed over microtopography features such as sub-ridges, talus, and collapses ( Kubo et al. 2001a). Only 46 of the 59 individuals, 20 females and 26 males (Table 4.1), were mature, and female and male trees were randomly distributed relative to each other ( Fig. 4.5). Individuals were distributed along a stream in the study area from 1200 to 1600 m in elevation, and were found co-occurring with C. magnificum on a talus slope next to a stream above 1600 m (Fig. 4.6). Cercidiphyllum japonicum was found at <1650 m in elevation, but C. magnificum was found at elevations !1600 m, extending to the ...
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... canopy of C. japonicum turns purple-red at the time of blooming (Fig. 4.7), when individuals produce a large number of female or male flowers that lack a perianth at the tips of short shoots (Fig. 4.8). Blooming occurs in late April or early May in ...
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... canopy of C. japonicum turns purple-red at the time of blooming (Fig. 4.7), when individuals produce a large number of female or male flowers that lack a perianth at the tips of short shoots (Fig. 4.8). Blooming occurs in late April or early May in ...
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... categorized the 59 individuals of C. japonicum according to the DBH of their main stem (Fig. 4.9). All female and male trees reached the canopy layer, and except for one individual, all immature trees were found in the subcanopy and shrub layers (Table 4.1). All immature trees were < 26 cm in DBH, but one immature tree with a DBH of only 21 cm reached the canopy layer (Kubo and Sakio, unpublished data), implying that stem size may ...
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... late October and early November, C. japonicum leaves turn yellow ( Fig. 4.11). This is followed by seasonal defoliation and seed dispersal ( Fig. 4.12). Seed dispersal may begin as early as July, but it has been suggested that most seeds are immature at that time (Mizui 1993). Seeds are wind-dispersed, with larger dispersal events observed after seasonal defoliation. In the study area, the number of seeds ...
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... AE 12.2 per individual, n ¼ 59) of all tree species in the Ooyamazawa study area (Table 4.1). Small sprout stems (shoot stems) occurred circularly around large (main) stems, although the distribution pattern of stems varied (Figs. 4.18 and 4.19). When considering stems arising from shoots, most C. japonicum stems are small in diameter (n ¼ 55; Fig. 4.20), and individuals with larger main stems generally have a higher number of sprouts (R ¼ 0.37, n ¼ 59; Fig. 4.21; Kubo et al. 2001b). The mean number of sprouts per individual female, male, and immature tree was 8.9 AE 9.5, 12.5 AE 15.5, and 2.9 AE 4.4, respectively. Reproductive trees, both male and female, had a greater number of ...
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... et al. 2005), which was harvested from between the slope and stream at the bottom of the study area valley. We cut all 29 associated stems at 50 cm high in autumn 2001. Across the study area, individual trees had large root systems from which many sprouts could originate. Sprouts frequently surrounded the main stems at the center of the stool (Fig. 4.22), and many individuals had coalesced stems (11 of 29 sprouts). The ages of coalesced stems tended to be similar (Fig. 4.22). Sprouts approximately 30 years old were usually clustered around main stems at the upstream slope site, whereas sprouts in Survival rate (%) Relative illuminance (%) Fig. 4.16 Seasonal change in Cercidiphyllum ...
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... 29 associated stems at 50 cm high in autumn 2001. Across the study area, individual trees had large root systems from which many sprouts could originate. Sprouts frequently surrounded the main stems at the center of the stool (Fig. 4.22), and many individuals had coalesced stems (11 of 29 sprouts). The ages of coalesced stems tended to be similar (Fig. 4.22). Sprouts approximately 30 years old were usually clustered around main stems at the upstream slope site, whereas sprouts in Survival rate (%) Relative illuminance (%) Fig. 4.16 Seasonal change in Cercidiphyllum japonicum seedling survival rate under different relative light intensities at Ooyamazawa ( Kubo et al. 2000, revised) the ...
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... the main stems at the center of the stool (Fig. 4.22), and many individuals had coalesced stems (11 of 29 sprouts). The ages of coalesced stems tended to be similar (Fig. 4.22). Sprouts approximately 30 years old were usually clustered around main stems at the upstream slope site, whereas sprouts in Survival rate (%) Relative illuminance (%) Fig. 4.16 Seasonal change in Cercidiphyllum japonicum seedling survival rate under different relative light intensities at Ooyamazawa ( Kubo et al. 2000, revised) the downstream area tended to be older, around 80 years. Eighty-year-old sprouts from the slope site showed increased growth approximately 30 years ago, as determined by growth-ring ...
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... 4.16 Seasonal change in Cercidiphyllum japonicum seedling survival rate under different relative light intensities at Ooyamazawa ( Kubo et al. 2000, revised) the downstream area tended to be older, around 80 years. Eighty-year-old sprouts from the slope site showed increased growth approximately 30 years ago, as determined by growth-ring analysis (Fig. 4.23). This suggests that light conditions may have improved around that time, allowing many shoots to sprout or grow rapidly, and potentially increasing stem coalescence. Growth patterns varied by ...
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... included. Letters indicate significant differences between the light densities in each figure (t-test, P < 0.05). The level of significance was adjusted using the Holm's method. No seedlings survived the 3.0% RPPFD treatment over both years, and only one seedling remained in the 100.0% RPPFD treatment in the second year individual and with age ( Fig. 4.23) and sprout stem diameter was positively correlated with age (R 2 ¼ 0.66, n ¼ 45; Fig. ...
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... (t-test, P < 0.05). The level of significance was adjusted using the Holm's method. No seedlings survived the 3.0% RPPFD treatment over both years, and only one seedling remained in the 100.0% RPPFD treatment in the second year individual and with age ( Fig. 4.23) and sprout stem diameter was positively correlated with age (R 2 ¼ 0.66, n ¼ 45; Fig. ...
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... stems cut in 2001 produced new sprouts from their stools the following September (Fig. 4.25). The number of current-year sprouts was positively correlated with the age and diameter of the parent stems ( Kubo et al. 2005). Smaller, younger stems also produced new sprouts, and new sprouts would survive on the periphery of the stand under favorable light conditions. ...
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... show a proposed scheme for self-maintenance of C. japonicum by sprouting in Fig. 4.26. Following germination and growth, sprouts are produced as a result of endogenous factors, such as aging, or in response to external factors, such as gap formation and physical damage. Following stem death, C. japonicum is able to fill in ensuing gaps by sprouting. Consequently, colonies containing sprouts of various ages are ...
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... in ensuing gaps by sprouting. Consequently, colonies containing sprouts of various ages are produced, which are circularly distributed around the stool. By this process, broad, extensive canopies and multi-stemmed individuals can be produced from a single main stem. (Fig. 4.6), almost all C. japonicum and C. magnificum produced numerous sprouts (Fig. 4. Relationship between stem age and diameter ( Kubo et al. 2005, revised). Age and diameter were measured 50 cm from the ground C. japonicum (Fig. 4.28). The average DBH of the main stems of C. japonicum was significantly larger than that of C. magnificum (t-test, P < 0.01). A number of C. magnificum individuals only reached the ...
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... stool. By this process, broad, extensive canopies and multi-stemmed individuals can be produced from a single main stem. (Fig. 4.6), almost all C. japonicum and C. magnificum produced numerous sprouts (Fig. 4. Relationship between stem age and diameter ( Kubo et al. 2005, revised). Age and diameter were measured 50 cm from the ground C. japonicum (Fig. 4.28). The average DBH of the main stems of C. japonicum was significantly larger than that of C. magnificum (t-test, P < 0.01). A number of C. magnificum individuals only reached the subcanopy layer, despite their being mature ( Kubo et al. 2010). Dead stems in this species were numerous and often >30 cm DBH; the maximum recorded diameter ...
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... DBH of the main stems of C. japonicum was significantly larger than that of C. magnificum (t-test, P < 0.01). A number of C. magnificum individuals only reached the subcanopy layer, despite their being mature ( Kubo et al. 2010). Dead stems in this species were numerous and often >30 cm DBH; the maximum recorded diameter of a live stem was 45 cm (Fig. 4.27). This suggests that stems are pressured to reach optimal size to ensure survival. Cercidiphyllum are distributed in the montane and subalpine zones in Japan, and differences in stool structure between C. japonicum and C. magnificum reflect differences in the climatic conditions these species experience. Severe conditions such as wind ...
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... japonicum reaches reproductive maturity at approximately 30 cm DBH (Fig. 4.9), which is estimated to occur after 100 years of growth. Larger individuals with many large sprouts flower heavily in early spring and then disperse a large amount of winged seeds following annual seasonal defoliation in autumn (Figs. 4.12 and 4.13). Suitable germination sites for C. japonicum seedlings include bare soil and fallen ...
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... bare soil and fallen trees; sites with high litter accumulation or gravel are unsuitable for germination (Figs. 4.14 and 4.15) due to the small size of seeds and seedlings (Figs. 4.10 and 4.14). Germination does not imply survival for this species, given the high observed first-year mortality, which was likely a result of desiccation or stream Fig. 4.28 DBH size class distribution of stems including main stems and sprouts of Cercidiphyllum japonicum and Cercidiphyllum magnificum, co-occurring on an upper stream talus slope at Ooyamazawa ( Kubo et al. 2010, revised) flow and precipitation events (Fig. 4.16). Larger seedlings growing under bright light conditions may have a survival ...
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... the expansive canopies and rootstocks of parent trees, saplings of C. japonicum are uncommon (Fig. 4.4). This species occurs at low density in its riparian habitat (Fig. 4.5), but produces many small stems per individual (Figs. 4.20 and 4.21). Sprouts are continually produced as a result of endogenous and external factors, and C. japonicum can self-maintain for several hundreds of years by sprouting (Fig. 4.26), potentially compensating ...
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... the expansive canopies and rootstocks of parent trees, saplings of C. japonicum are uncommon (Fig. 4.4). This species occurs at low density in its riparian habitat (Fig. 4.5), but produces many small stems per individual (Figs. 4.20 and 4.21). Sprouts are continually produced as a result of endogenous and external factors, and C. japonicum can self-maintain for several hundreds of years by sprouting (Fig. 4.26), potentially compensating for low sapling recruitment with high vegetative ...
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... oldest specimen of F. platypoda, a coexisting canopy species, is 254 years ( Sakio 1997), and the lifespan of P. rhoifolia is approximately 120 years ( Kisanuki et al. 1992). We found that C. japonica stands had live main stems dating to 226 years (Fig. 4.22). Many stools in Ooyamazawa contain the remains of previous main stems, meaning that many individuals are likely several hundred years old. It is possible, therefore, that C. japonica maintains its populations by outliving its ...
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... In this study, high numbers of canopy plant species, including many accidental epiphyte tree species, were found in the lower crown at heights ranging 5-10 m where multiple and large reiterated trunks occurred, as well as on large horizontal branches of the host tree. These multiple reiterated trunks in large trees of C. japonicum is characteristic of this species, which extends its longevity by basal sprouting (Kubo and Sakio 2020). On these host tree structures, we found depositions of arboreal soils (ca. 10 cm deep), which have similar nitrogen availability to ground soils (Tatsumi et al. 2021), suggesting long-term accumulation of organic matter facilitates establishment of accidental epiphyte by providing nutrients and substrate for roots. ...
Large, trees have survived for a long time, and their complex crown structure can serve as habitat for epiphytic plants. Canopy plants are not as well studied in the temperate zones as in the tropics, because many of them are accidental epiphytes, epiphytic individuals of normally terrestrial species. We hypothesized that the canopy can serve as a refuge for terrestrial species that have difficulty establishing on the ground (e.g., insufficient light, deer over-browsing), promoting and conserving forest species diversity. Terrestrial species may also vary in their ability to adapt to a wide range of canopy growth conditions. Here, we investigated canopy vascular plants hosted on a large Cercidiphyllum japonicum tree in a temperate old-growth forest, Japan. The canopy plant community was diverse comprising 39 vascular plant species (6 of which were threatened species), including 31 accidental epiphytes and 8 obligate epiphytes. We found at least 14 species could escape deer over-browsing by establishing in the canopy. Most accidental epiphytes were found on multiple and large reiterated trunks bifurcated from the main trunk as well as on large horizontal branches. Leaves of canopy plants had higher nitrogen concentration compared to those of plants on the ground, and higher water-use efficiency with increasing height was achieved by controlling transpirational water loss. Our results show one large tree crown can be defined as a local hotspot for current and future plant species diversity in a temperate old-growth forest, reinforcing its ecological value for conservation purposes.
Annual seedfall was measured over a 24‐year period for three canopy species, Fraxinus platypoda , Pterocarya rhoifolia , and Cercidiphyllum japonicum , in a riparian forest in the Chichibu Mountains, central Japan. We collected seeds from 20 traps approximately every month, except in the winter. Fraxinus platypoda and P. rhoifolia seedfall fluctuated, with a 2‐year cycle. The seedfall of C. japonicum also fluctuated, but this species produced seeds every year, with an approximate 4‐year cycle. The simple moving average over 5 years for F. platypoda and P. rhoifolia was maintained at a constant level over the 24‐year period, but it gradually increased for C. japonicum . Each year's seedfall for the three species was influenced by the previous year's seedfall. The results for F. platypoda suggest that the fluctuations in the amount of seedfall were almost entirely dependent on fluctuations in stored resources, regardless of the previous year's weather. Conversely, the amount of seedfall for P. rhoifolia and C. japonicum was influenced by fluctuations in stored resources and the average temperature or precipitation in the summer of the year before flowering. These differences in seed production patterns were associated with differences in life history characteristics such as seed size and regeneration mechanisms.
To investigate the reproductive traits of the dioecious canopy tree Cercidiphyllum japonicum, we performed visual surveys from 2000 to 2007 to assess the flowering and fruiting of 59 individuals in a riparian forest in central Japan. Flowering and fruiting were assigned to ranked categories, where 1 indicated none and 5 indicated abundant production throughout the canopy. In addition, we used conical seed traps to assess the seed production of this species at the center of the forest. Of the 59 study individuals, 20 female and 26 male trees were >26 cm in diameter at breast height (DBH) and classified into the canopy layer. The remaining smaller individuals were non-reproductive and, excluding one, were classified into the sub-canopy layer. We estimated that C. japonicum may require 100 years of growth to reach 26 cm in DBH in a riparian forest. Both sexes produced abundant flowers in all years, and males exhibited slightly higher flowering and lower variation than females. Individuals with a DBH > 50 cm tended to flower very abundantly, and this pattern was consistent with trees over 1 m in DBH. Seed-trap surveys indicated that C. japonicum seeds were dispersed annually, with inter-annual variation. We conclude that C. japonicum is quite slow to reach sexual maturity, but can produce very abundant flowers, fruits, and seeds annually throughout its long lifespan.
The lower Campanian members 2 and 3 of the Nenjiang Formation host a diverse array of cupressoid conifers (Glyptostrobus, Sequoiadendron, Cupressinocladus, Mesocyparis; commonly shoots and a few cones), a few unassigned conifers and putative ginkgophytes. True aquatics (Cobbania, Quereuxia) and water-bound plants (Nelumbites and Sparganiophyllum) have been recorded, as well as foliage and putative flowers of the deciduous angiosperm tree Trochodendroides. The floral composition is very similar to assemblages from the adjacent Zeya-Bureya Basin (Far East Russia), and it contains elements of contemporaneous floras from further away regions (Sakhalin, Honshu, Chukotka, Kamchatka), indicating an affiliation of the Nenjiang Flora to the Santonian–Campanian floras of north-east Asia, and corroborating a Campanian age for the examined strata of the Nenjiang Formation. Ecological assessments indicate a flora that thrived in and along the shores of a comparatively large lake that supported populations of three to four different species of free- or partly floating aquatics.
