林窗面积对香椿生长及光合生理特性的影响

研究了川中丘陵区柏木人工林内人造林窗面积[50 m2(Ⅰ),100 m2(Ⅱ)和200 m2(Ⅲ),旷地(CK)]对香椿生长、光合特性及相关生理特征的影响。结果表明:林窗面积对香椿的树高、地径和冠面积影响显著,以林窗Ⅱ最大,CK最小;随着林窗面积的增加,香椿的日均净光合速率增加,但午时的净光合速率初始量子效率、近光饱和点、最大净光合速率、光氮利用率、光能利用效率都显著降低,光合能力下降;香椿的叶绿素a/b、叶片厚度、比叶鲜重、比叶干重、单位面积叶氮含量、丙二醛、超氧化物歧化酶及可溶性蛋白含量随着林窗面积的增加而增加,而叶绿素a、叶绿素b和类胡萝卜素含量都降低,表明香椿对因林窗面积变化而导致的光强改变有较强的适应性,这在一定程度上促进了生长及光合能力的差异;旷地内香椿的"光合午休"及林窗内日均Pn的降低可归因于非气孔因素限制。说明香椿对生境的变化适应性较强,而且100 m2的林窗在其形成初期能有效促进香椿维持较高的生产力和光合能力。     

马尾松人工林林窗边缘效应对油樟化学计量特征的影响

马尾松(Pinus massoniana)是长江上游低山丘陵区退耕还林的一个主要造林树种。为了解林窗调控下,马尾松人工林更新优势种在化学计量特征水平上受边缘效应的影响,该文选取四川省宜宾市高县马尾松人工林人工砍伐形成的7种面积不等的林窗,以林下为对照,分析了人工更新优势植物油樟(Cinnamomum longepaniculatum)叶片化学计量特征及其季节动态。结果表明:马尾松人工林林下与不同面积林窗边缘相比,不同季节各林窗边缘油樟叶片碳(C)含量、碳氮比(C:N)和碳磷比(C:P)均显著高于林下。随着林窗面积增大,油樟叶片C含量、C:N、C:P和N:P均先升后降,在中型林窗(400–900 m~2)边缘出现最大值。从春季到冬季,油樟叶片N、P含量显著下降后上升,C:N和C:P显著上升后下降,均在夏季有拐点。夏秋季林窗边缘油樟对养分吸收利用优于春冬季,边缘效应更显著。主成分分析结果显示,马尾松人工林不同大小林窗边缘油樟化学计量特征主要受林窗面积、相对光强及月平均气温影响。研究结果表明:625 m~2林窗边缘油樟有机质储存及养分利用效率在不同季节均有最大值,在化学计量特征水平上具有更显著的边缘效应。     

常绿阔叶树种栲树幼苗对不同光环境的光合生理响应

常绿阔叶树种栲树是亚热带常绿闭叶林中的优势种之一.对比研究了林下、林窗和旷地3个光环境下栲树幼苗的光合特征参数、比叶质量、叶氮含量及叶解剖结构等,结果表明:林下栲树幼苗的光补偿点和暗呼吸速率低,比叶质量小,表观量子效率和光合氮素利用率高,利用弱光的能力强,在低光环境下保持了较高的光合能力;海绵组织发达且表皮细胞厚,是对凉爽环境的积极响应.在旷地,栲树的最大光合速率明显下降,暗呼吸速率显著升高,单位面积的叶氮含量较高但氮素利用率较低;叶厚度和栅栏组织厚度均增加,栅栏组织/海绵组织厚度比值也升高了,但栅栏组织不如海绵组织发达,不能适应高光环境.栲树幼苗在林窗有较高的光合能力,在林窗中能正常生长和更新.     

不同光环境对桤木幼苗生长和光合特性的影响

通过设置3个光照强度(100%、56.2% 和12.5%),模拟森林幼苗生长的旷地(采伐迹地)、林窗和林下光环境,研究不同光照强度对外来种台湾桤木和乡土种四川桤木幼苗的生长、光合特性以及生物量积累与分配的影响.结果表明： 低光环境限制了两种桤木幼苗形态指标的增长,适当遮荫的林窗环境比旷地更有利于幼苗的生长.台湾桤木幼苗具有较高的比叶面积和相对生长速率,较大的单叶面积、叶长、叶宽、株高和基径,较少的叶片数和较低的叶面积比、叶柄长.低光环境下,台湾桤木幼苗的最大净光合速率、光饱和点和表观光量子效率较高,光补偿点和暗呼吸速率较低.随着光照强度的降低,台湾桤木幼苗具有更高的根生物量比和更低的叶生物量比;四川桤木幼苗则相反,加剧了动物取食和机械损伤的风险.     

不同光环境对桤木幼苗生长和光合特性的影响

通过设置3个光照强度(100％、56.2％和12.5％),模拟森林幼苗生长的旷地(采伐迹地)、林窗和林下光环境,研究不同光照强度对外来种台湾桤木和乡土种四川桤木幼苗的生长、光合特性以及生物量积累与分配的影响.结果表明:低光环境限制了两种桤木幼苗形态指标的增长,适当遮荫的林窗环境比旷地更有利于幼苗的生长.台湾桤木幼苗具有较高的比叶面积和相对生长速率,较大的单叶面积、叶长、叶宽、株高和基径,较少的叶片数和较低的叶面积比、叶柄长.低光环境下,台湾桤木幼苗的最大净光合速率、光饱和点和表观光量子效率较高,光补偿点和暗呼吸速率较低.随着光照强度的降低,台湾桤木幼苗具有更高的根生物量比和更低的叶生物量比;四川桤木幼苗则相反,加剧了动物取食和机械损伤的风险.     

不同光强下生长的两种榕树叶片光合能力与比叶重、氮含量及分配的关系

观测了不同光强下生长的两种榕树幼苗叶片的光合能力、比叶重(LMA)、氮含量及在光合机构中的分配.结果表明,两种榕树幼苗的最大净光合速率(Pmax)均随生长光强的升高而升高,这与LMA、单位面积氮含量(NA)和光合氮利用效率随生长光强的升高而升高有关.除36%光强下外,相同光强下生长的喜光的斜叶榕(Ficus tinctoria)的Pmax均显著高于耐荫的假斜叶榕(Ficus subulata),这与其叶片中氮在羧化组份和生物力能学组份中的分配系数、LMA和NA较高有关.     

不同光环境下亚热带常绿阔叶树种和落叶阔叶树种幼苗的叶形态和光合生理特征

由于地球环境的演变,亚热带常绿阔叶林中常有落叶阔叶成分的存在,但从生态角度,其存在的机理尚不清楚.通过比较研究常绿阔叶优势树种大头茶与落叶阔叶树种枫香幼苗的叶形态和光合生理特征对不同光环境(旷地、林窗、林下)的响应,尝试解释落叶成分遗留的基础.结果表明:①枫香具有较小的比叶重(LMA)和较高的光合氮利用效率(PNUE),在高光环境中,并未受到光抑制,具有较高的光合可塑性,尤其在林窗,表现出较大头茶高3倍的光合能力(Pnmax),保证了其在较短的生长季节积累更多的光合产物,增强与常绿树种的竞争力;②常绿阔叶树种大头茶具有较大的LMA,在旷地受到严重的光抑制,将更多的氮用于化学防御中.在林窗和林下具有相对高的Pnmax,生长幅度较广,但在林下将更多的氮投入到比生长更为重要的生存消耗中.而枫香在林下响应表现为Rubisco活性降低,光合受阻;③两树种在林窗均表现出较高的光合适应性,具有最大的光合能力(Pnmax)及合理的氮在光合机构中的分配系数,说明林窗是它们更新的最佳环境.但枫香的光合作用更依赖于光照,而大头茶则更依赖于CO2浓度;④总之,落叶阔叶树种以其较高的形态可塑性和对高光的光合生理可塑性能够在常绿阔叶林种生存、生长,并成为常绿阔叶林中固有成分的主要原因之一.     

华南典型人工林凋落物的持水特性

对于华南地区的杉木 (Cunninghamialanceolata) 林、马尾松 (Pinusmassoniana) 林、湿地松 (Pinuselliottii) 林、马占相思 (Acaciamangium) 林和尾叶桉 (Eucalyptusurophylla) 林的凋落物的储量、持水量、持水率和吸水速率进行了研究。结果表明, 杉木林的凋落物的干重 (6.5× 10 3 kg·hm-2 ) 最大, 其次是马尾松林和马占相思林 (5.5× 10 3 kg·hm-2 ), 而湿地松林 (4.1× 10 3 kg·hm-2 ) 和尾叶桉林较小 (4.0× 10 3 kg·hm-2 ) 。凋落物持水量呈现杉木林 >马占相思林 >尾叶桉林 >马尾松林 >湿地松林。各林分凋落物的最大持水量为杉木林 17.9× 10 3 kg·hm-2, 马占相思林 14.8× 10 3 kg·hm-2, 尾叶桉林 14.0× 10 3 kg·hm-2, 马尾松林 10.6× 10 3 kg·hm-2, 湿地松林 9.8× 10 3 kg·hm-2 。尾叶桉林、杉木林、马占相思林、湿地松林和马尾松林的凋落物最大持水率分别为 35 1%、2 74 %、2 6 9%、2 35 %和 191%。凋落物持水量和凋落物持水率随着浸泡时间的增加按照对数方程增加。 5种林分中尾叶桉林的凋落物吸水速率在各浸泡时间后居首位, 杉木林和马占相思林中等, 湿地松林较小, 而马尾松林最小, 各林分的凋落物的吸水速率随浸泡时间的增长按方程Y =a +b·t-1下降。     

中亚热带典型人工林土壤酶活性及其化学计量特征

以中亚热带典型的马尾松林、湿地松林和马尾松-木荷混交林(针阔混交林)为研究对象,分析不同林分类型下0～10和10～20 cm土层的β-D-葡萄糖苷酶(BG)、β-N-乙酰氨基葡萄糖苷酶(NAG)、亮氨酸氨基肽酶(LAP)、酸性磷酸酶(AP)、多酚氧化酶(POX)、过氧化物酶(POD)6种土壤酶活性,以及酶化学计量比及土壤理化性质特征,分析驱动中亚热带典型林分类型土壤酶活性及其计量比变异的主要因素。结果表明: 林分类型显著影响了土壤BG和LAP活性,表现为湿地松林10～20 cm土层土壤BG显著高于马尾松林,而LAP在马尾松林最高;湿地松林10～20 cm土层土壤BG/(NAG+LAP)、BG/AP显著高于马尾松林,而马尾松林(NAG+LAP)/AP显著高于湿地松林和针阔混交林;林分类型间酶化学计量的向量长度在10～20 cm土层差异显著,表现为湿地松林>针阔混交林>马尾松林。3种人工林酶化学计量的向量角度均大于45°,其中在湿地松林10～20 cm土层向量角度显著大于马尾松林。冗余分析表明,土壤碳质量指数和有机碳与全磷的比值(C/P)以及土壤含水量和C/P分别是0～10和10～20 cm土层土壤酶活性及其化学计量的关键影响因素,土壤碳和磷的数量和质量,以及土壤含水量在调节中亚热带人工林生态系统养分循环中发挥关键作用。     

南亚热带尾巨桉人工林土壤种子库特征研究

为了解桉树人工林土壤种子库特征,对不同林龄尾巨桉(Eucalyptusurophylla×E.grandis)人工林种子库的储量、垂直分布特征及与林下植被的相似度进行了分析。结果表明,土壤活力种子储量最高的为1～2 a生桉林,显著高于其他林型;其次为3～4 a生桉林和马尾松(Pinus massoniana)林;最低的为杉木(Cunninghamia lanceolata)林,显著低于其他林型。随林龄增加,尾巨桉林土壤种子库储量快速下降。土壤种子库中植物种类最为丰富的是杂木林和马尾松林,显著大于其他林型。随林龄增加,尾巨桉林土壤种子库植物种类先增后降。所有林型中,0～5 cm土壤种子库的密度均显著高于5～10 cm土层。杂木林种子库和林下植被共存植物种类最多,其次是马尾松林,杉木林最少。尾巨桉人工林随林龄的增加,共存植物种数呈先升高后下降趋势,土壤种子库Jaccard (CJ)和Sorensens (CS)相似系数也呈先升后降的趋势。因此,在速生桉人工林经营中适当间种(保留)乡土树种,可增加森林生态系统的生物多样性和生态功能的稳定性。     

林隙对香樟和枫香幼苗早期生长阶段功能性状的影响

探究林隙对不同需光性树种早期生长特征和功能性状的影响,对揭示林隙微生境影响次生林内幼苗更新机制具有重要意义。以亚热带次生林中耐荫常绿树种香樟和阳性落叶树种枫香幼苗为试验对象,研究大林隙(D/H介于1.5—2.0)、中林隙(D/H介于1.0—1.5)和小林隙(D/H介于0.5—1.0)对不同需光树种幼苗早期(1—3年生)生长特征和功能性状的影响。结果表明：(1)林隙大小对两种幼苗的生长均有显著影响。其中,中林隙可显著促进香樟2—3年生幼苗的生长,大林隙对枫香1—3年生幼苗的生长均具有显著促进作用。(2)对林隙环境因子与幼苗功能性状的关系进行冗余分析表明,香樟幼苗功能性状的变化与林隙土壤有机质含量、水解性氮含量、酸碱度和有效磷含量密切相关,而枫香幼苗功能性状则主要受林隙土壤酸碱度、有机质含量、水解性氮含量、土壤含水率、冠层透光率和土壤有效磷含量的影响。(3)维持较高的根重比、细根比根长、叶碳氮比和叶碳磷比是幼苗应对林隙环境影响的重要生理生态调节机制。     

Different responses to shade of evergreen and deciduous oak seedlings and the effect of acorn size

An evergreen oak species, Cyclobalanopsis multinervis, and a deciduous oak species, Quercus aliena var. acuteserrata were grown from acorns under two light levels (full sunlight and shade at about 18 % of full sunlight, simulating the light intensities in forest clearings and gaps, respectively) for one growing season. Three hypotheses were tested: (i) the deciduous species grows faster than the evergreen species in forest gaps and clearings; (ii) the deciduous species responds more strongly in terms of growth and morphology to variation in light climate than the evergreen species; and (iii) seedling size is positively correlated to acorn size. The results showed: (i) at both light levels, the deciduous seedlings gained significantly more growth in biomass and height than the evergreen seedlings; (ii) both species produced significantly more biomass in full sunlight than in shade, without showing any significant difference in height between treatments. Increase in light intensity improved the growth of the deciduous seedlings more strongly; (iii) at a similar age, the deciduous seedlings showed a greater response in leaf morphology and biomass allocation to variation in light levels, but when compared at a similar size, biomass allocation patterns did not differ significantly between species; (iv) bigger acorns tended to produce larger seedlings, larger leaf sizes and more leaf area, between and within species. These differences demonstrate that the deciduous species is gap-dependent and has the advantage over the evergreen species in forest gaps and clearings.     

林窗对川西周公山柳杉人工林林下物种多样性的影响

采用典型样地法,以川西周公山柳杉人工林5种不同大小的林窗为研究对象,以林下非林窗为对照,研究了不同大小的林窗对柳杉人工林物种多样性的影响,同时分析了不同梯度林窗下林窗中心、林窗边缘、及林下群落的物种组成、物种多样性的变化情况。结果表明:(1)在所调查的18个样地231个样方中共记录到维管束植物141种,隶属于76科113属;随着林窗面积的增大,群落各层次的物种数呈现出先升高后降低的趋势,灌木层物种数在各林窗梯度上表现为林缘林下林窗中心,草本层物种数在各林窗梯度上表现为林缘林窗中心林下。(2)不同林窗优势种及其重要值不同,即在小林窗内,优势种为柳杉和野桐,其重要值之和高达0.292 3;在大林窗内,杉木及亮叶桦为群落优势物种,群落内出现大量其更新幼苗。(3)不同大小的林窗表现为灌木层物种丰富度指数(D)、Shannon-Wienner指数(H)、和Pielou均匀度指数(Jsw)值在400~450 m2面积的大林窗内达到一个均优水平,草本层物种的多样性在面积为100~150m2的小林窗内达到较高水平;不同梯度的林窗各层次群落D、H值整体表现为林缘林窗中心林下。研究认为:林窗的存在会改变群落物种组成,提高群落物种多样性水平,并且大林窗(400~450m2)更利于柳杉人工林林下树种更新及物种多样性的提高。     

林窗尺度对侧柏人工林土壤微生物量和功能多样性的影响

微生物活性是影响土壤碳循环等地下生态系统过程的重要因素。以徐州市侧柏(Platycladus orientalis(L.)Franco)人工林为研究对象,以未受干扰的侧柏林为对照(CK),设置半径分别为4、8、12m的3种尺度近圆形林窗,从林窗边缘(D1)到距林缘4m(D2)及8m处林下(D3)水平梯度上分析土壤微生物量碳(MBC)、氮(MBN)和代谢功能多样性的变化。结果表明:1)与CK相比,林窗样地土壤MBC总体降低,MBN含量显著下降(P0.05),MBC/MBN显著上升(P0.05)。在3种尺度林窗中,MBC在大林窗偏小,MBN在小林窗偏小;MBC/MBN总体上随林窗尺度增大而减小。2)与CK相比,大中林窗降低了土壤微生物代谢活性(AWCD),小林窗变化不大。从D1到D3点,小林窗的AWCD先降后升,中林窗呈上升趋势,大林窗则相反。而且林窗降低了土壤微生物对各类碳源的利用,主要利用聚合物类和氨基酸类碳源,中林窗样点对碳水化合物和氨基酸类小分子碳源的利用最低。3)林窗总体提高了土壤微生物功能多样性,其中多样性(H')、丰富度(S)和均匀度(E)3个指数在各点之间均无显著差异,小林窗和CK的优势度指数(D_s)显著大于(P0.05)大林窗。侧柏林人工林窗对土壤微生物量和功能多样性的影响有着明显的尺度和位置梯度效应,林窗有望促进侧柏林土壤碳固持和大分子物质降解,提高其应对全球气候变化的能力,综合而言,中尺度林窗对侧柏林生态功能的发挥较为有利。     

马尾松(Pinus massoniana)人工林林窗对土壤不同形态活性有机碳的影响

研究了四川盆地低山丘陵区马尾松人工林不同大小林窗对表层土壤活性有机碳(水溶性有机碳、微生物量碳、易氧化碳)含量、分配比例及碳库管理指数的影响。结果表明:(1)林窗下土壤微生物量碳含量与分配比例较林下土壤有所升高,而水溶性有机碳与易氧化碳含量及水溶性有机碳分配比例有所降低。(2)林窗大小显著影响林窗中心土壤活性有机碳含量与分配比例。随林窗面积增大,水溶性有机碳、微生物量碳与易氧化碳含量呈现较为一致的升高趋势;水溶性有机碳和微生物量碳分配比例也升高,易氧化碳分配比例先下降后升高,稳定态碳先升高后降低;总体表现为较大林窗(900—1225m2)微生物活性强,活性有机碳含量高,且有机碳库稳定性较好。(3)土壤碳库管理指数随林窗面积增大无显著变化,但与各形态活性有机碳含量及总有机碳含量显著相关,说明土壤碳库管理指数能够相对全面地反映林窗大小对土壤碳库的影响。     

马尾松人工林林窗大小对四种凋落叶质量损失和养分释放的影响

马尾松人工林乔木层植物凋落物的分解对林地养分平衡和系统物质循环具有重要意义,并可能受不同大小林窗下微环境差异的影响。采用凋落物袋分解法,以马尾松(Pinus massoniana)人工林人为砍伐形成的7个不同大小林窗(G1:100 m~2、G2:225 m~2、G3:400 m~2、C4:625 m~2、G5:900 m~2、G6:1225m~2、G7:1600 m~2)为研究对象,林下(G0)为对照,研究林窗大小对红椿(Toona ciliata)、桢楠(Phoebe zhennan)、香樟(Cinnamomum camphora)和马尾松4种乡土树种凋落叶质量损失及养分释放的影响。结果显示:1)林窗大小(G0-G7)显著影响林窗中心放置的红椿和桢楠凋落叶N和P释放率、香樟凋落叶失重率和N、P、K释放率以及马尾松凋落叶P和K释放率。相对于林下,中小型林窗(G1-G4)的凋落叶失重率和N、P释放率明显较大,而大型林窗(G6-G7)的凋落叶K释放率明显较大。2)林窗内放置位置显著影响红椿、桢楠和马尾松凋落叶的K释放率及香樟凋落叶的P释放率。红椿和桢楠的凋落叶K释放率从林窗中心到边缘显著减少,而马尾松凋落叶K释放率及香樟P释放率从林窗中心到边缘显著增加。3)4种凋落叶类型中红椿凋落叶分解最快,其分解50%和95%所需时间分别为5.29和23.14个月。上述结果表明,林窗大小和林窗内位置对凋落物质量损失及其养分释放具有显著影响,但影响大小及趋势随物种初始基质质量的差异具有明显变化,研究结果为亚热带低山丘陵区马尾松人工低效林的科学经营及管理提高了一定的科学依据。     

Invasion of broad-leaf tree species into a larch plantation: seasonal light environment, photosynthesis and nitrogen allocation

We have studied how tree seedlings with differing leaf phenological traits change their nitrogen allocation in seasonally changing light environments. Specifically, we have investigated seasonal changes in maximum leaf photosynthetic rates in situ, and changes in nitrogen, chlorophyll and ribulose-1,5-bisphosphate carboxylase oxygenase (Rubisco) content. We have also estimated nitrogen allocation characteristics from leaf nitrogen to photosynthetic proteins, light-harvesting protein and other proteins in four successional tree species growing in a larch plantation. The gap phase species, Magnolia hyporeuca Siebold et Zucc., displays higher photosynthetic rates to compensate for the short leafy period and allocates more nitrogen to Rubisco. In contrast, tree species that grow at the forest floor, Prunus ssiori F. Schmidt and Carpinus cordata Blume, use the leafless period of the canopy trees to change their allocation of nitrogen from Rubisco to chlorophyll in line with canopy closure. Quercus mongolica Fischer ex Ledeb. var. crispula (Blume) Ohashi is a mid-successional tree species, and maintains a high photosynthetic rate in September because it allocates more nitrogen to Rubisco. We conclude that phenological differences in tree seedlings are clearly reflected in the nitrogen allocation pattern at gaps and at the forest floor in larch plantations.     

林窗尺度对侧柏人工林土壤微生物生物量碳氮的短期影响

为了明晰林窗尺度对侧柏人工林土壤微生物生物量碳氮的短期影响,以徐州市50年生侧柏人工林为研究对象,探讨了开设林窗2a后3种林窗尺度(半径分别为4m,S林窗;8m,M林窗;12m,L林窗)和位置(林窗内、林缘和林窗外部)对土壤微生物生物量碳氮(MBC,Microbial Biomass Carbon;MBN,Microbial Biomass Nitrogen)的影响。结果表明:(1)与对照样地相比,L林窗显著提高了春季(207.1mg/kg)和夏季(169.5mg/kg)林窗外部的土壤MBN含量;M林窗显著提高了春季林窗内部(2959.3mg/kg)和林缘(3008.8mg/kg)位置土壤MBC含量和林缘位置(207.7mg/kg)土壤MBN含量,且显著提高了夏季林窗内部(144.4mg/kg)土壤MBN含量;S林窗显著降低了春季林窗外部和林缘位置土壤MBC(分别为2159.2mg/kg和1955.1mg/kg)和MBN(分别为153.1mg/kg和131.3mg/kg)含量。(2)土壤MBC含量与土壤全碳和土壤可溶性有机碳(DOC)含量呈极显著(P0.01)正相关,与土壤温度呈极显著负相关(在3—27℃之间);土壤MBN含量与土壤含水量和DOC含量呈极显著正相关,与土壤全碳呈显著正相关;土壤MBC和MBN含量与凋落物量没有显著相关关系。本研究中,相对于S和L林窗,M林窗对土壤微生物生物量的提高作用较为明显,可促进侧柏人工林土壤碳氮循环过程,在徐州侧柏人工林中开设M林窗有利于提高土壤肥力和林木生长。     

Gap dynamics and regeneration strategies in <Emphasis Type="Italic">Juniperus-Laurus</Emphasis> forests of the Azores Islands

We asked the following questions regarding gap dynamics and regeneration strategies in Juniperus-Laurus forests: How important are gaps for the maintenance of tree diversity? What are the regeneration strategies of the tree species? Thirty canopy openings were randomly selected in the forest and in each the expanded gap area was delimited. Inside expanded gaps the distinction was made between gap and transition zone. In the 30 expanded gaps a plot, enclosing the gap and transition zone, was placed. In order to evaluate the differences in regeneration and size structure of tree species between forest and expanded gaps, 30 control plots were also delimited in the forest, near each expanded gap. In the 60 plots the number of seedlings, saplings, basal sprouts and adults of tree species were registered. Canopy height and width of adult individuals were also measured. The areas of the 30 gaps and expanded gaps were measured and the gap-maker identified. Juniperus-Laurus forests have a gap dynamic associated with small scale disturbances that cause the death, on average, of two trees, mainly of Juniperus brevifolia. Gap and expanded gap average dimensions are 8 and 25 m², respectively. Gaps are of major importance for the maintenance of tree diversity since they are fundamental for the regeneration of all species, with the exception of Ilex azorica. Three types of regeneration behaviour and five regeneration strategies were identified: (1) Juniperus brevifolia and Erica azorica are pioneer species that regenerate in gaps from seedlings recruited after gap formation. However, Juniperus brevifolia is a pioneer persistent species capable of maintaining it self in the forest due to a high longevity and biomass; (2) Laurus azorica and Frangula azorica are primary species that regenerate in gaps from seedlings or saplings recruited before gap formation but Laurus azorica is able to maintain it self in the forest through asexual regeneration thus being considered a primary persistent species; (3) Ilex azorica is a mature species that regenerates in the forest.     

Photosynthetic plasticity of two rain forest shrubs across natural gap transects

Summary Photosynthetic plasticity of two congeneric shrub species growing under natural field conditions was compared along transects spanning two canopy gaps in a Costa Rican rain forest. Piper arieianum is a shadetolerant species common in successional and mature forests, whereas P. sancti-felicis is a pioneer species abundant in abandoned clearings and large gaps. Twenty potted cuttings of each species were placed at regular intervals along two east-west transects crossing a small branch-fall gap and a large tree-fall gap. Along the transects, the percent of full sun photon flux density varied from less than 2% to 45%. After six months of growth under these conditions, leaves were monitored for incident photon flux density, photographic measures of light availability, photosynthetic capacity (A_max), leaf nitrogen content, leaf chlorophyll content, and specific leaf mass. Although both species demonstrated considerable plasticity in A_max across gap transects, P. sancti-felicis leaves had a superior capacity to track closely variation in light availability, particularly in the larger gap. For regressions of A_max on measures of light availability, P. sancti-felicis consistently showed a 3.5 to 5-fold higher coefficient of determination (R²) and a 3 to 4-fold higher slope than P. arieianum. In both species leaf nitrogen content per leaf area increased significantly with light availability, although P. sancti-felicis, again, showed a much stronger relationship between these variables. Across the transects, mean chlorophyll content per unit leaf area did not differ significantly between the species, whereas mean chlorophyll content per unit leaf dry mass was 3-times greater in leaves of P. sancti-felicis. Piper arieianum exhibited highly significant increases in chlorophyll a:b ratio with increased light availability, whereas P. sancti-felicis lacked significant variation in this trait across a gradient of light availability. Mean specific leaf mass did not vary significantly between species across the gap transects. The nature of the light acclimatory response differs quantitatively and qualitatively between these species. An important constraint on light acclimation of the shade-tolerant P. arieianum is its inability to increase photosynthetic nitrogen-use efficiency under conditions of high light availability. The lack of plasticity in chlorophyll a:b ratios does not restrict light acclimation of A_max in P. sancti-felicis. Leaves of P. arieianum exhibited symptoms of chronic photoinhibition in exposed microsites within the large gap. Species differences in the capacity to finely adjust A_max across a wide range of light conditions may be attributed to their maximum growth potential. Light acclimation in species with low maximum growth potential may be constrained at the cellular level by rates of protein and chlorophyll synthesis and at the whole-plant level by low maximum rates of uptake and supply of nutrients and water. For P. arieianum, restriction of photosynthetic plasticity is likely to limit competitive abilities of plants in high-light conditions of large gaps and clearings, whereas observed habitat restrictions for P. sancti-felicis do not appear to depend upon the highly-developed capacity for adjustment of A_max observed in this species.