全世界約有800種榕屬植物 (Ficus, figs) 廣泛分佈於熱帶和亞熱帶地區，其生存於多樣的棲地類型並展現各種生活型態。 榕果 (syconium) 複雜的生活史中包含榕小蜂 (fig wasps) 授粉與食果動物 (frugivores) 協助傳播種子這兩個動−植物互動過程，並有雌雄同株 (monoecious) 和異株 (gynodioecious或functionally dioecious) 的種間生殖差異。 雌雄同株物種往往單一果期短、果量大、且榕果成熟同步，榕果兼顧產生種子與供養小蜂。 異株物種則通常果期長、果量小、且榕果成熟不同步，族群中部分植株負擔雌性功能，產出具有種子的榕果以完成種子傳播，其他植株則扮演雄性角色，產生花粉且生成供養小蜂的癭果。 由於不同生殖系統的結實特性和性別功能差異，相較於雌雄同株，雌雄異株之雌株可能投資較多的繁殖努力在最大化種子量，或者增加動物取食榕果的回饋，像是果肉量、水分、和營養成分等，以利於吸引食果動物並幫助種子傳播。然而，即使在相同的生殖系統中，種間榕果大小變異仍很大，可能因此限制了植株資源的分配，進而影響榕果的性狀。
此外，種內株間可能展現不同的生活史策略，在資源受限的前提下，植株對於繁殖方面的資源投資可能會隨著植株年齡或大小增加而提升。 另一方面，雌雄同株榕果在兼顧種子和榕小蜂的情況下，植株可能必須經由調整果期間的資源分配，使其結實量和榕果性狀產生變異，以適應季節性造成的榕小蜂、食果動物、及物理條件差異。 不過，雌雄同株榕樹植株每年結實的時間皆會有所變動，若能量的獲得與累積能量之時日有關，則可能影響每次果期所能獲得的能量，進而影響榕果的產出。
本研究針對恆春熱帶植物園區內的雌雄同株的大葉雀榕 (F. caulocarpa (Miq.) Miq., large leaf figs) 與雀榕 (F. subpisocarpa Gagnep., fruit fig trees)，及雌雄異株的金氏榕 (F. ampelas Burm. f., King's figs) 與澀葉榕 (F. irisana Elm., rough-leaved figs) 雌株，探討種間和種內之榕果功能性性狀 (functional traits) 差異與其各種生態特性之關聯，並測試以下假說。 假說 (一)：種間生殖系統和榕果大小差異與其榕果的功能性性狀有關。 因生殖系統差異之故，榕果在雌雄功能分配和結實模式方面皆有所不同，進而可能影響植物的繁殖投資，使雌雄異株物種可以投資更多資源在每一顆榕果上，增加榕果的種子量或對食果動物的吸引力，以利種子傳播。 因此我預期 (1) 相較於雌雄同株，雌雄異株雌株榕果會有較高的單位體積種子量和較低的果肉-種子比，或者單位體積種子量和果肉-種子比皆較高。 營養成分的部分，我則預期 (2) 相較於雌雄同株，雌雄異株雌株榕果有較高的水分和動物易消化利用的營養成分，而不易消化之成分含量則較低。
另一方面，榕果大小可能也會限制植物的資源投資，即使在相同的生殖系統中，不同榕果大小之物種會造成性狀產生變異，所以在榕果的形質方面我預期 (3) 相同生殖系統的物種間，較大的榕果有較高的單位體積種子量但果肉-種子比較低。 而水分和營養成分方面我則預期 (4) 相同生殖系統的物種間，較大的榕果有較高的水分及碳水化合物含量，而較小的榕果則有較高的脂質含量。
假說 (二)：種內株間結實量或榕果性狀差異與植株大小之相關特性有關。 根據生活史理論，植株在資源受限的情況下，隨著年齡和植株大小的增長，會逐漸將資源投資在繁殖方面，進而增加其結實量或果實和種子的品質。 於是我預期和植株大小相關之胸高直徑、樹高、或樹冠體積會與其結實量或是榕果大小及種子數量呈正相關。
假說 (三)：雌雄同株種內榕果性狀及結實量變異與天氣、榕小蜂、及食果動物的季節性變化有關聯。 雌雄同株榕果內可用之雌花數量與氣溫呈正向關係，且小蜂的數量和壽命也有季節性差異，進而可能影響了榕果與榕小蜂之間的交互作用，使榕果性狀產生變異。 因此我預期 (1) 相較於乾冷季節，濕熱季節果期的榕果內部種子量和種子-蟲癭比會較高。 此外，環境中的天氣因子和食果動物的季節性需求變化可能也影響了族群或個體內的季節性榕果性狀變異，所以我預期 (2) 相較於乾冷季節，濕熱季節果期的榕果體積較大，水分、果肉乾重、果肉-種子比、碳水化合物、蛋白質、及鈣含量較高，脂質含量則較低。 不過，榕果果肉所含之纖維為動物所不偏好的成分，而灰分則可能一年四季皆為影響動物生理的重要成分，從而使我預期 (3) 榕果果肉纖維和灰分含量沒有季節性差異。 另一方面，根據生活史策略和能量權衡的概念，依照前述的榕果季節性變化趨勢，我預期 (4) 相較於乾冷季節，濕熱季節果期的結實量較低。
最後我測試假說 (四)：雌雄同株每次果期的繁殖表現會被之前累積能量的時間長短影響。 植物產果會消耗大量的資源，因此其能量的獲得和累積會影響每次產果。 在榕屬植物中，雌雄同株榕樹個體每年結實的時間皆會有所變動，可能使每次果期累積能量的時間不同，進而影響其果實或結實量的生產結果。 因此，我預期當果期間隔時間和發育期時間越長時，該次果期的榕果體積會越大或結實量越多。
本研究從2013年3月至2015年2月在恆春熱帶植物園區進行四種榕屬植物的物候調查及榕果採集。 我搜尋園區內步道兩旁及林緣，選取會結果且能夠採集和觀測的植株，作為定期調查和取樣的個體。 本研究最終使用大葉雀榕13棵、雀榕11棵、金氏榕雌株9棵、及澀葉榕雌株8棵，並以每週一次的頻度調查其物候變化和結實量。 我於研究期間收集植株的成熟期榕果，雌雄異株兩物種植株皆僅採收一次果期，而雌雄同株兩物種植株皆盡可能採收至少兩次果期，以比較植株果期間季節性差異，最終獲得大葉雀榕27個果期和雀榕19個果期。 我將採集之榕果測量並記錄其長、寬、高、濕重、乾重、果肉乾重、種子量和總乾重、及蟲癭量，並計算出榕果體積、含水量、單位體積種子量、果肉-種子比、及種子-蟲癭比。 乾燥果肉則進行營養成分分析。
本研究結果顯示四種榕屬植物因生殖系統和榕果大小差異而反應出不同的榕果形質和營養成分性狀，展現出了不同的結實策略。 雌雄同株的大葉雀榕和雀榕在個體間結實不同步，因此幾乎終年皆有植株帶有成熟榕果，結實株數大致上平均分佈在每個月份。 雌雄同株個體內榕果成熟同步且果期短，並且在單位時間內的熟果量很多，但榕果單位體積種子量和水分含量少，不過果肉-種子比高，果肉營養成分含有較高的脂質和纖維。 雀榕的榕果大於大葉雀榕，但植株熟果期結實量較低且榕果單位體積種子量少，不過，榕果含水量和碳水化合物含量較高；而大葉雀榕榕果雖然水分含量低，且不易消化之纖維和脂質含量高，但結實量遠高於其他榕屬植物。
雌雄異株的部分，金氏榕和澀葉榕雌株在個體間結果較為同步，高度聚集在6至8月和10月至12月生產成熟榕果。 雌雄異株雌株個體內榕果成熟不同步且果期長，在單位時間內的熟果量不多，榕果果肉-種子比低，但水分和果肉碳水化合物含量高，榕果單位體積種子量多，使得每顆榕果的種子傳播量較高。 澀葉榕的榕果大於金氏榕，但植株熟果期結實量較低，且榕果單位體積種子量少，但榕果果肉-種子比、含水量、和碳水化合物含量較高；不過，金氏榕的果肉蛋白質和鈣含量是四種榕樹中最高的。
在探討種內植株大小相關之特性與其生產之結實量或果實性狀的關聯中，只有大葉雀榕植株結實量與其胸高直徑有顯著正相關。 而雀榕、金氏榕、和澀葉榕的結實量、榕果大小、及種子數量則與其植株大小相關之特性無顯著關聯，可能是由於其植株不如大葉雀榕，植株可長很大而使其大小差異明顯，故可找出其與結實量之關聯。 此外，雌雄異株也可能是受其可塑性及複雜的物候模式影響。
雌雄同株之大葉雀榕和雀榕榕果性狀部分有季節性變化，且親緣極近之兩物種展現完全不同的結實策略。 大葉雀榕濕季榕果有果肉-種子比較高之趨勢，而乾季產出的榕果則傾向體積較大、果肉乾重較重、及水分和脂質含量較高。 雀榕則為濕季榕果體積較大、水分含量較高、果肉乾重較重、種子-蟲癭比較高、及種子較多，而乾季產出的榕果則有果肉-種子比較高和蟲癭量較多的傾向。 此外，兩者的個體內結實量皆沒有一致的季節性改變。 兩物種種內植株個體變異極大，使不同植株之榕果性狀季節間變動程度和趨勢不一致，顯示出此物種結實模式的高度彈性。
不過，無論在整個族群或個體內，本研究中之大葉雀榕和雀榕榕果在反應性別功能的種子-蟲癭比上皆與榕果大小、種子量、及蟲癭量有直接和間接的相關，說明了雌雄同株與榕小蜂之間的緊密互動關係。 隨著榕果體積變大，種子量和蟲癭量增加，而種子-蟲癭比受種子量影響較大，故也因而隨之提升。
另一方面，本研究觀察到雌雄同株榕樹植株每年結實的時間確實有所變動，並發現大葉雀榕的果期間隔和發育期時間長度顯著影響之後果期的榕果體積。 而雀榕無論是榕果大小或結實量皆與果期間隔和發育期時間長度無關，其榕果大小的差異可能是受個體基因或生理所限制，但仍需更進一步的研究證實。 此外，兩物種之葉子與結實物候無明顯關聯，葉子狀態對繁殖結果的影響則有待更深入之生理研究。
本研究經由探討種間和種內榕果功能性性狀差異與榕屬植物其他特性和生態關聯之間的關係，提供了共域之四種榕屬植物在結實策略上的差異。 此研究結果不僅可說明在演化歷史上較古老之雌雄同株物種的族群生存優勢，還能更進一步探討榕屬植物結實策略對食果動物果實選擇和種子傳播效益之潛在影響。 不僅如此，在面對現今環境變遷影響，本研究也能提供熱帶和亞熱帶地區榕屬植物群聚之生態和演化動態相關的重要資訊。

Nearly half of fig species (Ficus, Moraceae) are monoecious. Their syconia support seed production and wasps rearing simultaneously, and generally emerge in large but shorter and synchronous crops. The rest species reproduce gynodioeciously with some plants as functional females produce only seeded syconia and in generally smaller but longer and asynchronous crops. The latter, compared to monoecious figs, may invest reproductive efforts to maximize seediness, or increase syconium pulp, water content, and nutritional rewards to better attract frugivores, either of which would aid to seed dispersal. Figs, however, vary in syconium size among species even in the same breeding system, thus may constraint their resource allocation. Within population, plants will invest more resource to reproduction with increasing age or tree size, and cause higher crop size or fruit and seed quality. Monoecious trees may also allocate the resource between crops that may encounter seasonal changes of weather, fig wasps, and frugivores, and their syconium size may be regulate by the time to accumulate reserves for crop production. This study conducted phenology survey, measured syconium morphological traits, and analyzed nutrient contents for monoecious F. caulocarpa and F. subpisocarpa and dioecious F. ampelas and F. irisana in Hengchun Tropical Botanical Garden, to examine the relationships of inter- and intra-specific fig functional traits and their ecological correlations. Dioecious syconia appeared to have higher seediness, carbohydrate, and water contents, whereas monoecious syconia tended to have higher pulp-seed ratio, fiber, and lipid contents. Within the same breeding system, large-sized F. irisana and F. subpisocarpa had lower seediness but higher carbohydrate and water contents. Within populations, only F. caulocarpa ‘s plant sizes positively correlated to their crop sizes. Moreover, the two monoecious figs both showed partially seasonally change in syconium traits, and their trends were converse but all good for seed dispersal. Ficus caulocarp’s syconium volumes were also affected by the durations of syconium developmental period and fruiting interval, which positive correlated to energy accumulation. The results generally support inter- and intra-specific differences in fig functional traits are correlated to their fruiting strategies aiding to syconium removal and seed dispersal by frugivores.

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