Diet of Southern Muriquis in Continuous Brazilian Atlantic Forest

We systematically collected data on feeding behavior for one group of 33–39 southern muriquis (Brachyteles arachnoides) in Parque Estadual Carlos Botelho (PECB), São Paulo State, Brazil (37,432.45 ha of continuous Atlantic Forest), between January and December 1995. We determined food item consumption from instantaneous scans of behavior. Fruits were the most eaten food items in all 12 mo (40–80% of scan in every mo, average = 71.3%). Muriquis ate young leaves more than mature leaves or flowers. Our results are consistent with previous findings at the same and neighboring forest sites that southern muriquis have a consistently frugivorous diet when inhabiting less disturbed habitats, but contrast with previous observations on oppportunistic frugivory in muriqui populations inhabiting fragmented forests. Sustained high levels of frugivory probably result from year-round availability of fruit within large continuous forests.     

Feeding ecology and activity pattern of black-fronted titi monkeys (<Emphasis Type="Italic">Callicebus nigrifrons</Emphasis>) in a semideciduous tropical forest of southern Brazil

Most aspects of the ecology and behavior of Callicebus nigrifrons are still unknown. The information available about this species is mainly based on a few studies that also focused on other Callicebus. We examined the feeding behavior and activity pattern of a free-ranging pair of C. nigrifrons between March and November 2007 in an area of semideciduous tropical forest of southeastern Brazil. The study site is located at the southern limit of the Tropical Zone and is characterized by pronounced seasonality. As observed for other Callicebus monkeys, fruits were the most consumed food resource, accounting for 53% of the diet, which was complemented mainly by leaves (16%) but also by invertebrates and flowers (10% of each). A great variety of plant families (28) and species (62) were included in the diet. The titis spent 35% of their time feeding, distributing the remaining time between resting (30%) and traveling (24%). Data presented here indicate that C. nigrifrons prefer high-quality food items (fruit pulp), adding low-quality food items (such as leaves) as the availability of the higher-quality foods decreases. The amount of time spent traveling and resting did not change between seasons, but the time invested in feeding increased during the lean period. The activity pattern was not related to fruit availability, but in months with lower temperatures, monkeys spent more time feeding. We suggest that the feeding ecology and activity pattern of C. nigrifrons reflect adaptations related to annual fluctuations in food availability and temperature, respectively.     

Seasonal variation in activity and diet in a small-bodied folivorous primate,Hapalemur griseus,in southeastern Madagascar

How small-bodied (500–1,200 g) folivorous prosimian primates cope with large amounts of foliage in their diet seasonally has yet to be determined for many species such as Hapalemur griseus, which specializes on bamboo. To address this issue, we present results on seasonal variation in activity and diet from a wild group of H. griseus in southeastern Madagascar. Throughout the study (which was conducted from July–November 1994 and July 1995–February 1996), H. griseus primarily fed on new growth from three species of bamboo: two species of liana bamboo and Cephalostachyum perrieri. Bamboo species were used in different ways seasonally; liana bamboo was consumed more during the dry, cool season, and C. perrieri was eaten more often during the wet, warm season. H. griseus also spent more of their time feeding and traveling than nocturnal folivores of similar body size during the dry season. During the warm wet season, H. griseus decreased the amount of time spent feeding and traveling and rested more often. We hypothesize that seasonal changes in activity may be primarily related to the distribution and availability of food sources and/or reproductive cycles. Am. J. Primatol. 43:211–223, 1997. © 1997 Wiley-Liss, Inc.     

Overwintering strategy of Yunnan snub-nosed monkeys: adjustments in activity scheduling and foraging patterns

Temperate forests are characterized by pronounced climatic and phenological seasonality. Primates inhabiting such environments experience prolonged resource scarcity and low ambient temperatures in winter and are expected to adjust time allocation and foraging behavior so as to maintain their energy balance. We analyzed the activity scheduling of a group of Yunnan snub-nosed monkeys (Rhinopithecus bieti) based on data collected over 20 months in the high-altitude (>3000 m) Samage Forest, Baimaxueshan Nature Reserve, PRC. The forest consists of evergreen conifers and oaks and deciduous broadleaf trees. The diet varied seasonally, with young leaves preferentially exploited in spring and fruits in summer. The monkeys subsisted on readily available fallback resources (mainly lichens) in winter [Grueter et al. in (Am J Phys Anthropol 140:700–715, 2009)]. We predicted that this switch to a relatively low-quality diet would prompt an increase in feeding effort and decrease in moving effort. We found that the monkeys spent significantly more time feeding in winter than in the other seasons. The monthly time devoted to feeding was also negatively correlated with temperature and positively with percentage of lichens in the diet. Time spent on moving did not vary among seasons or with temperature, but day-journey length was found to be longer on hotter days. Time spent resting was lower in winter and under colder conditions and was also negatively correlated with time spent feeding, indicating that resting time is converted into feeding time during times of ecological stress. These results indicate a strong effect of seasonality on time allocation patterns, constraints on inactivity phases, and the prevalence of an energy-conserving foraging strategy in winter, when costs of thermoregulation were high and the availability of preferred food was low.     

Graminivory and Fallback Foods: Annual Diet Profile of Geladas (<Emphasis Type="Italic">Theropithecus gelada</Emphasis>) Living in the Simien Mountains National Park,Ethiopia

All primates show some dietary flexibility, particularly during food shortages. Foods consumed during times of scarcity (i.e., fallback foods) strongly influence the ecology and evolution of a species. Geladas (Theropithecus gelada) eat primarily graminoid leaves (i.e., grasses and sedges), but also consume other diet items (e.g., underground storage organs), especially in the dry season. We investigated the feeding ecology of wild geladas in the Sankaber region of the Simien Mountains National Park, Ethiopia across 12 mo. We asked how the gelada diet in this region, which is disturbed by livestock and agriculture, correlated with food availability and whether underground foods are fallback foods. We quantified the monthly diets of adults from eight reproductive units using instantaneous scan sampling, and seasonal aboveground and underground food availability using point-intercept transects and soil core sampling. Geladas primarily consumed graminoid leaves year-round (76.3% of the annual diet, 36.2–93.2% of the monthly diet) but spent considerable time consuming underground foods in the dry season (14.0% of the annual diet, 11.1–49.7% of the diet across dry season months). Graminoid consumption increased with graminoid availability, and underground food consumption decreased with graminoid availability. Underground food availability did not vary significantly between the dry and wet season sampling months, supporting the hypothesis that underground foods are fallback foods for geladas. We then compiled data from gelada feeding studies and found that underground foods are an important dry season diet item across study sites, but geladas rely more heavily on underground foods in habitats more heavily influenced by humans. Understanding the range and effects of primate dietary flexibility in human-modified habitats will contribute to a better understanding of how changing environments shape primate ecology and evolution.     

Ecological correlates to activity and habitat use of two prosimian primates: Eulemur rubriventer and Eulemur fulvus rufus in Madagascar

In this paper, I provide data on the possible effects of group size and seasonal changes in food availability on the activity and habitat use patterns of two species of prosimian primate: the rufous lemur (Eulemur fulvus rufus) and the red-bellied lemur (Eulemur rubriventer). General and subtle seasonal differences were observed between lemur species in (1) group size and composition, (2) activity profiles, and (3) habitat use. Rufous lemur groups were larger (mean = 8 individuals) and contained more adults than red-bellied lemur groups (mean = 3 individuals). The overall degree and distribution of diurnal activity differed between lemur species and varied within species with seasonal changes in food availability. In general, rufous lemurs traveled more often than red-bellied lemurs. During food scarcity, both species increased the amount of time spent feeding, although peak feeding and traveling times differed. Both species also preferred horizontal substrates; however, during food scarcity, red-bellied lemurs used terminal branches more often while feeding than rufous lemurs. In addition, red-bellied lemurs used the lower middle story forest and rested in taller, upper canopy trees more often than rufous lemurs. Differences in activity and habitat use patterns between lemur species were more related to seasonal changes in food availability than overall differences in group size. These behavioral patterns may represent different strategies used by each lemur species to avoid directly competing for similar resources during times of food scarcity. © 1996 Wiley-Liss, Inc.     

Foraging strategies of black-fronted titi monkeys (<Emphasis Type="Italic">Callicebus nigrifrons</Emphasis>) in relation to food availability in a seasonal tropical forest

Many primates have to cope with a temporal scarcity in food availability that shapes their foraging strategies. Here we investigated the changes in diet, activity, and ranging behavior of a group of black-fronted titi monkeys (Callicebus nigrifrons) according to the availability of the main high-nutritional-density item in their diet and the foraging strategy adopted when this food is scarce. We monitored one habituated group using instantaneous scan sampling over 1 year (533 h of observation, 61 days) in a seasonal tropical forest fragment (245 ha). We simultaneously collected data on food availability with fruit traps. The titi monkeys consumed fleshy fruits, the main high-nutritional-density item of their diet, in accordance with its availability, and the availability of this item modulated the ingestion of vegetative plant parts, a relatively low-nutritional-density food. During high fleshy fruit availability, the titi monkeys consumed more fleshy fruits, flowers, and invertebrates. They also traveled more, but concentrated their activity in a central area of their home range. Conversely, during fleshy fruit scarcity, they increased the breadth of their diet, switching to one richer in seeds and vegetative plant parts, and with greater plant diversity. At the same time, they reduced most energy-demanding activities, traveling less and over shorter distances, but using their home range more broadly. Corroborating the optimal foraging theory, titi monkeys altered foraging strategies according to temporal food fluctuations and responded to low fleshy fruit availability by changing their diet, activity, and ranging behavior. The adoption of a low-cost/low-yield strategy allowed us to classify them as energy minimizers.     

Behavioral responses of Cao Vit gibbon (Nomascus nasutus) to variations in food abundance and temperature in Bangliang, Jingxi, China

The Cao Vit gibbon is a critically endangered species with only about 110 individuals remaining in a degraded karst forest along the China-Vietnam border. Behavioral data from this site are particularly useful in understanding gibbon behavioral adaptations to different sets of ecological conditions and will contribute to the conservation of the species. We studied seasonal variation in the time budget and diet of the Cao Vit gibbon in response to variation in food availability and ambient temperature by observing two groups for 1,379 hr between January and December 2009. We used 5-min scan samples to record the activity of gibbons. Both ambient temperature and food availability varied from month to month. Gibbon groups increased resting time and huddled together in sleeping places in cold months. Gibbons spent more time feeding on fruit when fruit was more abundant suggesting that fruit was their preferred food. Alternatively, leaf eating was negatively correlated with leaf availability which suggested that leaves may be used as a fallback food. Gibbons increased their diet diversity when they ate more leaves. This might be a strategy to cope with toxins or digestion inhibitor accumulation associated with feeding from a limited number of leaf species. Individuals consumed more buds when Broussonetia papyrifera produced buds in March and April. During this period, they decreased traveling time and engaged in less frequent social interactions. Gibbons spent more time searching for and feeding on invertebrates during June and October. However, we did not collect data on invertebrate abundance and therefore cannot determine the relationship between invertebrate feeding and availability. We conclude that flexibility in consuming diverse food types and food species, and in responding to the availability of preferred foods, has enabled the Cao Vit gibbon to survive in a degraded karst forest habitat.     

Activity Patterns, Intergroup Encounters, and Male Affiliation in Free-Ranging Bearded Sakis (Chiropotes sagulatus)

The assessment of how primates divide their daily activities is one of the foundations of primate behavioral ecology but the activity patterns and social behavior of the Pitheciines, including bearded sakis (genus Chiropotes), are poorly understood. During a 15-mo study, I collected 560 h of data on subgrouping, activity patterns, social behavior, and intergroup encounters of a group of free-ranging Guianan bearded sakis (Chiropotes sagulatus) in Guyana. The study group consisted of at least 65 individuals but showed a high degree of flexibility in grouping patterns (mean group size 39 ± 10). They were highly active, spending ca. 70% of their diurnal activity budget traveling and feeding. Activity patterns were relatively consistent throughout the year, although time spent feeding correlated significantly with fruit availability. The most common social behaviors were social resting and grooming. Agonism was rare (2.56% of social behavior) but did occur in the context of intergroup encounters, with males from the same group cooperating in intergroup agonism. Bearded sakis showed a high level of within group male affiliation, with male–male partners making up 65% of grooming dyads and males having another male as their nearest-neighbor 66% of the time. These results show that bearded sakis are characterized by egalitarian male–male and male–female relationships, highly fluid group sizes, and high levels of male affiliation. Similarities in the social behavior of bearded sakis and muriquis suggest several possible explanations for male–male bonding including cooperative defense of females from other groups, kinship, and maintenance of social cohesion after subgroup coalescence.     

Seasonal variation of diet and time budget of Eastern hoolock gibbons (Hoolock leuconedys) living in a northern montane forest

Most gibbons dwell in the tropical forests of Southeastern Asia, but eastern hoolock gibbons (Hoolock leuconedys) survive in high montane forest ranging from 1,600 to 2,700 m a.s.l. in Gaoligongshan (>24°30′N), Yunnan, China. To assess the behavioral adaptations of hoolock gibbons to the montane forest, we related temperature and food availability within the habitat to the seasonal behavioral patterns of a family group and a solitary female between August 2010 and September 2011 in Nankang, Gaoligongshan National Nature Reserve. The maximum temperature was 29.2 °C and the minimum temperature was ?0.3 °C during the period. The monthly mean temperature was <10 °C between December and February, making Nankang the coldest gibbon habitat reported so far. Nonfig fruit and fig availability declined to nearly zero in cold months. The family group increased resting and decreased travel and social behaviors when the monthly mean temperature was low. Compared with other gibbon populations, the hoolock gibbons spent proportionally less time feeding on figs and other fruit than other gibbon populations except Nomascus concolor and Symphalangus syndactylus. Only 36 species of plants provided nonfig fruit or figs, which is less than the number of fruit species consumed by any other gibbon population observed during a similar period of time (about 1 year). Hoolock gibbons shifted their diet to leaves and increased feeding time when fruit was not available. We conclude that diet flexibility and an energy-conserving strategy during the cold season when fruit is scarce have enabled the hoolock gibbons to survive in a northern montane forest.     

贵州麻阳河地区黑叶猴的食性观察

１９９５￣１９９７年对黑叶猴进行野处食性观察,结果表明：（１）对农作物的取食时间是影响其取食植物种类多少的主要因子;（２）其取食部位季节性明显;（３）各季节取食时间长短与了食农作物时间呈负相关;（４）各季节食物交叉性不强。     

Phylogeographic evidence for two species of muriqui (genus Brachyteles)

The taxonomy of muriquis, the largest extant primates in the New World, is controversial. While some specialists argue for a monotypic genus (Brachyteles arachnoides), others favor a two‐species classification, splitting northern muriquis (Brachyteles hypoxanthus) from southern muriquis (B. arachnoides). This uncertainty affects how we study the differences between these highly endangered and charismatic primates, as well as the design of more effective conservation programs. To address this issue, between 2003 and 2017 we collected over 230 muriqui fecal samples across the genus’ distribution in the Brazilian Atlantic Forest, extracted DNA from these samples, and sequenced 423 base pairs of the mitochondrial DNA (mtDNA) control region. Phylogenetic and species delimitation analyses of our sequence dataset robustly support two reciprocally monophyletic groups corresponding to northern and southern muriquis separated by an average 12.7% genetic distance. The phylogeographic break between these lineages seems to be associated with the Paraíba do Sul River and coincides with the transition between the north and south Atlantic Forest biogeographic zones. Published divergence estimates from whole mitochondrial genomes and nuclear loci date the split between northern and southern muriquis to the Early Pleistocene (ca. 2.0 mya), and our new mtDNA dataset places the coalescence time for each of these two clades near the last interglacial (ca. 120–80 kya). Our results, together with both phenotypic and ecological differences, support recognizing northern and southern muriquis as sister species that should be managed as distinct evolutionarily significant units. Given that only a few thousand muriquis remain in nature, it is imperative that conservation strategies are tailored to protect both species from extinction.     

Reproductive Energetics of Female Mantled Howlers (<Emphasis Type="Italic">Alouatta palliata</Emphasis>)

The energetic costs of reproduction have an important influence on the life histories of female primates. At present, however, the interplay of female reproductive state, food availability, and strategies aimed at maintaining energy balance has been described for only a few species, limiting our ability to understand intra- and interspecific variation in female life histories. We assessed how female mantled howlers (Alouatta palliata) are affected by reproductive seasonality, and whether they alter their behavior to cope with the energetic demands of reproduction. From August 2013 to July 2015 we measured the reproductive state, behavior (1100 h of focal animal observations), and energetic condition (312 urine samples collected for C-peptide analysis) of 7 adult females, and assessed food availability (weekly phenological sampling of 397 food trees). Female behavior did not vary with reproductive state or reproductive seasonality. There were, however, differences in how females responded to variation in food availability according to reproductive state. Cycling and gestating females spent more time feeding than lactating females, and cycling females less time resting than females in other reproductive states, when food was more available. C-peptide concentrations were unaffected by either individual or overall variation in reproductive state, except for cycling females, whose concentrations increased during periods of high food availability. The energetic condition of female mantled howlers is broadly maintained over different stages of reproduction, but is sensitive to variation in food availability.     

Leaf chemistry and food selection by common langurs (Presbytis entellus) in Rajaji National Park,Uttar Pradesh,India

Food selection in folivorous primates has been hypothesized to be correlated with leaf chemistry. To test this hypothesis, we conducted a 5-month study on Presbytis entellusin a moist deciduous forest in the Rajaji National Park,Uttar Pradesh, India. The study period covered two seasons, winter and spring. We used the percentage time spent feeding on each food item as an index of food selection, which we estimated from group scan data collected from one study group for 6 days each month. We estimated the selection ratio for each item as the ratio of time spent feeding to availability. We estimated food availability from vegetation sampling and phenological data in the home range of the study group. We estimated crude protein (CP), acid detergent fiber (ADF) and tannins in mature and young leaves of 12 major food species in the laboratory. Food selection is positively correlated with CP in winter and negatively correlated with ADF in both seasons. It also is correlated with CPJADF ratio, but to a lesser extent than the best predictor in winter and spring. Selection ratio is not correlated with CP and ADF in both seasons. It is probable that the inclusion of other factors, such as micronutrients,condensed tannins, and digestibility, might give a better prediction of food selection.     

Activity budgets of woolly spider monkeys,or muriquis (Brachyteles arachnoides)

One group of woolly spider monkeys, or muriquis (Brachyteles arachnoides), was observed from June 1983 through July 1984 at Fazenda Montes Claros, Minas Gerais, Brazil. The study subjects spent an average of 49% of their daylight time resting, 29% traveling, and 19% feeding. They shifted their diurnal schedule of activities in response to seasonal changes in temperature and rainfall. The activity budgets of adult males and females were similar. However, a comparison of three adult females in different reproductive conditions revealed that the lactating female spent a greater proportion of time feeding than did both the pregnant and nonreproductive females. Intraspecific differences in group size and diet appear to be important to understanding differences in the activity budgets of Brachyteles. Differences between locomotion patterns of woolly spider monkeys and sympatric howler monkeys may explain activity budget differences based on group size and diet.     

Feeding ecology and diet of the southern geladas (Theropithecus gelada obscurus) in human‐modified landscape,Wollo, Ethiopia

Studying the dietary flexibility of primates that live in human‐modified environments is crucial for understanding their ecological adaptations as well as developing management and conservation plans. Southern gelada (Theropithecus gelada obscurus) is an endemic little‐known subspecies of gelada that inhabits human‐modified landscapes in the northern central highlands of Ethiopia. During an 18‐month period, we conducted this intensive study in an unprotected area of a human‐modified landscape at Kosheme in Wollo to investigate the feeding ecology of southern geladas and their dietary responses to seasonal variations. We quantified the monthly and seasonal diet data from a band of southern geladas using instantaneous scan sampling method at 15‐min intervals, and green grass phenology and availability using visual inspection from the randomly selected permanent plots. The overall average diet of southern geladas at Kosheme constituted grass blades 55.4%, grass undergrounds 13.2%, grass bulbs 5.6%, grass seeds 5.4%, herb leaves 4.0, fruits 7.3%, and cereal crops 5.6%. Grass blade consumption increased with increasing green grass availability, while underground food consumption increased with decreasing green grass availability, and vice versa. Southern geladas spent significantly more time feeding on the grass blades and herb leaves and significantly less time on bulbs during the wet season than the dry season. Underground grass items (rhizomes and corms) were not consumed during the wet season, but made up 22.3% of the dry season diet. Thus, although grass blades are staple diet items for geladas, underground diet items are important “fallback foods” at Kosheme. Our result shows insights into the dietary flexibility southern geladas adopt to cope with human‐modified landscapes of the north‐central Ethiopian Highlands. Thus, the study contributes to a better understanding of how changing environments shape primate ecology and evolution.     

Seasonal changes in food resource distribution and feeding sites selected by Japanese macaques on Koshima Islet,Japan

Feeding sites of Japanese macaques (Macaca fuscata) change according to seasonal fluctuations in food resource distribution. To examine what characteristics of food items affect feeding site selection, I describe herein the seasonal changes in food items, feeding sites, and food resource distributions of this species. Feeding behavior of monkeys and their food resource distributions were investigated on Koshima Islet, southern Japan, for four study periods (i.e., seasons) in 2002. Monkeys showed large variations in their diet between seasons. To weigh the relative influence of the distribution and abundance of food items on feeding site selection in each season, multiple regression analyses were performed by 100 m × 100 m grid. In the analyses, feeding time was a dependent variable and the abundance of staple food items, for which feeding time was over 5% in each season, in each grid square was an independent variable. There was no correlation between the resource distribution of most food items and the distribution of feeding time by monkeys in each season. Monkeys spent more feeding time where multiple staple food items were available. Food items that affected feeding site selection by monkeys had the following three characteristics: (1) clumped distribution, (2) seasonal availability, and (3) fruit. This suggests that monkeys are likely to select feeding sites to consume food items whose availability is limited temporally and spatially, which may enable them to simultaneously use other widely distributed, abundant food items efficiently.     

Subgrouping Patterns in a Group of Wild <Emphasis Type="Italic">Cebus apella nigritus</Emphasis>

Seasonal patterns of group fragmentation, including the size of subgroups and percentage of time spent in subgroups, may provide information on individual decision-making in response to resource distribution. Age-sex class composition of subgroup membership can offer insights into the social dynamics of the group as a whole. At most field sites, capuchins (Cebus spp.) form stable groups with no evidence of group fragmentation. Here I describe seasonal subgrouping patterns, including proportion of time spent in subgroups, subgroup size, age-sex membership, dyadic fidelity, stability of membership, and the effect of subgrouping on individual foraging efficiency, in a group of wild Cebus apella nigritus. From September 1996 to August 1997 the study group at the Estação Biológica de Caratinga, Brazil divided into 148 different subgroups, on 99 of 194 census days. In contrast to expectations for subgrouping patterns as a response to seasonal distribution of resources, the proportion of days spent in subgroups did not vary significantly by season. Subgroup composition was relatively fluid, with multimale multifemale subgroups the most common throughout the year. Unimale multifemale subgroups were restricted to the wet season; in contrast, all-male subgroups and unimale unifemale subgroups occurred in the dry season. For both males and females, low rank predicted membership in smaller subgroups. For males, but not females, subgrouping coincided with increased foraging efficiency, as measured by increased time spent ingesting food and decreased time spent traveling on days with subgrouping compared to days with the group in a cohesive unit.     

Gonadal growth and reproduction in the commercial sea urchin Paracentrotus lividus (Lamarck, 1816) (Echinodermata: Echinoidea) from southern Spain

The reproductive state of six populations of the commercial sea urchin P. lividus was investigated. Specimens were sampled monthly from six sites of southern Spain. The development of the gonad during the period of study was measured using histological methods and categorised by four stages for females and two for male specimens. A Gonadosomatic Index was also established. The six populations exhibited similar reproductive patterns, having mature gonads during almost all months while reconstitution and spent were present mainly in autumn. The population from Sancti Petri had the highest Gonadosomatic Index value. Higher food availability may explain this difference.     

Phenological traits of the mangrove Rhizophora stylosa Griff. at the northern limit of its biogeographical distribution

The vegetative and reproductive phenology of the subtropical mangrove species Rhizophora stylosa was investigated at Manko Wetland, Okinawa Island, Japan. We assessed phenology using litterfall data over four years. Leaf and stipule litterfall occurred throughout the year, with distinct seasonal patterns. Kendall’s coefficient of concordance, W, revealed that the monthly changes in leaf and stipule litterfall were strongly and significantly concordant among years. Leaf litterfall was significantly correlated with monthly maximum wind speed and monthly day length, and stipule litterfall was significantly correlated with monthly mean air temperature and relative humidity. Branch litterfall showed no clear monthly pattern and correlated well with monthly maximum wind speed. Mean total litterfall was 11.2 Mg ha^?1 year^?1, with the largest component being vegetative organs (78.7 %). Flower, fruit, and propagule litterfall were all highest in summer and lowest in winter. The W values revealed that most reproductive organs in litterfall had significant monthly trends. Flower and fruit litterfall were significantly correlated with monthly day length and monthly mean air temperature, respectively. The average development periods from flower buds to flowers, fruits, and mature propagules were approximately 2–3 months, 4–5 months, and 11–12 months, respectively. Except for branches, all vegetative and reproductive components of litterfall had approximately one year cycles.