Philippa Kaur

The data used here provides evidence that particular relationships may determine sex-specific patterns of witchcraft accusation. Cases where women were targeted frequently came from affinal kin, while those directed at men were often from unrelated individuals and blood relatives. Most previous research on factors that determine the sex of accused ‘witches’ has largely consisted of qualitative studies of a single society or a few societies48, or historical studies that have not tested for correlations49. Our findings, in support of the overarching hypothesis that accusations may be driven by various forms of competition, can be tentatively aligned with evolutionary literature on patterns of intrasexual and kin competition, intersexual conflict and polygamous mating30,31,50.Men were more often accused than women in our sample, although we did not have a prediction in relation to this. But the finding suggests how overall patterns of competition within relationships may contribute to societal ‘phenotypes’ of witches as male or female. The ethnography of the Ndembele perhaps indicates why women were less frequently targeted in Bantu societies: ‘in a case of witchcraft, the complainant is actuated by caprice, jealousy or pique; and the defendant is a person of wealth or popularity, and is always a man, for the women have neither wealth nor honor worth coveting’51.Our predictions about how the sex of accused ‘witches’ might be associated with particular relationship categories were supported. The majority of accusations targeting men came from unrelated individuals, which is unsurprising, as inclusive fitness52 would not mitigate the effects of competition between them. Blood relatives were the next most common relationship category directing accusations at men. This aligns with more recent studies indicating that witchcraft fears between family members are significant in parts of Africa, to the extent that they can be construed as ‘the dark side of kinship’53. In evolutionary terms, kin may compete with one another in environments where resources are limited30,31,50 and in societies with patrilineal inheritance related males, and particularly brothers, compete for resources in order to marry31. This aligns with an ethnographic observation that among the Banyoro witchcraft accusations often occurred between brothers over inheritance, but not between brothers and sisters, whose interests did not conflict21. The situations relating to accusations of men were also often connected to the acquisition of wealth and status, such as rivalry over village headmanships32, power struggles between a chief’s counsellors54 or disputes over inheritance55. These connections can be found in more recent contexts such as twentieth century Ghana, where notions of obtaining political power and wealth through occult means involving human sacrifice were pervasive56.Accusations of women were more likely to come from affines. Husbands were the largest category of affinal kin to accuse women (Supplementary Fig. 2). The higher rate of accusations from husbands to wives than wives to husbands aligns with evolutionary perspectives suggesting male coercion of females is a strategy to maximize male reproductive success39,41. Accusations of wives who were suspected of being unfaithful can be interpreted as a strategy for reducing investment in unrelated offspring35,41. In a case from the Shona a woman gave birth to a stillborn child. This was attributed to an affair before marriage, and was followed by divorce and the repayment of bridewealth to her husband, who commented she was ‘a witch, a woman who had killed her own child’48. Other ethnographic accounts suggest accusations of wives by husbands were an attempt to gain control within the marital relationship55.A significant number of accusations of women by affinal kin were from co-wives in polygynous marriages, and these were often notably associated with jealousy connected to a husband’s attention and investment32. Evolutionary models predict competition for reproductive resources would occur among co-resident breeding women57, as has been found to occur among the Mosuo of southwest China58. In the patrilocal social systems that are predominant in our sample, women disperse at marriage and are isolated from kin, so conflict may be more extreme30. This is consistent with ethnographic observations reporting that the relationship between co-wives in polygynous marriages was often (although not always) marked by conflict, and liable to produce witchcraft accusations38,59.There were accusations of women from other categories of their affinal kin (Supplementary Fig. 2). These again may result from competition for a husband’s time and resources between his kin and wife. New wives may be vulnerable in environments where they enter their husband’s families as unrelated strangers, and are potentially expendable, at least before the arrival of offspring. Some accounts of accusations indicate that accusations of wives by in-laws in patrilocal households are common29.Accusations directed at elderly individuals targeted women more often than men. This may form part of a broader pattern of geronticide: societies close to subsistence-level are documented as sometimes accepting the abandonment or killing of elderly people19,60. In modern Tanzania, ‘witches’ are mostly post-reproductive women, who are more likely to be murdered in periods of income shock19. This is also the case in contemporary Ghana, where accusations are frequently directed at middle-aged or elderly women, whose families may subsequently cease to provide them with financial or material assistance61. In our sample, elderly women may have been targeted more frequently as a result of longer female lifespans: in a polygynous society, men may marry younger women, so wives would be widowed at an earlier age than husbands. Among the Bantu, older men were accused, but some were possibly protected by their status.Accusers’ payoffs from accusations are not always explicit but they can be inferred. The most common outcome of accusations in our sample was that accused ‘witches’ were exiled from their communities or forced to move from where they were living. This would mean resources and cooperative assistance they would have used became available to their accusers or others nearby. Where the accused acquires a negative reputation, which was the second most common outcome, there may be a subtle removal of benefits, which may be preferred to direct ‘punishment’ as it is less costly62. Accusers’ gains need not be direct, as harming behaviours may reduce the overall pressure of competition in an environment28. 8% of accusations in the sample resulted in the acquisition of either resources or political positions from the accused, or in preventing the accused from acquiring them. Where the accused were penalised in other ways, such performing ceremonies to reconcile with accusers, this is perhaps akin to classic cooperation models involving the punishment of defectors (although the accused may not actually be uncooperative)11, providing accusers with subordinate partners who offer fitness benefits to avoid more serious allegations63. Where an accusation does not ‘stick’, ethnographic accounts sometimes indicate it was reversed through divination or ordeal54. In other cases, for various reasons accusations are short-lived and forgotten about4. Finally, although not tested in this dataset, accusers may gain informal prestige and dominance, an outcome analogous to competitive punishment63.Not all of the cases in our dataset support the hypothesis that witchcraft accusations are a mechanism for competition. There is a significant proportion where the accusation of a particular individual appears to be incidental, or dependent a on circumstantial association between the ‘witch’ and a negative event. Such accusations are unlikely to provide accusers with a competitive advantage. There are several possible explanations for such cases. They are in line with the hypothesis that witchcraft belief arises from attempts to identify the cause of an impactful misfortune3,4. Cultural evolutionary explanations of witchcraft beliefs suggest that they are a maladaptive attempt to explain misfortune. Although it is inaccurate, belief in witches is maintained through bias and selective inattention to evidence that would otherwise counter it64. Alternatively this could be viewed under the contention that superstitious beliefs (or errors in attributing cause and effect) are broadly adaptive if they occasionally lead individuals to acts which provide them with fitness benefits65.Although witchcraft accusations may be a mechanism for mitigating the damage to accusers’ reputations in harmful competitive acts, as with any behavioural strategy it is not without risks. Accusers may suffer costs in the form subsequent reputational damage or counter-accusations, as with punishment63, depending on factors such the level of support for an accusation by other members of the community.One limitation of our dataset is that it contains realized allegations of witchcraft, that cannot be tested against baseline population measures. We could not examine the risk that a particular individual, such as an elderly woman, would be accused. Instead, the analysis shows the odds, given an accusation occurred, that the ‘witch’ was male or female, given certain predictors. For example, if the accused was elderly, there are increased odds they were female rather than male.A dataset using historic witchcraft cases is almost certainly affected by selection bias. Cases with sensational outcomes are more likely to be reported, and cases that are dismissed or where the accused removes themselves from their accusers are liable to be overlooked19. Most incidents in our sample were reported anecdotally. Obtaining a random sample of witchcraft accusations within a population is challenging, if not impossible1,66. Attempts to systematically collect cases within a given location and timeframe cannot guarantee that all are brought to the attention of researchers19. Comparative studies of this kind usually use all the data that is available and control for confounding effects. Our sensitivity analyses suggest the large number of accusations of men in the dataset probably reflects patterns of accusations in these societies, rather than male-focused bias from ethnographers. There are many accounts of cultures where witches are predominantly male33,34,49. But the accuracy of historic ethnographic accounts cannot be verified, especially in relation to one-off events such as witchcraft accusations, just as it is unclear how much uncertainty there is in the ethnographic record overall67. Ethnographers may not always have noted the characteristics of the individuals involved, or there may be times where they were mistaken in reporting the circumstances surrounding an accusation. There are several explanations for cases where the identities of accusers or purported victims of witchcraft were not reported. Not all cases had identifiable ‘victims’, for example when the accused was thought to have used witchcraft to promote their own success, or ethnographers could not denote the relationship between the accused and their accusers when suspicions of witchcraft were communicated through general gossip. In a small number of cases, ethnographer perspectives on accusations (and possible inability to access further information) are salient, as they may ascribe more importance to one relationship over another in reporting a case, such as a witch’s envy of their victim, or a witch’s argument with an accuser.However, it is likely that ethnographers were for the most part accurate in documenting variables of interest such as the sex of an accused individual and their relationships with accusers. There is less certainty in relation to the situation connected to an accusation, especially taking into recent research that indicates the prevalence of phenomena such as the misperception of causation68,69. Our attempts to account for such possibilities with sensitivity analyses and meta-data on the production of ethnographies cannot conclusively provide reassurance that bias has not affected results, and so this section of the analysis should be treated with caution and regarded as exploratory. The situations documented in our study do however align with accounts of accusations from more contemporary observers and studies from different geographic locations, suggesting that similar causes of accusations arise convergently in different societies. For example in modern contexts accusations have led to accusers gaining land or property in India6 and cessation of the obligation to provide material and financial assistance to elderly relatives in Ghana61. One advantage of our cross-cultural data being drawn from numerous ethnographies is that it is not reliant on the perspective of one individual, meaning that random perceptual error or individual (as opposed to cultural) bias is more likely to be mitigated in the results than would be the case in the study of a single culture by one ethnographer.As a further limitation, we were reliant on accessible ethnographic records from the best-documented societies. Although selection bias in favour of better described societies is present in our sample, this should not impact the main aim of this research, which is to understand the determinants of witchcraft accusations being directed at male or female targets.Overall our findings may indicate allegations of witchcraft stem from diverse forms of competition between individuals. This aligns with evolutionary approaches to competition and conflict. Accusations may provide fitness benefits by allowing individuals to target competitors, but the exact form and direction of competition is determined by aspects of socio-ecology. This in turn influences which sex is most likely to be accused and the overall portrayal of witches in a society. Accusations may be more likely to occur in some relationships rather than others, when there is a gain for the accuser, as in disputes over inheritance and property, or where another individual may pose a threat, or by simply reducing numbers of competitors. The success of witchcraft accusations in removing competitors and their flexibility as an adaptive strategy may explain their widespread distribution. More

In the version of this article initially published, there were mistakes in affiliations 1, 2 and 6. The corrected affiliations should read as follows: 1. Department of Earth System Science, Ministry of Education Key Laboratory for Earth System Modeling, Institute for Global Change Studies, Tsinghua University, Beijing, China; 2. Ministry of Education Ecological Field Station for East Asian Migratory Birds, Department of Earth System Science, Tsinghua University, Beijing, China; 6. Department of Geography, Department of Earth Sciences, and Institute for Climate and Carbon Neutrality, The University of Hong Kong, Hong Kong, China. The affiliations have been corrected in the HTML and PDF versions of the article. More

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Data collection of flowering and fruiting phenologyMonthly reproductive phenology data recorded over 35 years (from April 1976 to September 2010) were collected from the Bulletin Fenologi Biji Benih dan Anak Benih (Bulletin of Seed and Seedling Phenology), which was deposited at the FRIM library. The bulletin reported seed and seedling availabilities and the flowering and fruiting phenology of trees at several research stations in Malaysia. The present study collected flowering and fruiting records of trees grown in FRIM arboretums located approximately 12 km northwest of Kuala Lumpur, Malaysia (latitude 3°24 ‘N, longitude 101°63 ‘E, elevation 80 m). There are both dipterocarp and non-dipterocarp arboretums in FRIM, both of which were founded in 1929. These arboretums preserve and maintain living trees for research and other purposes. Each month, three research staff members of FRIM with sufficient phenology monitoring training made observations with binoculars to record the presence of flowers and fruits on trees of each species on the forest floor from April 1976 to September 2010. The phenological status of the trees was recorded as flowering during the developmental stages from flower budding to blooming and as fruiting during the developmental stages from the occurrence of immature fruit to fruit ripening. Because only one or two individuals per species are grown at the FRIM arboretums, the flowering and fruiting phenology were monitored using these individuals. The resultant flowering and fruiting phenology data included a time series of binary data (1 for presence and 0 for absence) with a length of 417 months.The original data included 112 dipterocarp and 240 non-dipterocarp species. We excluded 17 dipterocarps and 125 non-dipterocarp species based on the following five criteria for data accuracy.

1.

Percentage of missing values is ≤50%: If the monthly flowering or fruiting phenology data of a species included a substantially large number of missing values ( >50%), the species was excluded.

2.

Stable flowering period: We considered an observation to be unreliable if the flowering period was significantly different among flowering events (if the coefficient of variation in the flowering period was larger or equal to 1.0).

3.

Flowering period is shorter than or equal to 12 months: we considered an observation to be unreliable if the flowering period was longer than 12 months because it was unlikely that the same tree would flower continuously for longer than 1 year.

4.

The flowering and fruiting frequencies were not significantly different between the first and second half of the census period: when the flowering frequency was zero for the first half of the observation period but was larger than 0.1 for the second half of the observation period, or when the flowering frequency was zero for the second half of the observation period but was larger than 0.1 for the first half of the observation period, we removed these species because data are not reliable (e.g., physiological conditions may have changed significantly). We adopted the same criteria for the fruiting phenology data.

5.

We removed overlapping species, herb species, and specimens with unknown species names.

After removing unreliable species based on the five criteria explained above, we obtained 95 dipterocarp and 115 non-dipterocarp species (Supplementary Data 1). We used these species for further analyses. It is unlikely that our final data includes trees that were replaced by young trees during the census period because newly planted seedlings do not flower over 20–30 years until they are fully grown to the reproductive stage ( >20–30 cm DBH)45.Detection of seasonality in reproductive phenologyTo compare the flowering and fruiting phenology seasonality among different families, nine families that included at least five species were used. The number of flowering or fruiting events was counted for each month from January to December during a census, and then the frequency distribution was drawn as a histogram. Similarly, we also generated a histogram for the seed dispersal month, which was calculated as the month when fruiting ended (i.e., when the binary fruiting phenology data changed from one to zero).Classification of phenological patternsTo classify the phenological patterns, we performed time-series clustering using the R package TSclust46 with the hierarchical clustering method based on the Dynamic Time Warping distance of the flowering phenology data of each species. For this analysis, time points at which there were missing values for at least one species were excluded. Because of the large number of missing values in non-Dipterocarpaceae species, we performed time-series clustering only for the Dipterocarpaceae species based on 394 time points in total. The number of phenological clusters was estimated based on AIC, as explained below.Climate dataDaily minimum, mean, and maximum temperatures and precipitation data monitored at the FRIM KEPONG (3° 14’ N, 101° 42’ E, elevation 97 m) weather station were provided by the Malaysian Meteorological Department. We used the daily minimum temperature for our analysis because there were fewer missing values compared to the numbers of missing daily mean and daily maximum temperature values. The periods in which climate data were available were from 1 March 1973 to 31 March 1996, and from 23 July 1997 to 20 April 2005. We removed periods in which there were missing values spanning longer than 5 days. When the range of missing values spanned a period shorter than 3 days, we approximated these missing values using the mean minimum temperatures recorded on the adjacent three days. Although solar radiation data were not available for our study, the use of precipitation is sufficient for model fitting because there is a significant negative correlation between solar radiation and precipitation in Southeast Asia47.Climate data generated by GCMsAs the future climate inputs, we used bias-corrected climate input data from 1 January 2050 to 31 December 2099, with a daily temporal resolution and a 0.5° spatial resolution, provided by the ISI-MIP project48; these data are based on the Coupled Model Intercomparison Project Phase 5 outputs from three GCMs: GFDL–ESM2M, IPSL–CM5A-LR, and MIROC5. To compare the flowering phenology between 1976–1996 and 2050–2099, bias-corrected GCM data from 1 May 1976 to 31 March 1996, were also used. This period (1 May 1976–31 March 1996) is consistent with the period used for model fitting. We selected daily minimum temperature and precipitation time series from the 0.5° grid cells corresponding to the study site for phenology monitoring at FRIM. To compare flowering phenology among regions, we also used the same set of data from three other regions in Southeast Asia: Trang Province in Thailand (7° 4’ N, 99° 47’ E), Lambir Hills National Park in Malaysia (4° 2’ N, 113° 50’ E), and central Kalimantan in Indonesia (0° 06’ S, 114° 0’ E). Because the study site in FRIM was not in the center of a 0.5° grid cell, we interpolated the data using four grid cells in the vicinity of the observation site. We used the weighted average according to the distance between each observation site and the center of each corresponding grid cell.Although the climate input data provided by ISI-MIP were already bias-corrected, we conducted additional bias correction at FRIM using a historical scenario for each GCM data set and the observed weather data from 1 January 1976 to 31 December 2004 based on previously presented protocol49. We did not implement any bias correction for the frequency of dry days or precipitation intensity of wet days49 because we only focused on the average precipitation.The variances in the annual fluctuation of the monthly mean precipitation were not the same between the observation data and historical GCM runs at FRIM. For all three GCMs (GFDL–ESM2M, IPSL–CM5A-LR, and MIROC5), the variances in the yearly fluctuation output by the GCMs tended to be larger than that of the observed data at the FRIM KEPONG weather station during winter and spring. On the other hand, during summer and fall, the variances output by the GCMs tended to be smaller than that of the observed data. These biases could not be corrected using the previous method49. Therefore, we conducted the following bias correction for these data:$${p}_{i,m,y}^{{{{{{rm{GCM}}}}}}* }={r}_{i,m,y}^{{{{{{rm{GCM}}}}}}}cdot left[{F}_{Gamma }^{-1}left({F}_{Gamma }left({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}|{k}_{m,y},{theta }_{m,y}right)|{k}_{m,y}^{* },{theta }_{m,y}^{* }right)cdot {rho }_{m,y}^{{{{{{rm{GCM}}}}}}}right],$$
(1)
where ({p}_{i,m,y}^{{{{{{rm{GCM}}}}}}* }) is the bias-corrected precipitation value of the target GCM at year y, month m, and date i. In the equation, ({r}_{i,m,y}^{{{{{{rm{GCM}}}}}}}) is the ratio of the precipitation value of the GCM relative to the monthly mean value. Then, the following equation is used:$${r}_{i,m,y}^{{{{{{rm{GCM}}}}}}}=frac{{p}_{i,m,y}^{{{{{{rm{GCM}}}}}}}}{{bar{p}}_{m,y}^{{{{{{rm{GCM}}}}}}}},$$
(2)
where ({p}_{i,m,y}^{{{{{{rm{GCM}}}}}}}) is the precipitation value (not bias-corrected) of the GCM at year (y), month (m), and date i and ({bar{p}}_{m,y}^{{{{{{rm{GCM}}}}}}}) is the monthly mean precipitation value of the GCM at year (y) and month (m). In Eq. 1, ({F}_{Gamma }) represents the cumulative distribution function of a gamma distribution, ({F}_{Gamma }^{-1}) represents the inverse function of the cumulative distribution function of the gamma distribution, and ({k}_{m,y}) and ({theta }_{m,y}) are the shape parameters. In Eq. 1, ({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}) indicates the deviation of the monthly mean from the normal climate value of the corresponding period, and this value is calculated as follows:$${delta }_{m,y}^{{{{{{rm{GCM}}}}}}}=frac{{bar{p}}_{m,y}^{{{{{{rm{GCM}}}}}}}}{{rho }_{m,y}^{{{{{{rm{GCM}}}}}}}},$$
(3)
where ({rho }_{m,y}^{{{{{{rm{GCM}}}}}}}) is the normal climate value during the target period. In this method, we defined the normal climate value as the mean of the monthly mean precipitation values over 31 years.$${rho }_{m,y}^{{{{{{rm{GCM}}}}}}}=frac{1}{31}mathop{sum }limits_{j=y-15}^{y+15}{bar{p}}_{m,j}^{{{{{{rm{GCM}}}}}}}.$$
(4)
When the mean of a gamma distribution is fixed at one, the shape parameters are represented as follows:$${k}_{m,y}=frac{1}{Vleft({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}right)},$$
(5)
$${theta }_{m,y}=frac{1}{{k}_{m,y}},$$
(6)
where (Vleft({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}right)) indicates the variance in ({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}) at month (m) over 31 years.In this method, we assumed that the ({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}) value follows a gamma distribution and that the ratio of the variance of ({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}) to the variance of ({delta }_{m,y}^{{{{{{rm{obs}}}}}}}) is maintained even in the future scenario. Here, ({delta }_{m,y}^{{{{{{rm{obs}}}}}}}) represents the deviation of the monthly mean in the observation data from the normal climate value.$${delta }_{m,y}^{{{{{{rm{obs}}}}}}}=frac{{bar{p}}_{m,y}^{{{{{{rm{obs}}}}}}}}{{rho }_{m}^{{{{{{rm{obs}}}}}}}},$$
(7)
$${rho }_{m}^{{{{{{rm{obs}}}}}}}=frac{1}{28}mathop{sum }limits_{j=1976}^{2004}{bar{p}}_{m,y}^{{{{{{rm{obs}}}}}}}.$$
(8)
In the above equations, ({bar{p}}_{m,y}^{{{{{{rm{obs}}}}}}}) indicates the monthly mean precipitation value in the observed data. As mentioned above, because we assume that the ratio of the variance in ({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}) to the variance in ({delta }_{m,y}^{{{{{{rm{obs}}}}}}}) is maintained, ({k}_{m,y}^{* }) and ({theta }_{m,y}^{* }) are calculated as follows:$${k}_{m,y}^{* }=frac{{k}_{m,y}}{alpha },$$
(9)
$${theta }_{m,y}^{* }=frac{1}{{k}_{m,y}^{* }},$$
(10)
where$$alpha =frac{Vleft({delta }_{m,y}^{{{{{{{rm{GCM}}}}}}}^{{{{{{rm{h}}}}}}}}right)}{Vleft({delta }_{m,y}^{{{{{{rm{obs}}}}}}}right)}.$$
(11)
In Eq. 11, ({delta }_{m,y}^{{{{{{{rm{GCM}}}}}}}^{{{{{{rm{h}}}}}}}}) is the deviation of the monthly mean of the historical GCM precipitation data from the normal climate value. Here, we defined the normal climate value as the average monthly mean during 1976–2004.The method proposed here is an original bias correction method, but the above equations are easily derived if we assume that the ({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}) value follows a gamma distribution and that the ratio of the variance in ({delta }_{m,y}^{{{{{{rm{GCM}}}}}}}) to the variance in ({delta }_{m,y}^{{{{{{rm{obs}}}}}}}) is maintained even in the future scenario. Notably, because we combined this method with the bias correction method described previously49, Eq. 2 should be expressed as follows:$${r}_{i,m,y}^{{{{{{rm{GCM}}}}}}}=frac{{widetilde{p}}_{l,m,y}^{{{{{{rm{GCM}}}}}}}}{{bar{p}}_{m,y}^{{{{{{rm{GCM}}}}}}}},$$
(12)
where ({widetilde{p}}_{l,m,y}^{{{{{{rm{GCM}}}}}}}) is the precipitation data that are bias-corrected using the method described previously49. Bias-corrected data were compared with the data without bias correction (Supplementary Figs. 8–11).Statistical analyses and reproducibilityWe adopted previously presented models in which environmental triggers for floral induction accumulate for n1 days prior to the onset of floral induction21 (Supplementary Fig. 2). Flowers then develop for n2 days before opening (Supplementary Fig. 2). The model assumption of the time lag between floral induction and anthesis, which is denoted as n2, was validated by a previous finding in which the expression peaks of flowering-time genes, which are used as molecular markers of floral induction, were shown to occur at least one month before anthesis in Shorea curtisii19. S. curtissi is included in our data set. The CU at time t, ({{{{{rm{CU}}}}}}left(t|{theta }^{C}right)), is calculated as follows:$${{{{{rm{CU}}}}}}left(t|{theta }^{C}right)=mathop{sum }limits_{n={n}_{2}}^{{n}_{2}+{n}_{1}-1}{{{{{rm{max }}}}}}{bar{C}-xleft(t-nright),0},$$
(13)
where ({theta }^{C}=left{{n}_{1},{n}_{2},bar{C}right}) is the set of parameters and x(t) is the temperature at time t. Here, (bar{C}) indicates the threshold temperature. The term max{x1, x2} is a function that returns a larger value for the two arguments. Similarly, given ({theta }^{D}={{n}_{1},{n}_{2},bar{D}},) the DU at time t, ({{{{{rm{DU}}}}}}left(t|{theta }^{D}right)), is defined as the difference between the mean daily accumulation of rainfall over n1 days and a threshold rainfall level ((bar{D})):$${{{{{rm{DU}}}}}}left(t|{theta }^{D}right)={{{{{rm{max }}}}}}left{bar{D}-mathop{sum }limits_{n={n}_{2}}^{{n}_{2}+{n}_{1}-1}yleft(t-nright)/{n}_{1},0right},$$
(14)
where y(t) is the rainfall value at time t. The term max{x1, x2} is defined similarly as in Eq. 13.Logistic regression was performed using only the DU and using the product of CU and DU (CU × DU) as the explanatory variables and using the presence or absence of a first flowering event as the dependent variable for each phenological cluster. Because the number of phenological clusters is unknown, we performed forward selection on the cluster number based on the AIC. Let m be the number of phenological clusters based on the dendrogram drawn from the time-series clustering explained above (Supplementary Fig. 5). Given m phenological clusters, let ({G}_{k}^{m}) be the kth set of clusters in which the DU model is adopted for model fitting. Here, ({G}_{k}^{m}) indicates the set of cluster IDs, and k ranges from 0 to m(m+1)/2. For example, when m = 2 (i.e., there are two clusters, clusters 1 and 2), there are four cluster sets, calculated as follows:$${G}_{0}^{m=2}={},{G}_{1}^{m=2}={1},{G}_{2}^{m=2}={2},{G}_{3}^{m=2}={1,2},$$
(15)
where the element in the bracket indicates the ID of the cluster in which the DU model is adopted for model fitting. When k = 0, the DU model is not used; instead, the CU × DU model is adopted for model fitting for both clusters 1 and 2. Let i be the ith element of the vector E, which is defined as follows:$${{{{{bf{E}}}}}}={{t}_{1}^{1},,{t}_{2}^{1},…,,,{t}_{n}^{1},,…,,,{t}_{1}^{m},,{t}_{2}^{m},…,,{t}_{n}^{m}},$$
(16)
where n is the length of the time-series data for each cluster. Notably, n = 223 is the same for all species and clusters. The term ({t}_{1}^{m}) in the above equation denotes the first time point of the time series of length n for the species included in cluster m. Given m and k, let ({p}^{(m,k)}(i)) be the flowering probability of element i of vector E. The term ({p}^{(m,k)}(i)) is expressed as follows:$${{log }}left[frac{{p}^{left(m,kright)}left(iright)}{1-{p}^{left(m,kright)}left(iright)}right]= mathop{sum }limits_{j=1}^{m}{alpha }_{m,j}cdot {Z}_{m,j}left(iright)+mathop{sum }limits_{jin {G}_{k}^{left(mright)}}^{m}{beta }_{m,j}cdot {Z}_{m,j}left(iright)cdot {{{{{{rm{DU}}}}}}}_{m,j}left(i|{theta }_{j}^{D}right)\ +mathop{sum }limits_{jnotin {G}_{k}^{left(mright)}}^{m}{beta }_{m,j}cdot {Z}_{m,j}left(iright)cdot {{{{{rm{CU}}}}}}left(i|{theta }_{j}^{C}right)times {{{{{{rm{DU}}}}}}}_{m,j}left(i|{theta }_{j}^{D}right),$$
(17)
where ({Z}_{m,j}(i)) is the dummy variable indicating a cluster for i; ({Z}_{m,j}(i)) equals 1 if the ith element of E belongs to the jth cluster, otherwise it is zero, and ({alpha }_{m,j}) and ({beta }_{m,j}) in Eq. (5) are regression coefficients for the jth cluster when the species are grouped into m clusters. We estimate the parameters and the number of clusters based on a finite number of observations. Given the number of clusters m, for each of m clusters, the parameters were estimated by maximizing the loglikelihood value calculated for all combinations of potential parameter values for ({n}_{1},{n}_{2},bar{C},) and (bar{D}) within the ranges of [1 (min), 50 (max)] for n1, [1,50] for n2, [19,25] for (bar{C}), and [1,9] for (bar{D}). We varied the days (n1 and n2) by integers, temperature ((bar{C})) by tenths of a degree C, and daily precipitation ((bar{D})) by tenths of a mm. Regression coefficients (({alpha }_{m,j}), ({beta }_{m,j})) for all j values under a given m value and associated likelihoods were determined using generalized linear models with binomial error structures.With the results of the parameter estimations, we determined the number of clusters in two steps. For the first step, for a given m, we obtained (hat{k}(m)) according to the following equation:$$hat{k}(m)={arg }mathop{{min }}limits_{k}{{{{{{rm{AIC}}}}}}{m,k(m)},,k(m),=,0,,…,{2}^{m}}.$$
(18)
For the second step, with the results of (hat{k}) obtained from the first step, we obtained the estimate of the number of clusters according to forward selection by searching for the (hat{m}) value that satisfies the following inequalities:$${{{{{rm{AIC}}}}}}(hat{m},,hat{k}(hat{m})), < ,{{{{{rm{AIC}}}}}}(hat{m}+1,,hat{k}(hat{m}+1))cap {{{{{rm{AIC}}}}}}(hat{m},,hat{k}(hat{m})), < ,{{{{{rm{AIC}}}}}}(hat{m}-1,,hat{k}(hat{m}-1)).$$ (19) For model fitting, the first flowering month was extracted from the flowering phenology data. When flowering lasted more than 1 month, the month after the first flowering month was replaced by a value of zero (absence of flowering). If the month before the first flowering month was a missing value, the first flowering month was treated as a missing value and was not used for further analyses. We assumed that phenology monitoring was performed on the first date of each month.Projections of 21st-century changes in flowering phenologyWe used two scenarios (RCP2.6 and RCP8.5) to forecast future reproductive phenology in dipterocarp species for each of the three GCMs (GFDL–ESM2M, IPSL–CM5A-LR, and MIROC5). We predicted the flowering probability per month for each phenological cluster during the periods from 1 May 1976–31 March 1996 and from 1 January 2050–31 December 2099 based on the best model (Supplementary Table 2). The predicted flowering probability during the 2050–2099 period was normalized to that during the 1976–1996 period for each climate scenario and for each of three GCMs. To compare the seasonal patterns between 1976–1996 and 2050–2099, the predicted flowering probability was averaged for each month from January to December and plotted for each month in Fig. 6. R version 3.6.3 was used for all analyses.Reporting summaryFurther information on research design is available in the Nature Research Reporting Summary linked to this article. More

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