• Studies of floral polymorphisms have focused on heterostyly, while stigma-height dimorphism has received considerably less attention. Few studies have examined the reproductive biology of species with stigma-height dimorphism to understand how factors influencing mate availability and pollen transfer are related to morph ratios in populations. • Floral morphological traits, especially herkogamy and reciprocity, pollinator visitation, breeding system and spatiotemporal mate availability, are known to affect intermorph pollination and morph ratios in species with stigma-height dimorphism. In this study, we investigated the presence of stigma-height dimorphism and estimated morph ratios in four naturally occurring populations of Jasminum malabaricum. We quantified morph-and population-specific differences in the abovementioned factors in these populations to understand the observed morph ratios. • The positions of anthers and stigmas were characteristic of stigma-height dimorphism, the first report of this polymorphism in the genus. All study populations were isoplethic, implying equal fitness of both morphs. Herkogamy was higher in the shortstyled morph, while reciprocity was higher between the long-styled stigma and shortstyled anthers. Long-and short-tongued pollinators were common floral visitors, and we observed no differences between morphs in spatiotemporal mate availability or pollinator visitation. Neither morph exhibited self-or heteromorphic incompatibility. • The short-styled stigma had lower reciprocity but likely receives sufficient inter-morph pollen from long-tongued pollinators, and also by avoiding self-pollination due to higher herkogamy. These results highlight the importance of sufficient effective pollinators and floral morphological features, particularly herkogamy, in maintaining isoplethy in species with stigma-height dimorphism.

Inter-morph pollen transfer and its dependence on herkogamy and reciprocity are not completely understood in species with stigma-height dimorphism. We asked whether total stigmatic pollen loads, inter-morph fraction of pollen load and reproductive success differed between morphs in Jasminum malabaricum, a species exhibiting stigmaheight dimorphism. We tested whether higher herkogamy and reciprocity resulted in higher inter-morph pollen deposition and reproductive success.• We quantified individual-level estimates of herkogamy, reciprocity, total stigmatic pollen load, inter-morph stigmatic pollen fraction and fruit set for both morphs in naturally occurring populations of J. malabaricum. • Total pollen load was higher in the long-styled morph, inter-morph pollen fraction was higher in the short-styled morph, but fruit set did not differ between morphs. Higher herkogamy resulted in a higher inter-morph fraction of pollen load and fruit set in the long-styled morph of one population. In the other population, only reciprocity was found to be related to inter-morph pollen deposition.• This study is the first to quantify and report natural inter-morph stigmatic pollen load in a species with stigma-height dimorphism. Morph-specific differences in pollen load were similar to patterns commonly observed in heterostylous species. The results highlight the importance of both herkogamy and reciprocity in facilitating inter-morph pollen transfer. Population-specific patterns indicate that local environmental factors determine the relative functional importance of herkogamy and reciprocity.

In style length polymorphism, morph-specific sequence of encountering male and female sex organs within a flower by pollinators can cause differences in the need to avoid self-pollination and encourage inter-morph pollination. We asked if this difference can lead to disparity in stigma-anther separation (herkogamy) between morphs and spatial match between sex organs of complementary morphs (reciprocity). Further, we tested if herkogamy, and hence the level of selfing, is fairly constant among individuals of a population. Additionally, we examined the relationship between herkogamy and reciprocity among individuals of a population to understand functional interactions between these two morphological traits. Using data on sex organ heights for >200 heterostylous species from the literature, we observed that the short-styled morph had higher herkogamy as compared to the long-styled morph indicating a higher need to avoid selfing. Reciprocity did not show a consistent difference between the upper and lower sex organs implying a strong influence of local ecological factors. In most populations, allometric relationships suggested that herkogamy and hence the level of selfing remains constant. Finally, we observed that herkogamy and reciprocity can be related among individuals of a population, sometimes indicating a potential trade-off between avoidance of self-pollination and facilitation of inter-morph pollination.

The reciprocal position of sexual organs in complementary floral morphs is central to our understanding of heterostyly. Reciprocity indices are used to quantify the spatial match between complementary sex organs, but previous indices fail to appropriately account for intra‐population variation in sex organ positions. In this study, we examine how an increase in intra‐population variation in sex organ heights affects reciprocity and consequently reproductive success. We formulated a reciprocity index that incorporates this variation and asked if estimates of reciprocity can predict reproductive success in naturally occurring heterostylous populations. We developed a reciprocity index that assumed pollen transfer success equalled one for a perfectly matched stigma–anther pair, and decreased to zero with increasing mismatch. We examined the relationship between intra‐population variation in organ position and reciprocity, compared previously proposed indices using simulated populations and empirical data from natural populations, and tested the ability of the indices to predict reproductive success. We observed that when differences between mean complementary sex‐organ heights are small, increasing intra‐population variation in heights resulted in a decrease in reciprocity. However, when this difference is larger, reciprocity increased, reached a peak, and then decreased with increasing variation. Previous indices failed to capture this behavior. Seed set was positively related to reciprocity for our index. These results challenge the current understanding that increasing variation in sex‐organ heights will always decrease reciprocity in heterostylous populations. This may help explain why heterostylous systems exhibit and tolerate high amounts of intra‐population variation in sex organ heights.

AimsThe reciprocal position of sexual organs in complementary floral morphs is central to our understanding of heterostyly. Reciprocity indices are used to quantify the spatial match between complementary sex organs, but previous indices fail to appropriately account for variation in sex organ positions among individuals in a population. The objective of this study was to examine how reciprocity and consequently reproductive success change with an increase in intra-population variation in sex organ heights. To this end, we formulated a reciprocity index that incorporates variation in sex organ positions among individuals in a population and asked if estimates of reciprocity can predict reproductive success in naturally occurring heterostylous populations.MethodsWe developed a reciprocity index that assumed pollen transfer success equalled one for a perfectly matched stigma-anther pair, and decreased to zero with increasing mismatch. Reciprocity was quantified as the average pollen transfer success for all pair-wise combinations of complementary sex organs in the population. We examined the relationship between intra-population variation and reciprocity using simulated populations that varied in the distribution of sex organ positions, and with empirical data from natural populations. We compared previously proposed indices using the simulated and natural populations, and for a subset of natural populations we tested the ability of the indices to predict reproductive success.Important FindingsIn both simulated and natural populations we observed that when differences between mean anther and stigma heights of complementary morphs are small, increasing intra-population variation in heights resulted in a monotonous decrease in reciprocity. However, when differences between mean complementary anther and stigma heights are larger, reciprocity increased, reached a peak, and then decreased with increasing variation. Previous indices failed to capture this behaviour and were largely insensitive to variation or differences in mean complementary sex organ heights. Seed set was consistently positively related to reciprocity for our index, and for two of the four previous indices. These results highlight the importance of incorporating intra-population variation in sex organ dimensions in quantifying reciprocity, and challenge the current understanding that increasing variation will always decrease reciprocity in heterostylous populations. These results may help explain why heterostylous systems exhibit, and tolerate high amounts of intra-population variation in sex organ heights.