Fruit traits reflect adaptation to dispersers along a tropical elevational gradient

Fruit traits reflect adaptation to dispersers along a tropical elevational gradient | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL Ecology and Evolution This is a preprint and has not been peer reviewed. Data may be preliminary. 4 September 2025 V1 Latest version Share on Fruit traits reflect adaptation to dispersers along a tropical elevational gradient Authors : Richard Hazell 0000-0001-6347-3209 [email protected] , Graham Kaina 0000-0002-3806-8698 , Katerina Sam 0000-0002-3436-0579 , Daniel Souto-Vilarós , Wulan Koagouw , Vojtech Novotny 0000-0001-7918-8023 , Mika Peck , and Alan Stewart Authors Info & Affiliations https://doi.org/10.22541/au.175697755.52347980/v1 Published Ecology and Evolution Version of record Peer review timeline 425 views 258 downloads Contents Abstract Supplementary Material Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Fruit traits have the potential to influence disperser communities and vice versa. Here we examine the trends in three fruit traits related to dispersal by frugivores: diameter, colour and presentation (i.e. location of displayed fruits on the trunk or on the branches) along a 200 – 2700 m asl rainforest elevation gradient in Papua New Guinea. We found fruit diameter to be lower at higher elevations. While specific fruit colours showed few strong elevational patterns, colours typically attributed to attracting avian dispersers were more prevalent at higher elevations. The proportion of ramiflorous species (bearing fruits from branches) increased with elevation. Finally, we use phylogenetic information to test the “dispersal syndromes” hypothesis: that combinations of fruit traits have evolved in accordance with the preferences and sensory abilities of different frugivore guilds. All fruit traits except presentation showed little evidence of phylogenetic signal but we found fruits displaying colours attributed to mammal frugivory to be larger than “bird colour” fruits. We found evidence for the correlated evolution of fruit size and colour, in support of the dispersal syndromes hypothesis. Fruit traits reflect adaptation to dispersers along a tropical elevational gradient Richard J Hazell* 1 , Graham S Kaina 2,3 , Katerina Sam 4,5 , Daniel Souto-Vilarós 6 , Wulan Koagouw 7 , Vojtech Novotny 4,5 , Mika R Peck 1 , Alan J A Stewart 1 1 School of Life Sciences, University of Sussex, Falmer, United Kingdom. 2 New Guinea Binatang Research Centre, Nagada Harbour, Madang, Papua New Guinea. 3 Food and Agriculture Organization of the United Nations, Project Office in Conservation and Environment Protection Authority, Port Moresby, Papua New Guinea. 4 Biology Centre of Czech Academy of Sciences, Institute of Entomology, Ceske Budejovice, Czech Republic. 5 University of South Bohemia, Faculty of Sciences, Ceske Budejovice, Czech Republic. 6 Science Research Initiative, College of Science, University of Utah, Salt Lake City, Utah, USA. 7 National Research and Innovation Agency (BRIN), KST B.J. Habibie Puspiptek, Tangerang Selatan, Banten, Indonesia. * Corresponding author: [email protected] ; [email protected] +44 7817 646804 John Maynard Smith Building, School of Life Sciences, University of Sussex, Falmer, Brighton, BN1 9QG, United Kingdom. Abstract Fruit traits have the potential to influence disperser communities and vice versa. Here we examine the trends in three fruit traits related to dispersal by frugivores: diameter, colour and presentation (i.e. location of displayed fruits on the trunk or on the branches) along a 200 – 2700 m asl rainforest elevation gradient in Papua New Guinea. We found fruit diameter to be lower at higher elevations. While specific fruit colours showed few strong elevational patterns, colours typically attributed to attracting avian dispersers were more prevalent at higher elevations. The proportion of ramiflorous species (bearing fruits from branches) increased with elevation. Finally, we use phylogenetic information to test the “dispersal syndromes” hypothesis: that combinations of fruit traits have evolved in accordance with the preferences and sensory abilities of different frugivore guilds. All fruit traits except presentation showed little evidence of phylogenetic signal but we found fruits displaying colours attributed to mammal frugivory to be larger than “bird colour” fruits. We found evidence for the correlated evolution of fruit size and colour, in support of the dispersal syndromes hypothesis. Key Words: Fruit traits, elevation, dispersal syndromes, frugivory, gape limitation, phylogeny 3 September, 2025 Dear Editors - Ecology and Evolution, We submit our manuscript examining a simple question central to plant-frugivore coevolution: do fruit traits shift with elevation in ways that track changes in disperser guilds? Along a 200–2700 m a.s.l. rainforest transect in Papua New Guinea, we quantify three dispersal-linked traits (fruit diameter, colour, and presentation) and test predictions from the dispersal-syndromes framework. Our analyses reveal (i) systematic declines in fruit size with elevation, (ii) greater prevalence of “bird-attracting” colours at higher elevations where avian frugivores dominate, and (iii) an increase in ramiflory (fruits presented away from the stem) with elevation. Using comparative phylogenetic methods, we find little phylogenetic signal for size and colour (presentation excepted), and correlated evolution between fruit size and colour, with mammal-associated colours tending to occur in larger fruits. Together, these results support the view that elevational turnover in frugivore assemblages, via gape limitation and sensory biases, has repeatedly and predictably shaped fruit phenotype. We believe this work is an excellent fit for Ecology and Evolution’s broad readership in ecology, evolution, and conservation science. It combines trait-based ecology, macroecological gradients, and phylogenetic inference to provide general insights into how mutualisms drive phenotypic evolution, especially in biodiversity-rich tropical systems. The manuscript is written for accessibility across subfields and emphasizes testable predictions of dispersal syndromes relevant beyond our study region. All authors approve this submission; the work is original and not under consideration elsewhere. We declare no competing interests. Consistent with the journal’s policies, all data, metadata, and analysis code are provided for peer review and will be archived in a public repository (Figshare) with an accompanying Data Availability Statement upon acceptance, enabling full reproducibility. We appreciate your consideration and would be pleased to address any questions. Sincerely, Dr. Richard Hazell Visiting Research Fellow University of Sussex [email protected] Introduction The ability of a plant to disperse its seeds constitutes an important factor determining its survival (Janzen 1970, Connell 1971, Howe and Smallwood 1982, Beckman and Rogers 2013). In response, fruits have evolved into a variety of different forms in order to maximise seed dispersal ability in differing environments. In tropical regions, an estimated 70-90% of plant species have evolved fleshy fruits which are adapted to dispersal by vertebrate frugivores (Muller-Landau and Hardesty 2005). The frugivory mutualism thus has important implications for the evolution of fruit traits in tropical plants – traits related to accessibility and potential attractiveness to frugivores are of clear importance in determining a plant’s fitness. An obvious example concerns fruit size: while larger seeds confer evolutionary advantages related to seedling survival (Mack 1998, Pizo et al. 2006, Souza and Fagundes 2014), a plant’s animal dispersers are limited in the maximum size of seeds they can consume. This sets an upper limit on the size of fruits a plant can produce if it is to be successfully dispersed (Wheelwright 1985). Tropical montane habitats may hold important clues into the relationships between fruit traits and dispersers. Elevational gradients in the tropics are characterised by steady but rapid changes in climatic conditions across relatively small geographical distances. This typically leads to high species turnover of both plant species and their potential dispersers, and a corresponding high turnover of functional traits along tropical elevational gradients. Fruit trait profiles thus have the potential to influence, and in turn to be influenced by, local assemblages of frugivorous species occurring at different elevations (Burns 2013, Dehling et al. 2014, Bender et al. 2018). Nevertheless, while changes in many plant functional traits across elevational gradients are well studied (Swenson and Enquist 2007, Swenson et al. 2011, Hulshof et al. 2013, Read et al. 2014, Asner et al. 2017), surprisingly little is currently known about how fruit traits change with elevation on a community level (but see Chen et al. 2016, Lu et al. 2019). Data on fruit traits are often less readily available than for other plant characteristics because fruits are physically difficult to reach, and present on plants only for short periods of time. If we are to fully understand the functional roles of plants in tropical forests, knowledge of fruit traits is a key component. Of course, individual fruit traits do not exist in isolation. Numerous studies have attempted to detect the presence of “dispersal syndromes”. Akin to pollination syndromes (Fenster et al. 2004), the dispersal syndrome hypothesis states that combinations of fruit traits have evolved as a response to selective pressures from different frugivore guilds (Gautier-Hion et al. 1985, Hererra 1992, Fischer and Chapman 1993, Lomascolo et al. 2008, 2010, Florchinger et al. 2010). For example, birds have acute colour vision and their gape size commonly limits the maximal size of fruits they can consume (Wheelwright 1985, Alcantara and Rey 2003). Mammals are typically larger than birds and can consume and disperse larger fruits. However, outside of the simian primates, mammals generally lack colour vision and rely more on olfactory cues to find fruits (Nevo et al. 2018). Finally, the smaller size of birds means they can easily feed on “ramiflorous” fruits presented among the leaves at the ends of branches, while “cauliflorous” fruits borne on the stem are more likely to be fed on by mammals such as bats (Whittaker and Jones 1994). The island of New Guinea lacks primates, resulting in an exclusively nocturnal mammalian fauna. This creates a clear division of frugivores into a diurnal avian guild and a primarily nocturnal mammalian guild comprising bats, marsupials, and rodents. If the matching of functional traits occurs here between fruits and their frugivore dispersers, we may expect to observe evidence of dispersal syndromes related to fruit colour and size. Bird-dispersed fruits are hypothesised to be smaller and display colours that strongly contrast with a background of bark and foliage, aiding visual detection (Schmidt et al. 2004, Lomascolo et al. 2008). In contrast, as mammals are generally not gape-limited, mammal-dispersed fruits may be larger on average and display a greater range of sizes, reflecting the greater range of feeding techniques used by mammals (Howe 1986). Additionally, since visual cues are likely less important for mammals than for birds, colour is expected to be more variable in mammal-dispersed fruits and duller colours may occur. Evidence for dispersal syndromes is mixed (e.g. Fischer and Chapman 1993, Lomascolo et al. 2008, 2010, Valenta and Nevo 2020, Green et al. 2022). An alternative non-adaptive hypothesis is phylogenetic inertia, whereby fruit size and colour are determined by the size and colour of ancestral species, and frugivores disperse fruits according to pre-determined preferences for certain combinations in a process known as ecological fitting (Janzen 1985, Jordano 1995, Florchinger et al. 2010). If dispersal syndromes occur based on trait combinations selected for by dispersers, then we would expect evidence of correlated evolution of the traits in question, and the appearance of these traits independently in different clades. In contrast, the phylogenetic inertia hypothesis should predict phylogenetic clustering of these traits, indicative of shared evolutionary histories. In this study, we use a fruit trait dataset from a continuously forested elevational gradient in Papua New Guinea to address the following questions: i) How do three key fruit traits related to dispersal by vertebrates change with elevation? ii) Can fruits along the elevational gradient be categorised into dispersal syndromes based on size and colour? iii) To what extent are fruit traits phylogenetically conserved? We hypothesise the following: 1) Fruit size will decrease with increasing elevation, as large dispersers, such as larger birds and frugivorous bats, are more abundant in the lowland areas than at higher elevations (Sam et al. 2019, Sivault et al. 2023). 2) Fruit colours associated with avian dispersal will be more prevalent at higher elevations, where birds are likely to play a dominant role in seed dispersal due to the relative scarcity of mammalian frugivores such as bats (Hazell et al. 2023, Sivault et al. 2023). 3) Ramiflorous fruits will similarly be more prevalent at higher elevations, reflecting an adaptation to avian frugivory. 4) We will detect an association between fruit size and fruit colour categories corresponding to dispersal by either birds or mammals. 5) Fruit traits associated with dispersal by birds vs. mammals will arise independently across the plant phylogeny. Materials and Methods Study Site The study was conducted along the north-eastern slopes of Mt Wilhelm in the northern watershed of the Central Range of Papua New Guinea (Figure 1). The study area is located in the Usino-Bundi district of southern Madang province, and comprises six study sites separated by 500 m elevation, ranging from 200 – 2,700 m above sea level (asl) (5 o 44’ S, 145 o 20’ E; 5 o 49’ S, 145 o 09’ E). The sites represent the lower portion of a complete rainforest transect spanning from the lowland floodplains of the Ramu River to the treeline at 3,700 m asl (Sam et al. 2019). The habitats at the surveyed sites range from lowland alluvial forest (200 m asl) through foothill forest (700 and 1,200 m asl) and lower montane forest (1,700 and 2,200 m asl) to mid montane forest (2,700 m asl) (Paijmans 1976). Mean annual temperature recorded using data loggers decreases from 24.89 °C at 200 m to 14.34 °C at 2,700 m. Average annual precipitation measured by local weather stations is 3,288 mm at 200 m asl, rising to 4,400 mm at 3,700 m asl, with a distinct condensation zone around 2,500 – 2,700 m asl (Sam and Koane 2014, Marki et al. 2016, Sam et al. 2019). Fruit Surveys We collected data on fruiting plants and their fruits using transect surveys (March – July 2016). Ten transects were created at each elevation, each 20 m wide and 500 m in length. This provided a total of 10 ha of sampled area per elevational site. Surveyors walked the transect route, searching carefully for any fruiting woody plants or fallen fruits. Fruiting plants were recorded only if the base of the stem occurred at least partially within the transect. When a fruiting plant was located, we collected data on plant location, diameter at breast height (DBH), growth form and taxonomy (identifying to species level where possible). When necessary, leaf voucher specimens were collected and photographs of stems taken to allow subsequent detailed identification. We also collected information on the method of fruit presentation: either cauliflorous (fruiting directly from the stem) or ramiflorous (fruiting from the branches). In the case of fallen fruits encountered on the ground, we located the most likely source plant using binoculars. In most cases this was not difficult as the plant was still displaying fruits. We collected up to 10 ripe fruits at random from each fruiting plant we encountered (mean = 3.9, SD = 2.8). In cases where fewer than 10 ripe fruits were reachable, we collected as many as safely possible. In cases where fruits were completely unreachable (N = 26 fruiting plants, 2.4% of total), we estimated mean fruit length and width and the colour of ripe fruits, using binoculars. Fruit Measurement We measured the collected fruits as soon as possible after collection to ensure that fruit traits were recorded prior to decomposition. Fruit dimensions were measured to the nearest 0.1 mm using digital callipers. Fruit diameter was defined as the secondary (longest orthogonal to the primary) axis, regardless of the fruit’s morphological characteristics such as stem location or the orientation of seeds. Fruits were weighed using digital scales to the nearest 0.01 g. Fruit colour was defined subjectively using the basic colour categories of “red”, “orange”, “yellow”, “green”, “blue”, “purple”, “pink”, “brown”, “black” and “white”. For bicoloured fruits, the dominant colour (covering >50% of the fruit surface) was considered for analyses. Analyses To enable interspecific analyses of fruit traits across elevations, we first calculated the mean trait values per species at each elevation. For species occurring across multiple elevations, we calculated separate mean values for each elevation at which it was recorded, to allow for any potential intraspecific variation in fruit traits depending on elevation. In addition, we calculated fruit traits weighted by number of individual fruiting plants at each elevation. Analyses of fruit traits across elevations We used generalised linear models (GLMs) to test for the effect of elevation on various fruit traits for both species and abundance weighted data. Analyses were conducted in R version 3.5.2 (R Core Team 2013). For fruit diameter, we used a GLM with a Gaussian error distribution and included plant DBH as a fixed effect. To test the effect of elevation on the proportion of fruiting plants bearing fruits of different colours and different presentation types, we ran separate GLMs for each colour and for each presentation type, using binomial error distributions. Additionally, we tested the effect of elevation on fruit colours when grouped into two categories (see “Fruit Syndromes” below), again using a GLM with binomial error. Tukey pairwise comparisons were performed to adjust p-values for multiple comparisons, calculated using the ‘emmeans’ function in emmeans package (Lenth 2018). Phylogenetic analyses For phylogenetic analyses of fruit traits, we used a global phylogeny adapted by Smith and Brown (2018) from GenBank release 218 (ftp://ftp.ncbi.nlm.nih.gov/genbank) and the Open Tree of Life synthetic tree (taxonomy version 3; https://tree.opentreeoflife.org/about/synthesis-release/v9.1). The phylogeny was subjected to hierarchical analysis with individual phylogenies constructed for major clades, and using a backbone provided by Open Tree of Life version 9.1. We used the prune.sample function in the R package picante (Kembel et al. 2010) to subset this global phylogeny to include only species found in our dataset from the Mt. Wilhelm study sites. In order to determine whether the categorical traits of colour and presentation were clustered or randomly distributed across the phylogeny, we used a null model analysis. We first calculated the mean phylogenetic distance (MPD) between individuals of each colour and of each presentation type, and then compared this to a distribution of values generated by shuffling the tip labels across the phylogeny 999 times. We then assessed the deviation of observed and null values. Null models were implemented using functions in the package picante . For the continuous trait of fruit diameter, we used Pagel’s lambda (λ) (Pagel 1999a) to test for phylogenetic signal in the trait data. Pagel’s λ uses phylogenetic data to assess whether a trait has evolved independently of phylogeny (low phylogenetic signal) or if it conforms to an evolutionary model expected under Brownian motion (high phylogenetic signal) (Molina-Venegas and Rodriguez 2017). Fruit Syndromes The fruit syndrome hypothesis predicts correlated evolutionary change in fruit size and colour according to dispersal guild (Lomascolo et al. 2008). To test the hypothesis that fruit traits corresponding to dispersal by birds vs. mammals evolved together, we first divided fruits into binary size and colour categories corresponding to each dispersal syndrome. Fruits were divided by colour into “mammal colour” (brown, green, yellow and orange) and “bird colour” (red, pink, purple, blue, black, white), according to Janson’s (1983) classification. Fruits were divided by size based on mean fruit diameter for all fruiting plant species recorded in our data. We performed Pagel’s likelihood ratio test of binary correlations (Pagel 1994, 1999b), using the subset of fruiting plant species for which phylogenetic data was available, on the two binary categories of colour (mammal vs. bird colour) and size (large vs. small), using the ‘fitPagel’ function in the package phytools (Revell 2012). Pagel’s test compares the goodness of fit of a model of correlated evolution to one of independent evolution, considering phylogenetic branch lengths. To test whether mammal colour fruits were larger overall than bird colour fruits (regardless of phylogeny), we performed a Mann-Whitney U test using fruit diameter per fruiting plant as the response variable. To test whether variation in fruit diameter was greater for mammal colour fruits than for bird colour fruits, we used a Fligner-Killeen test, which is a non-parametric test for assessing the homogeneity of variances (Fligner and Killeen 1976). Results We collected and measured ripe fruits from a total of 1062 fruiting plant individuals across all elevations, representing 167 plant species and morpho-species. Twenty-one species and morpho-species were recorded from 200 m, 43 from 700 m, 38 from 1200 m, 33 from 1700 m, 38 from 2200 m and 28 from 2700 m asl (Table A1). Of these, 83 species were sufficiently identified to be used in phylogenetic analyses. Outside of phylogenetic analyses, we used data from all fruiting plant species and morpho-species. Mean fruit diameter showed a significant decrease towards higher elevations when weighted both by species (p < 0.01; Figure 2a) and individual fruiting plants (p < 0.01; Figure A1a). Fruit diameter was also positively correlated with fruiting plant DBH (p < 0.01; Figure A2). Fruit colour when divided into mammal vs. bird colour also showed a significant trend – the proportion of species bearing bird colour fruits increased with elevation (p < 0.01, Figure 2b, Table 1). A similar trend was observed for individual-weighted data (p < 0.01, Figure A1b), although a high number of plant individuals displayed bird colour fruits at 200 m, primarily attributable to white fruits (which may also be attractive to mammals). Other fruit colours showed differing patterns depending on whether weighted by species or individual, although most colours were represented at all or nearly all elevations. Using species-weighted data, most individual fruit colours did not show strong elevational trends (Table 1, Table A2), although richness and abundance of species bearing green fruits peaked at 700 m, and purple fruits at 2700 m. Fruit presentation showed a clear trend with elevation: less than 40% of fruiting species were exclusively ramiflorous at 200 m, increasing to 100% at 2700 m (p < 0.01; Figure 2c). We observed a similar increase with elevation in individual-weighted data (p < 0.01; Figure A1c). Phylogenetic analysis on the categorical traits of fruit colour and presentation showed broadly contrasting patterns. In general, fruit colour did not differ from random expectations, suggesting that fruit colour is not phylogenetically constrained and rather subject to selection (Figure 3a, Table A4). Of all colour categories, only brown fruits appear to be phylogenetically clustered (SES.MPD = -2.21, p = 0.03). Fruit presentation however showed evidence of significant phylogenetic clustering: cauliflorous fruits were clustered significantly more than expected under null models (SES.MPD = -7.12, p < 0.01; Figure 3b). For the continuous trait of fruit diameter, a low phylogenetic signal indicated a lack of phylogenetic clustering (λ = 0.29, p = 0.02). Both phylogenetic and non-phylogenetic analysis revealed significant relationships between fruit colour and size. We observed a significant phylogenetic association between “large” and “small” fruits, and mammal colour and bird colour fruits respectively (likelihood ratio = 14.33, p < 0.01; Figure 4a), showing evidence of correlated evolution of fruit colour and size. Regardless of phylogeny, we found mammal colour fruits to have significantly greater median diameter than bird colour fruits (median mammal colour = 14.06 mm, median bird colour = 10.2 mm, W = 6026, p < 0.01; Figure 4b), and to show a greater variation in size (Fligner-Killeen median χ2 = 18.04, p < 0.01; Figure 4b). The mean diameters of each fruit category were 17 mm (mammal colour) and 12.8 mm (bird colour). Discussion In this study, we examined three key fruit traits – diameter, colour, and presentation – associated with dispersal by vertebrate frugivores along the elevational gradient of Mt. Wilhelm in Papua New Guinea. To our knowledge, this work provides the first comprehensive analysis of fruit traits across a tropical fruiting plant community spanning an elevational gradient. By analysing these traits, we also tested the dispersal syndromes hypothesis, which posits that specific fruit characteristics are shaped by the preferences of their primary dispersers. Fruit Diameter and Elevation We found mean fruit diameter to decrease towards higher elevations, after a plateau between 200 and 1200 m. Our result mirrors a trend observed by Almeida-Neto et al. (2008), who found fruit diameter to be lower at higher elevations across 135 sites in the Brazilian Atlantic rainforest. A few potential factors could explain the observed pattern: i) A decrease in productivity with decreasing temperature at higher elevations limits the production of large fruits. ii) Fruit size is constrained by plant size, which in turn decreases with elevation. iii) A lack of large-gaped frugivores at high elevations prevents the seed dispersal of large-fruited plants. These hypotheses are not mutually exclusive, and our data likely reflect a combination of factors; here we briefly discuss each one. As temperature decreases consistently with elevation, as recorded by dataloggers at the study site (Sam et al. 2019), one might expect fruit diameter to show a corresponding steady decline if productivity were the dominant factor influencing seed size. However, Almeida-Neto et al. (2008) found that differences in fruit diameter with elevation were primarily not attributable to temperature. Specifically, low-elevation communities had greater mean fruit diameter than higher-elevation communities experiencing similar annual temperatures (i.e., those from warmer areas). Combined with our observation that fruit size does not follow a similarly steady decrease with elevation, the evidence suggests that temperature alone cannot account for the pattern in fruit diameter observed here. Broader productivity factors, such as rainfall, may help to account for fruit size trends, but while Sam et al. (2024) report higher annual rainfall at 2700 m than at 200 m, we lack rainfall data for intermediate sites to adequately test the relationship. Regarding the potential influence of plant size, we found a positive correlation between fruit diameter and fruiting plant DBH, yet elevation still explained variation in fruit size even after accounting for DBH. The relationship between fruit size and disperser size at the community level is complex, making it difficult to determine evolutionary cause and effect, although an association between the two groups has been noted by several studies (Burns 2013, Dehling et al. 2014, Muñoz et al. 2017, Bender et al. 2018). Our data, which show a low-elevation plateau in fruit diameter followed by a decline towards mid-elevations, is consistent with a change in the avian frugivore community observed between lowland and highland forest. Sam et al. (2019) documented a mid-elevational shift in the distribution of the large-gaped frugivores, such as the Papuan hornbill Rhyticeros plicatus and the majority of pigeon species, which are primarily found between 200 and 1200 m asl. Furthermore, species richness of bats (which are not gape-limited and may consume larger fruits than birds) is known to decline with elevation in Papua New Guinea (McCain 2007), and in the study area specifically (Sivault et al. 2023). Hazell et al. (2023) noted a decline in bat frugivory with elevation here, as has also been noted elsewhere (McCain 2007, Almeida-Neto et al. 2008). Fruit Colour and Elevation Prevalence of combined bird colour fruits (red, pink, purple, blue, black, white) increased with increased elevation, specifically driven by their dominance at 2700 m. This trend supports our second hypothesis, which states that the increased relative importance of avian frugivory at higher elevations would be associated with higher prevalence of these fruit colours, in line with the dispersal syndromes hypothesis (see below). Meanwhile there were few clear trends in specific fruit colours along the elevational gradient, although green, purple and black fruits each showed some significant variations in prevalence. A recent study found no relationship between fruit colour and specific avian colour preferences along the same gradient (Hazell et al. 2023), suggesting birds are exerting little selective pressure on specific fruit colours. Fruit colour may also reflect an adaptation to abiotic factors (Burns 2015, Valenta et al. 2018). Anthocyanins, which are responsible for blue, deep red, purple and black colours in fruit, additionally serve an important role in protecting plants from abiotic stressors such as heat, drought, and salinity (Kaur et al. 2023). Thus, their presence in other plant tissues and in fruits may be coupled, meaning fruit colours characterised by high anthocyanin content could simply be a by-product of this effect (Stournaras and Schaefer 2017). Fruit presentation and Elevation Elevation had a significant effect on the presentation method of fruits. Cauliflorous fruits were relatively common at 200 m but steadily declined with elevation and were absent at 2700 m. This pattern aligns with the sharp decline in bat frugivory with increasing elevation. Unlike birds, which typically require a branch on which to perch while feeding, bats are known to preferentially forage on fruits borne away from foliage, including cauliflorous fruits (Whittaker and Jones 1994). The above-mentioned elevational decrease in bat species richness (Sivault et al. 2023) and frugivory (Hazell et al. 2023) in the study area may therefore help to explain the parallel decline in cauliflorous fruits. In contrast, arboreal mammals, including rodents and marsupials, do not show a similar decline across the elevations in this study (Hazell et al. 2023, Vejmělka In Prepatation ). Arboreal mammals may easily access fruit presented on the main stem of a fruiting plant, and thus could potentially support dispersal of cauliflorous fruits at higher elevations. However, the relatively local scale of non-volant mammalian dispersal, combined with the higher probability of seed destruction by rodents in particular (Denslow & Moermond 1982) may serve to weaken any evolutionary effect on fruits. It should be noted that cauliflory showed significant phylogenetic clustering in this study, meaning the effects of phylogenetic inertia cannot be discounted in structuring the observed patterns. Nevertheless, as with fruit diameter, the decrease in cauliflory with elevation suggests a relatively greater importance of avian frugivores at higher elevations, as also observed by Almeida-Neto et al. (2008). Fruit Syndromes Our results support the dispersal syndrome hypothesis. Across elevations, we demonstrated a clear relationship between fruit colour category (the groups of colours commonly attributed to dispersal by either mammals or birds) and fruit diameter. We found mammal colour fruits (brown, green, yellow and orange) to be larger than bird colour fruits (red, pink, purple, blue, black and white) and to show a greater variation in diameter. Both of these factors are consistent with adaptation to dispersal by mammalian vs. avian frugivores. The association between fruit colour category and diameter appears across separate clades within the phylogeny of fruiting plants, indicating correlated evolution of size and colour category. This occurs despite limited evidence of phylogenetic clustering of individual fruit colours, which are known to be evolutionarily labile. In a wide-ranging study of avian frugivory in a subtropical Andean forest, Ordano et al. (2017) found only weak phylogenetic effects on fruit chromatic contrast and conspicuousness of fruiting displays. A similarly broad study on fruit traits in the Brazilian Atlantic forest found no significant phylogenetic signal in fruit colour, although it was present in several other fruit traits (Cazetta et al. 2012). On a broader scale, Stournaras et al. (2013) found minimal evidence of phylogenetic constraints on fruit colour at local and global scales, despite fruit colour being limited by other factors such as chemical constraints. The fact that individual fruit colours are spread across the phylogeny, but that colour still correlates with fruit size at the broader level of colour category (mammal vs. bird), supports the hypothesis that fruit colours have adapted independently to dispersal by different frugivore guilds. Thus, we discount the hypothesis of phylogenetic inertia as the means of explaining observed fruit colour/size combinations. Summary We observed that fruit traits related to mammalian frugivory (large size, dull colours, ramiflorous presentation) decreased at higher elevations, while traits related to avian frugivory generally increased. A significant relationship between fruit size and colour categories across sites, independent of phylogeny, supports the dispersal syndromes hypothesis. The fruit traits investigated are clearly important in determining frugivory by different guilds, influencing seed dispersal and ultimately plant community assembly. However, the shifting distributions of plants and frugivores resulting from climate change has the potential to disrupt existing relationships and should be a focus of further study. Author Contributions Richard J Hazell: Conceptualization; data curation; formal analysis; investigation; methodology; project administration; resources; visualization; writing – original draft; writing – review and editing. Graham S Kaina: Investigation; methodology; project administration; resources; writing – review and editing. Katerina Sam: Conceptualization; funding acquisition; methodology; project administration; resources; supervision; visualization; writing – review and editing. Daniel Souto-Vilarós: Conceptualization; data curation; formal analysis; methodology; resources; writing – review and editing. Wulan Koagouw: Formal analysis; project administration; visualization; writing – review and editing. Vojtech Novotny: Conceptualization; funding acquisition; methodology; writing – review and editing. Mika R Peck: Funding acquisition; methodology; project administration; supervision; writing – review and editing. Alan J A Stewart: Funding acquisition; methodology; project administration; supervision; writing – review and editing. Acknowledgements We are grateful to the communities along the Mt Wilhelm elevational gradient, especially to landowners Francis Yama, Simon Yama, Markus Bundikana, Ava Memeku, Ambros Gena and Peter Sai for allowing us access onto their land. Thank you also to the local field assistants for all their help with fruit collection and measurement, and also to Salape Tulai and Luda Paul for helping to manage collection activities when needed. Thanks also go to Martin Jung for his help in organising the fruit data back in the UK, and to Simon Segar for providing some of the code necessary for performing phylogenetic analyses. Finally we thank František Vejmělka for sharing data on mammalian diets on the Mt Wilhelm gradient. We acknowledge the funding support of the UK Darwin Initiative (19-008). RH would like to thank the Sir Richard Stapley Educational Trust for its generous financial support. VN was funded by Praemium Academie from CAS. 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Results of generalised linear models testing the effect of elevation on fruit diameter, fruit colour (both for individual colours and colour category, i.e., colours grouped according to bird vs. mammal dispersal syndromes) and presentation type (ramiflorous vs. cauliflorous) for fruiting plant species along the Mt Wilhelm elevational gradient. Significant (p < 0.05) individual pairwise interactions between elevations are presented, after correcting for multiple comparisons using Tukey pairwise comparisons. Significance values for all pairwise comparisons are displayed in full in the appendix (Table A2). Fruit Diameter 2338.1 <0.01 700-2700; 1200-2700 Colour Red 9.89 0.08 None Orange 11.99 0.03 None Yellow 21.69 <0.01 None Green 21.79 <0.01 700-1700; 700-2200 Blue 8.62 0.13 None Purple 45.47 <0.01 700-2700; 1700-2700; 2200-2700 Pink 37.13 <0.01 1700-2200 Brown 18.35 <0.01 None Black 15.68 <0.01 1700-2700 White 2.7 0.75 None Colour Category 35.73 <0.01 700-2700; 1200-2700; 1700-2700 Presentation Cauliflorous 36.79 <0.01 None Ramiflorous 55.09 <0.01 200-700; 200-1200; 200-1700; 200-2200; 200-2700; 700-2700 Figure Legends Figure 1. Map of the Mt Wilhelm survey area, including location within Papua New Guinea (inset). White numbers next to survey sites denote metres above sea level. Figure 2. Effect of elevation on three fruit traits related to dispersal by frugivores. a) Mean fruit diameter per fruiting plant species is represented by squares, with error bars displaying 95% confidence intervals (CIs). Letters above the points denote significant differences after adjusting for multiple comparisons using Tukey pairwise tests. b) Proportion of fruiting plant species displaying fruits of each colour (top to bottom at 1200 m asl: white, black, red, pink, purple, blue, green, yellow, orange, brown) at each elevation. Mammal colour fruits are hatched to highlight the proportions of mammal vs. bird colour fruits. c) Proportion of fruiting plant species bearing cauliflorous fruits (white bars), ramiflorous fruits (dark grey bars) and a combination of both presentation types (light grey bars) at each elevation. Figure 3. Phylogenies of the 83 fruiting plant species for which phylogenetic data was available, including fruit colour (a) and the presentation method of fruits (b). In (a), colours of squares represent the predominant colours of the fruits of that species (see Figure 2b for colour list). In (b), blue squares represent ramiflorous fruiting species, while orange squares represent cauliflorous species. Species without squares attributed to them indicate a combination of ramiflory and cauliflory within a single species. Numbers correspond to species names listed in Table A3. Prominent genera (represented by more than two species) are displayed on the right. Figure 4. Relationships between fruit colour and diameter for all fruiting plant species and morpho-species recorded along the Mt Wilhelm gradient. a) Visualisation of data from Pagel’s likelihood ratio test between fruit colour (left phylogeny) and diameter (right phylogeny), where each trait is represented by two binary categories. For colour, grey circles represent “mammal colour” (green, yellow, orange and brown) fruits, and black circles “bird colour” (red, pink, purple, blue, black and white) fruits. For mean diameter, grey circles represent “large-fruited” species, and black circles “small-fruited” species. b) Relationship between fruit colour category (as above) and fruit diameter, this time displayed as a continuous variable. Grey boxes span the first and third quartiles of fruit diameter and the horizontal line within each box represents the median diameter. Vertical lines indicate maximum and minimum observations falling within 1.5 times the interquartile range. Remaining observations are displayed as black circles. Appendix 1 Note on plant phylogeny: The phylogeny used in this study is adapted from Open Tree of Life version 9.1, following hierarchical analysis with individual phylogenies constructed for major clades, and can be found at https://github.com/FePhyFoFum/big_seed_plant_trees/releases (labelled “ALLOTB”), in addition to the Figshare repository for project data. Appendix 1 Figure Legends Figure A1. Effect of elevation on four fruit traits related to dispersal by frugivores, here weighted by plant individual. a) Mean fruit diameter per fruiting plant is represented by squares, with error bars dispalaying 95% confidence intervals. Letters above points denote significant differences after adjusting for multiple comparisons using Tukey pairwise tests. b) Proportion of individual fruiting plants displaying fruits of each colour (top to bottom: white, black, red, pink, purple, blue, green, yellow, orange, brown) at each elevation. “Mammal colour” fruits are hatched to highlight the proportions of mammal vs. bird colour fruits. c) Proportion of fruiting plants bearing cauliflorous fruits (white bars) and ramiflorous fruits (dark grey bars) at each elevation. Figure A2. Relationship between plant diameter at breast height (DBH) and fruit diameter for all plant species recorded along the elevational gradient. Each circle represents mean DBH and fruit diameter values for a single species. The x-axis is log-transformed for display purposes. The blue line represents a line of best fit, with grey bars representing 95% confidence intervals. R2 is 0.064. Appendix 1 Tables Table A1. List of fruiting plant species and morpho-species recorded at each elevation, together with information on fruit traits. Species means are displayed for fruit diameter and for the proportion of fruit weight attributable to seeds. NA values are given where seeds could not easily be separately weighed (e.g. compound fruits). Presentation types include cauliflorous (C), ramiflorous (R) and “C+R” (species with individuals displaying either presentation type). NA values are given for species for which individuals displayed both types of presentation simultaneously. The mode fruit colour is also listed for each species. 200 m Aglaia sp. 1 21.4 0.452556818 NA orange 200 m Allophylus cobbe 6.466666667 0.397959184 C red 200 m ANNONACEAE sp. 1 13.98 0.537081024 C black 200 m Artocarpus lakoocha 7.228571429 NA C yellow 200 m Casearia clutiifolia 16.05 0.405228758 NA orange 200 m Erythrospermum candidum 9 0.269230769 NA green 200 m Ficus congesta 38 0.325098039 C brown 200 m Ficus sp. 5 20.16 0.208609272 C green 200 m Ganalium sp. 1 22.06666667 0.430965682 R black 200 m Harpullia ramiflora 22.33333333 0.07753051 C red 200 m Leea indica 11.17142857 0.191366906 C+R black 200 m Litsea collina 14.25714286 0.493804956 NA red 200 m Micromelum minutum 5.9 0.361111111 NA orange 200 m Myristica sp. 3 20.5 0.139720559 NA yellow 200 m Myristica sp. 4 19.5 0.452399685 NA brown 200 m Pittosporum sinuatum 11.5 NA R red 200 m Psychotria leptothyrsa 11.94 0.25 R red 200 m Psychotria sp. 1 5.533333333 0.459016393 R white 200 m Psychotria sp. 2 12.91666667 0.176056338 R white 200 m SAPINDACEAE sp. 1 25.63 0.5944 C+R red 200 m Wenzelia dolichophylla 35.225 0.139579148 NA orange 700 m Aglaia sp. 2 13.82857143 0.59268762 R red 700 m ANNONACEAE sp. 2 9.43 0.441558442 R red 700 m Archidendron aruense 19.3 0.114798206 C orange 700 m Archidendron sp. 1 27.1 0.070609812 C red 700 m Ardisia imperialis 6.5 0.454545455 R purple 700 m Ardisia sp. 2 11.06086957 0.493767313 R green 700 m Ardisia sp. 1 10.87777778 0.300306435 R green 700 m Clerodendrum sp. 1 9.8 0.574468085 R black 700 m Cryptocarya sp. 1 11.26666667 0.513368984 R green 700 m Cupaniopsis acuticarpa 20.975 0.375780275 R orange 700 m Cupaniopsis sp. 1 13.7 0.15503876 R green 700 m Ficus badiopurpurea 5.1 0.235294118 R green 700 m Ficus bernaysii 18.8015873 0.310492776 C green 700 m Ficus hahliana 15.4 0.217252396 C green 700 m Ficus morobensis 39.55 0.129220509 C brown 700 m Ficus pungens 6.555 0.367454068 C brown 700 m Ficus saccata 18.3 0.125364431 NA orange 700 m Ficus sangumae 13.8 0.218023256 C red 700 m Ficus sp. 3 29.83333333 NA R brown 700 m Ficus sp. 4 47.05 0.258940112 C green 700 m Ficus wassa 14.7 0.317567568 C red 700 m Glochidion angulatum 5.8 0.210526316 R green 700 m Gymnacranthera paniculata 20.85 0.095317726 R orange 700 m Harpullia longipetala 23.63131222 0.215984645 C+R orange 700 m Harpullia sp. 1 24.5 0.068085106 R green 700 m Harpullia sp. 2 20.16 0.030376671 C green 700 m Homalanthus novoguineensis 5.9 0.125 R red 700 m Kibara sp. 2 9.7 0.481927711 R black 700 m LAMIACEAE sp. 1 8.2 0.285714286 R red 700 m Leucosyke australis 14.7 0.255093003 R white 700 m Leucosyke sp. 1 10.57 NA R white 700 m Magnolia tsiampacca 13.7 0.691442629 R orange 700 m Medinilla crassinervia 17.75714286 0.41503268 C+R purple 700 m Melastoma sp. 1 9.3 NA R red 700 m Myristica subalulata 5.466666667 0.466666667 R orange 700 m Pandanus kaernbachii 7.85 0.152091255 R orange 700 m Pittosporum sinuatum 9.2 NA R red 700 m Planchonella sp. 1 61.32 0.562448582 R black 700 m Popowia pisocarpa 13.51333333 0.235361653 C+R black 700 m Psychotria micrococca 6.78 0.339233038 R white 700 m Sterculia schumanniana 26.73333333 0.174298045 R green 700 m Syzygium gonathantum 50.1 0.246326476 R red 700 m Syzygium sp. 1 27.6 0.238843931 R red 1200 m Aglaia tomentosa 12.36666667 0.275862069 R green 1200 m Annesijoa novoguineensis 19.58 0.085042521 R yellow 1200 m ANNONACEAE sp. 3 11.76666667 0.153488372 R purple 1200 m Ardisia imperialis 6.891666667 0.680208333 R black 1200 m Ardisia lanceolata 14.5 0.293144208 R pink 1200 m Ardisia sp. 2 10.75555556 0.430156472 R red 1200 m Ardisia sp. 1 9.85 0.684039088 R red 1200 m Areca sp. 1 10.5125 0.732732733 R red 1200 m Caryota rumphiana 14.23 NA C black 1200 m Cyclophyllum brevipes 7.3 0.416666667 R blue 1200 m Cyrtandra erectiloba 12.45 0.56119403 R white 1200 m FAGACEAE sp. 1 17.8125 NA R black 1200 m Garcinia maluensis 28.8 0.532967033 R orange 1200 m Harpullia longipetala 18.72857143 0.563312229 NA red 1200 m Harpullia sp. 3 13.22 0.581772334 C red 1200 m Kibara sp. 1 9.06 0.47826087 R black 1200 m Magnolia tsiampacca 24.93333333 NA R green 1200 m Medinilla crassinervia 15.1 NA C green 1200 m Melastoma sp. 1 16.15 0.320366133 R pink 1200 m Myristica fatua 58.4 0.027943934 R orange 1200 m Myristica filipes 65.9 0.224500898 R brown 1200 m Myristica sp. 1 16.375 0.519174041 R orange 1200 m Myristica sp. 2 19.8 0.319218241 R brown 1200 m Myristica subalulata 27.5 NA R brown 1200 m Planchonella macropoda 43.2 0.31828784 R black 1200 m Prunus dolichobotrys 24.9 NA R orange 1200 m Rapanea involucrata 10.3 0.46875 R red 1200 m RUBIACEAE sp. 1 9.45 0.254901961 R purple 1200 m RUBIACEAE sp. 2 11.42 0.460893855 R white 1200 m RUBIACEAE sp. 3 9.842857143 0.088328076 R purple 1200 m RUBIACEAE sp. 3 10.4826087 0.270524582 R blue 1200 m RUBIACEAE sp. 3 10.57142857 0.270306258 R purple 1200 m SOLANACEAE sp. 1 29 0.524161074 NA orange 1200 m Solanum sp. 1 12.93333333 NA R orange 1200 m Solanum sp. 2 15.3 0.646586345 R red 1200 m Terminalia sp. 1 31.1 0.687538493 R green 1200 m ULMACEAE sp. 1 5.031578947 NA R white 1200 m Ziziphus angustifolia 12.7125 0.568421053 R purple 1700 m Ardisia sp. 4 26.3 NA R pink 1700 m Canarium sp. 1 9.8 0.224299065 R black 1700 m Chisocheton sp. 1 33.4 0.197997775 R red 1700 m Ficus arfakensis 16.85 0.41285297 R brown 1700 m Ficus hahliana 20.83333333 0.343481138 C+R red 1700 m Ficus sangumae 12.42857143 0.427622842 C yellow 1700 m Ficus subulata 4.4 0.147058824 R yellow 1700 m Ficus trachypison 14.15 0.324324324 R orange 1700 m Ficus trichocerasa 12.25 0.201834862 C+R red 1700 m Ficus wassa 9.8 0.24 C+R red 1700 m Kibara coriacea 10.23333333 0.421276596 R purple 1700 m Lepidopetalum comesperma 9.8 0.478021978 R green 1700 m Melicope elleryana 17.4 0.026359143 R brown 1700 m Myristica filipes 44.3 NA R brown 1700 m Osmoxylon sp. 1 1.433333333 NA R orange 1700 m Osmoxylon sp. 2 5.7 0.625 R black 1700 m Piper interruptum 7.468292683 NA R red 1700 m Piper macropiper 2.4 NA R brown 1700 m Piper recessum 6.775 NA R yellow 1700 m Piper sp. 1 8.5 NA R red 1700 m Piper sp. 2 9.2 NA R red 1700 m Piper sp. 3 10.4 NA R orange 1700 m Pittosporum sp. 1 10.53636364 0.620147874 R orange 1700 m Psychotria micrococca 9.85 0.548148148 R white 1700 m Psychotria sp. 3 4.6 0.5 R white 1700 m RHAMNACEAE sp. 1 10.2 0.507246377 R black 1700 m RUBIACEAE sp. 4 4.014285714 0.511415525 R white 1700 m Saurauia conferta 29.66666667 NA R green 1700 m Smilax nova-guineensis 13 0.325581395 R black 1700 m Sterculia schumanniana 15.54285714 0.54754289 R black 1700 m Tabernaemontana pandacaqui 10.2 NA R red 1700 m VITACEAE sp. 1 2.283333333 NA R purple 1700 m Xanthophyllum papuanum 10.6 0.375838926 R brown 2200 m ANNONACEAE sp. 1 28.9 0.126213592 R green 2200 m Ardisia sp. 2 9 0.346153846 R pink 2200 m Ardisia sp. 1 7.42 0.233918129 R black 2200 m Astronidium sp. 1 12.8 0.172506739 R white 2200 m Cayratia trifolia 9.5 0.26 R black 2200 m Cyrtandra erectiloba 10.7 NA R green 2200 m Decaspermum alpinum 5.05 0.588235294 R orange 2200 m Decaspermum forbesii 5.2 0.25 R orange 2200 m Embelia cotinoides 12.5 NA R pink 2200 m Ficus iodotricha 41.84 NA C red 2200 m Ficus microdictya 19.8 NA NA brown 2200 m Ficus sangumae 15.55 0.291666667 C+R pink 2200 m Ficus sp. 1 16.05 0.138790036 R green 2200 m Ficus sp. 2 22.35 0.259932245 R red 2200 m Ficus wassa 7.483125 0.399505728 C+R pink 2200 m Hydriastele sp. 1 14.22 0.493333333 C+R red 2200 m Ichnocarpus frutescens 9.4 0.266666667 R black 2200 m Kibara coriacea 10.73333333 0.652892562 R black 2200 m Maesa haplobotrys 3.966666667 0.333333333 R brown 2200 m Medinilla crassinervia 5.2 NA C+R white 2200 m Medinilla lorentziana 5.5 0.25 R brown 2200 m Melastoma sp. 2 7.5 0.428571429 R green 2200 m Melicope sp. 1 16.575 0.018034265 R yellow 2200 m Myrsine womersleyi 5.351443001 0.366424337 R purple 2200 m Osmoxylon sp. 1 1.385714286 NA R orange 2200 m Pandanus sp. 1 9.42 0.042666667 C+R red 2200 m Piper celtidiforme 12.8 NA R orange 2200 m Piper macropiper 3.7544 NA R red 2200 m Piper sp. 5 7.1 NA R orange 2200 m Pittosporum sp. 1 7.74375 NA R red 2200 m Polyosma cunninghamii 8.1 0.125 R black 2200 m Psychotria murmurensis 5.2 NA R orange 2200 m Scaevola oppositifolia 5.65 0.5 R black 2200 m Smilax nova-guineensis 10.2 0.4375 R brown 2200 m Timonius sp. 1 31.4 NA R red 2200 m Tinospora cordifolia 5.6 0.363636364 R yellow 2200 m Tripetalum cymosum 34.25 0.22165069 R red 2200 m Zehneria mucronata 6.4 0.153846154 R brown 2700 m Ardisia sp. 1 5.777192982 0.224238579 R black 2700 m Ardisia sp. 3 5.625 0.347826087 R black 2700 m Bhesa archboldiana 3.6 0.6 R red 2700 m Breynia cernua 6.6 0.434782609 R black 2700 m Decaspermum alpinum 9.3 NA R red 2700 m Elaeocarpus nymanii 14.16 0.806060606 R green 2700 m Eurya tigang 9.2 NA R purple 2700 m Glochidion sp. 2 4.3 0.375 R red 2700 m Glochidion sp. 1 4.22 0.396825397 R red 2700 m Kibara coriacea 11.12 0.516332982 R black 2700 m Kibara sp. 3 19.08 0.266677409 R red 2700 m Maesa edulis 4.166666667 0.4 R red 2700 m Myrsine involucrata 3.964788732 0.405444887 R purple 2700 m Myrsine womersleyi 7.007894737 0.428751576 R purple 2700 m Palmeria sp. 1 7.866666667 0.448275862 R red 2700 m Piper sp. 4 16.2 NA R orange 2700 m Polyosma cunninghamii 8.633333333 0.419548872 R purple 2700 m Prunus oligantha 13.15714286 0.481472081 R red 2700 m Prunus sp. 1 14.7 0.452631579 R black 2700 m Psychotria multicostata 9.990909091 0.299354244 R white 2700 m RUBIACEAE sp. 1 7.8 NA R orange 2700 m Saurauia sp. 1 12.01428571 0.291233284 R green 2700 m Solanum sp. 1 8.6 0.625 R red 2700 m Steganthera sp. 1 9.24 0.631141345 R black 2700 m Streblus glaber 5.6 0 R green 2700 m Streblus sp. 1 7.425 0.379310345 R black 2700 m Zygogynum haplopus 5.216666667 0.295081967 R black 2700 m Zygogynum oligostigma 15.25 NA R red Table A2. Results of Tukey pairwise comparisons from generalised linear models testing the effect of elevation on different fruit traits (left column). Estimate and standard error values are displayed, together with z ratios and p values adjusted for multiple comparisons. Significant comparisons are displayed in bold. Fruit Diameter 200 - 700 -0.885 2.76 -0.321 0.9995 200 - 1200 -1.672 2.81 -0.594 0.9915 200 - 1700 4.148 2.89 1.436 0.7051 200 - 2200 4.819 2.81 1.713 0.5234 200 - 2700 7.781 2.99 2.605 0.0961 700 - 1200 -0.787 2.3 -0.342 0.9994 700 - 1700 5.032 2.39 2.101 0.2865 700 - 2200 5.703 2.3 2.475 0.1314 700 - 2700 8.665 2.51 3.448 0.0075 1200 - 1700 5.819 2.46 2.363 0.1693 1200 - 2200 6.49 2.37 2.734 0.0687 1200 - 2700 9.452 2.58 3.667 0.0033 1700 - 2200 0.671 2.46 0.272 0.9998 1700 - 2700 3.633 2.66 1.366 0.7473 2200 - 2700 2.962 2.58 1.149 0.8606 Seed Proportion 200 - 700 0.17847 0.599 0.298 0.9997 200 - 1200 -0.34155 0.611 -0.559 0.9936 200 - 1700 -0.20215 0.662 -0.305 0.9996 200 - 2200 0.17547 0.648 0.271 0.9998 200 - 2700 -0.33951 0.645 -0.527 0.9951 700 - 1200 -0.52003 0.51 -1.019 0.9117 700 - 1700 -0.38062 0.57 -0.668 0.9854 700 - 2200 -0.003 0.554 -0.005 1 700 - 2700 -0.51799 0.55 -0.942 0.9355 1200 - 1700 0.13941 0.582 0.239 0.9999 1200 - 2200 0.51703 0.567 0.912 0.9437 1200 - 2700 0.00204 0.563 0.004 1 1700 - 2200 0.37762 0.621 0.608 0.9905 1700 - 2700 -0.13737 0.617 -0.223 0.9999 2200 - 2700 -0.51499 0.603 -0.854 0.9571 Colour Category: Mammal 200 - 700 -0.883 0.452 -1.954 0.3692 200 - 1200 -0.2311 0.486 -0.475 0.997 200 - 1700 -0.6705 0.428 -1.566 0.6213 200 - 2200 0.0873 0.455 0.192 1 200 - 2700 1.2217 0.539 2.268 0.2072 700 - 1200 0.6519 0.381 1.709 0.5257 700 - 1700 0.2125 0.304 0.699 0.9821 700 - 2200 0.9704 0.341 2.843 0.0509 700 - 2700 2.1047 0.446 4.716 <.0001 1200 - 1700 -0.4394 0.353 -1.244 0.8149 1200 - 2200 0.3185 0.386 0.826 0.9629 1200 - 2700 1.4528 0.481 3.02 0.0304 1700 - 2200 0.7578 0.309 2.451 0.1391 1700 - 2700 1.8922 0.422 4.481 0.0001 2200 - 2700 1.1343 0.45 2.521 0.1178 Colour Category: Bird 200 - 700 0.883 0.452 1.954 0.3692 200 - 1200 0.2311 0.486 0.475 0.997 200 - 1700 0.6705 0.428 1.566 0.6213 200 - 2200 -0.0873 0.455 -0.192 1 200 - 2700 -1.2217 0.539 -2.268 0.2072 700 - 1200 -0.6519 0.381 -1.709 0.5257 700 - 1700 -0.2125 0.304 -0.699 0.9821 700 - 2200 -0.9704 0.341 -2.843 0.0509 700 - 2700 -2.1047 0.446 -4.716 <.0001 1200 - 1700 0.4394 0.353 1.244 0.8149 1200 - 2200 -0.3185 0.386 -0.826 0.9629 1200 - 2700 -1.4528 0.481 -3.02 0.0304 1700 - 2200 -0.7578 0.309 -2.451 0.1391 1700 - 2700 -1.8922 0.422 -4.481 0.0001 2200 - 2700 -1.1343 0.45 -2.521 0.1178 Colour: Red 200 - 700 0.9948 0.491 2.026 0.3276 200 - 1200 0.7372 0.517 1.426 0.7112 200 - 1700 0.0291 0.425 0.068 1 200 - 2200 0.1723 0.447 0.386 0.9989 200 - 2700 0.3619 0.467 0.776 0.9717 700 - 1200 -0.2576 0.476 -0.541 0.9945 700 - 1700 -0.9657 0.374 -2.585 0.101 700 - 2200 -0.8225 0.398 -2.064 0.3063 700 - 2700 -0.6329 0.421 -1.504 0.6619 1200 - 1700 -0.7081 0.407 -1.74 0.5053 1200 - 2200 -0.5649 0.43 -1.314 0.7775 1200 - 2700 -0.3752 0.451 -0.833 0.9615 1700 - 2200 0.1432 0.313 0.458 0.9975 1700 - 2700 0.3329 0.341 0.977 0.9255 2200 - 2700 0.1897 0.368 0.516 0.9956 Colour: Orange 200 - 700 -0.5055 0.613 -0.824 0.9631 200 - 1200 -0.4432 0.651 -0.681 0.9841 200 - 1700 -0.2566 0.594 -0.432 0.9981 200 - 2200 -0.1898 0.621 -0.306 0.9996 200 - 2700 1.5404 0.896 1.719 0.519 700 - 1200 0.0623 0.475 0.131 1 700 - 1700 0.2489 0.394 0.632 0.9887 700 - 2200 0.3158 0.433 0.73 0.9783 700 - 2700 2.046 0.778 2.631 0.0899 1200 - 1700 0.1866 0.45 0.414 0.9984 1200 - 2200 0.2534 0.485 0.523 0.9953 1200 - 2700 1.9837 0.808 2.455 0.1376 1700 - 2200 0.0669 0.406 0.165 1 1700 - 2700 1.7971 0.763 2.355 0.1722 2200 - 2700 1.7302 0.784 2.208 0.234 Colour: Yellow 200 - 700 19.14 3926.922 0.005 1 200 - 1200 1.176 1.247 0.943 0.9354 200 - 1700 -0.554 0.792 -0.699 0.9821 200 - 2200 0.977 1.023 0.954 0.9322 200 - 2700 19.048 3919.617 0.005 1 700 - 1200 -17.964 3926.922 -0.005 1 700 - 1700 -19.694 3926.922 -0.005 1 700 - 2200 -18.163 3926.922 -0.005 1 700 - 2700 -0.092 5548.343 0 1 1200 - 1700 -1.73 1.056 -1.639 0.5728 1200 - 2200 -0.199 1.239 -0.161 1 1200 - 2700 17.872 3919.617 0.005 1 1700 - 2200 1.53 0.779 1.965 0.3624 1700 - 2700 19.602 3919.617 0.005 1 2200 - 2700 18.071 3919.617 0.005 1 Colour: Green 200 - 700 -1.7381 0.777 -2.236 0.2211 200 - 1200 -0.5224 0.87 -0.6 0.9911 200 - 1700 0.0339 0.829 0.041 1 200 - 2200 0.0205 0.865 0.024 1 200 - 2700 -0.3884 0.847 -0.458 0.9975 700 - 1200 1.2157 0.541 2.246 0.2166 700 - 1700 1.772 0.471 3.761 0.0023 700 - 2200 1.7585 0.532 3.305 0.0122 700 - 2700 1.3497 0.503 2.682 0.0788 1200 - 1700 0.5563 0.613 0.908 0.9447 1200 - 2200 0.5429 0.661 0.821 0.9637 1200 - 2700 0.134 0.638 0.21 0.9999 1700 - 2200 -0.0134 0.605 -0.022 1 1700 - 2700 -0.4223 0.58 -0.728 0.9785 2200 - 2700 -0.4088 0.63 -0.649 0.9872 Colour: Blue 200 - 700 0.554 16245 0 1 200 - 1200 -20.1244 12246 -0.002 1 200 - 1700 0.9521 16048 0 1 200 - 2200 0.6282 16133 0 1 200 - 2700 0.4619 16232 0 1 700 - 1200 -20.6784 10674 -0.002 1 700 - 1700 0.3981 14883 0 1 700 - 2200 0.0743 14975 0 1 700 - 2700 -0.092 15082 0 1 1200 - 1700 21.0765 10371 0.002 1 1200 - 2200 20.7527 10502 0.002 1 1200 - 2700 20.5864 10655 0.002 1 1700 - 2200 -0.3238 14760 0 1 1700 - 2700 -0.4901 14869 0 1 2200 - 2700 -0.1663 14961 0 1 Colour: Purple 200 - 700 -16.139 1657.352 -0.01 1 200 - 1200 -17.314 1657.352 -0.01 1 200 - 1700 -15.253 1657.352 -0.009 1 200 - 2200 -16.329 1657.352 -0.01 1 200 - 2700 -18.449 1657.352 -0.011 1 700 - 1200 -1.175 0.734 -1.601 0.5978 700 - 1700 0.886 0.926 0.957 0.9312 700 - 2200 -0.19 0.782 -0.243 0.9999 700 - 2700 -2.31 0.648 -3.563 0.0049 1200 - 1700 2.061 0.836 2.465 0.1347 1200 - 2200 0.985 0.674 1.463 0.6882 1200 - 2700 -1.135 0.512 -2.214 0.2309 1700 - 2200 -1.076 0.879 -1.224 0.825 1700 - 2700 -3.196 0.762 -4.192 0.0004 2200 - 2700 -2.12 0.579 -3.66 0.0034 Colour: Pink 200 - 700 0.554 9853.381 0 1 200 - 1200 -19.552 7427.736 -0.003 1 200 - 1700 -18.253 7427.736 -0.002 1 200 - 2200 -20.725 7427.736 -0.003 1 200 - 2700 0.462 9845.471 0 1 700 - 1200 -20.106 6474.4 -0.003 1 700 - 1700 -18.807 6474.4 -0.003 1 700 - 2200 -21.279 6474.4 -0.003 1 700 - 2700 -0.092 9147.671 0 1 1200 - 1700 1.299 0.93 1.397 0.7288 1200 - 2200 -1.173 0.665 -1.763 0.4898 1200 - 2700 20.014 6462.356 0.003 1 1700 - 2200 -2.472 0.772 -3.202 0.0171 1700 - 2700 18.715 6462.356 0.003 1 2200 - 2700 21.187 6462.356 0.003 1 Colour: Brown 200 - 700 0.1442 0.894 0.161 1 200 - 1200 0.0339 0.943 0.036 1 200 - 1700 -0.9535 0.777 -1.227 0.8237 200 - 2200 0.0205 0.865 0.024 1 200 - 2700 18.0477 2377.368 0.008 1 700 - 1200 -0.1103 0.788 -0.14 1 700 - 1700 -1.0977 0.578 -1.899 0.4029 700 - 2200 -0.1238 0.692 -0.179 1 700 - 2700 17.9035 2377.368 0.008 1 1200 - 1700 -0.9874 0.652 -1.515 0.6547 1200 - 2200 -0.0134 0.754 -0.018 1 1200 - 2700 18.0138 2377.368 0.008 1 1700 - 2200 0.974 0.532 1.831 0.4454 1700 - 2700 19.0012 2377.368 0.008 1 2200 - 2700 18.0273 2377.368 0.008 1 Colour: Black 200 - 700 -0.11 0.54 -0.204 1 200 - 1200 0.519 0.63 0.823 0.9633 200 - 1700 1.138 0.584 1.948 0.3729 200 - 2200 0.904 0.603 1.499 0.665 200 - 2700 -0.308 0.538 -0.572 0.9928 700 - 1200 0.629 0.524 1.199 0.8377 700 - 1700 1.248 0.468 2.665 0.0823 700 - 2200 1.013 0.491 2.063 0.3068 700 - 2700 -0.198 0.409 -0.484 0.9967 1200 - 1700 0.619 0.57 1.086 0.8873 1200 - 2200 0.385 0.589 0.653 0.9868 1200 - 2700 -0.827 0.523 -1.581 0.6111 1700 - 2200 -0.234 0.54 -0.434 0.9981 1700 - 2700 -1.446 0.466 -3.1 0.0238 2200 - 2700 -1.212 0.49 -2.475 0.1316 Colour: White 200 - 700 1.2114 0.797 1.52 0.6513 200 - 1200 0.7985 0.802 0.996 0.9194 200 - 1700 0.7985 0.663 1.205 0.8348 200 - 2200 0.7851 0.707 1.11 0.8776 200 - 2700 1.119 0.798 1.403 0.7256 700 - 1200 -0.4128 0.839 -0.492 0.9965 700 - 1700 -0.4128 0.707 -0.584 0.9921 700 - 2200 -0.4263 0.749 -0.569 0.993 700 - 2700 -0.0924 0.835 -0.111 1 1200 - 1700 0 0.713 0 1 1200 - 2200 -0.0134 0.754 -0.018 1 1200 - 2700 0.3205 0.84 0.382 0.999 1700 - 2200 -0.0134 0.605 -0.022 1 1700 - 2700 0.3205 0.708 0.452 0.9976 2200 - 2700 0.3339 0.75 0.445 0.9978 Presentation: 200 - 700 0.055 0.472 0.116 1 Cauliflorous 200 - 1200 1.7698 0.714 2.478 0.1306 200 - 1700 0.9987 0.475 2.1 0.287 200 - 2200 1.0899 0.517 2.106 0.2839 200 - 2700 19.8174 2363.894 0.008 1 700 - 1200 1.7148 0.651 2.633 0.0894 700 - 1700 0.9437 0.374 2.522 0.1177 700 - 2200 1.0349 0.426 2.428 0.1465 700 - 2700 19.7624 2363.894 0.008 1 1200 - 1700 -0.7711 0.653 -1.18 0.8466 1200 - 2200 -0.6799 0.685 -0.993 0.9203 1200 - 2700 18.0476 2363.894 0.008 1 1700 - 2200 0.0912 0.43 0.212 0.9999 1700 - 2700 18.8187 2363.894 0.008 1 2200 - 2700 18.7275 2363.894 0.008 1 Presentation: 200 - 700 -2.1332 0.498 -4.287 0.0003 Ramiflorous 200 - 1200 -3.0406 0.592 -5.139 <.0001 200 - 1700 -2.3611 0.474 -4.982 <.0001 200 - 2200 -2.3408 0.494 -4.739 <.0001 200 - 2700 -3.6083 0.605 -5.967 <.0001 700 - 1200 -0.9075 0.482 -1.884 0.4118 700 - 1700 -0.2279 0.326 -0.699 0.9822 700 - 2200 -0.2076 0.355 -0.586 0.992 700 - 2700 -1.4752 0.497 -2.965 0.0358 1200 - 1700 0.6795 0.457 1.486 0.6731 1200 - 2200 0.6998 0.478 1.465 0.687 1200 - 2700 -0.5677 0.592 -0.96 0.9306 1700 - 2200 0.0203 0.321 0.063 1 1700 - 2700 -1.2472 0.474 -2.632 0.0897 2200 - 2700 -1.2675 0.494 -2.567 0.1055 Table A3. List of fruiting plants identified to species level and thus included in phylogenetic analyses. Numbering of species corresponds to the phylogeny as depicted in Figure 2. Table A4. Results of phylogenetic null model analysis on individual fruit colours (top) and presentation types (bottom). Standardised effect sizes (SES) were obtained for each colour and presentation type by comparing the observed mean phylogenetic distance (MPD) between species displaying the trait with a null value obtained by randomising the tips of the phylogeny 999 times. Negative SES values indicate phylogenetic clustering, while positive values indicate phylogenetic evenness. Significant results (p < 0.05) are displayed in bold. Red 16 241.5 239.73 9.28 0.19 0.54 Orange 17 243.05 239.46 8.97 0.40 0.62 Yellow 5 242.55 239.27 21.57 0.15 0.48 Green 9 232.21 240.18 13.85 -0.58 0.23 Blue 1 NA NA NA NA NA Purple 5 210.87 239.26 20.84 -1.36 0.07 Pink 3 228.09 236.6 36.73 -0.23 0.25 Brown 12 213.91 239.16 11.43 -2.21 0.03 Black 11 259.85 238.72 13.1 1.61 1 White 4 209.53 240.1 24.33 -1.26 0.11 Presentation N Observed MPD Randomised MPD Mean Randomised MPD SD Standardised Effect Size p-value Cauliflorous 13 161.48 238.92 10.88 -7.12 <0.01 Ramiflorous 63 245.65 239.50 2.52 2.44 1 Supplementary Material File (image2.emf) Download 62.51 KB Information & Authors Information Version history V1 Version 1 04 September 2025 Peer review timeline Published Ecology and Evolution Version of Record 2 Dec 2025 Published Copyright This work is licensed under a Non Exclusive No Reuse License. Collection Ecology and Evolution Keywords comparative evolutionary ecology plants terrestrial Authors Affiliations Richard Hazell 0000-0001-6347-3209 [email protected] University of Sussex View all articles by this author Graham Kaina 0000-0002-3806-8698 New Guinea Binatang Research Center View all articles by this author Katerina Sam 0000-0002-3436-0579 Biology Centre Czech Academy of Sciences View all articles by this author Daniel Souto-Vilarós The University of Utah College of Science View all articles by this author Wulan Koagouw National Research and Innovation Agency Republic of Indonesia View all articles by this author Vojtech Novotny 0000-0001-7918-8023 Biology Centre Czech Academy of Sciences View all articles by this author Mika Peck University of Sussex View all articles by this author Alan Stewart University of Sussex View all articles by this author Metrics & Citations Metrics Article Usage 425 views 258 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Richard Hazell, Graham Kaina, Katerina Sam, et al. Fruit traits reflect adaptation to dispersers along a tropical elevational gradient. Authorea . 04 September 2025. DOI: https://doi.org/10.22541/au.175697755.52347980/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu . 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