Natural Colonization Research Articles

Fish and coral communities along the seawall of Sutera Harbour Marina, Kota Kinabalu, Sabah, Malaysia

Coastal development involves the introduction of artificial substrates into the natural marine environment, thereby altering and causing the loss of natural habitat. Nevertheless, such artificial structures are known to provide novel habitat for the recruitment and growth of epifauna. Seawalls serve as hard substrata on which assemblages of benthic communities have been observed to settle and proliferate. This study investigated the diversity and abundance of marine fauna, with a primary focus on fish and hard coral communities along the seawall of Sutera Harbour Marina, Kota Kinabalu, Malaysia, in 2015 and 2022. Additionally, benthic cover was assessed in 2022. The assessment was conducted along 12 belt transects (each measuring 10 m × 3 m) placed on the seawall approximately 2–3 m from the bottom. Data on the diversity and abundance of fish and hard coral colonies were collected. Results showed that 105 fish species from 32 families, and 48 genera of hard corals from 17 families inhabited the marina. There was an increase in the abundance and colony size of hard corals over time. Additional benthic data collected in 2022 indicated that the seawalls had a fair coral cover of 31.7%. Although the taxa richness for fish and hard corals was lower compared to surrounding natural reef habitats, this study presented that artificial structures such as seawalls in the marina can support the natural colonization of marine fauna. This study underscores the importance of artificial structures as possible refuges for marine organisms, particularly in urbanized coastal areas.

Pneumococcal population dynamics: Investigating vaccine-induced changes through multiscale modelling.

The mechanisms behind vaccine-induced strain replacement in the pneumococcus remain poorly understood. There is emerging evidence that distinct pneumococcal lineages can co-colonise for significant time periods, and that novel recombinants can readily emerge during natural colonisation. Despite this, patterns of post-vaccine replacement are indicative of competition between specific lineages. Here, we develop a multiscale transmission model to investigate explicitly how within host dynamics shape observed ecological patterns, both pre- and post-vaccination. Our model framework explores competition between and within strains defined by distinct antigenic, metabolic and resistance profiles. We allow for strains to freely co-colonise and recombine within hosts, and consider how each of these types may contribute to a strain's overall fitness. Our results suggest that antigenic and resistance profiles are key drivers of post-vaccine success.

A simple genetic method to distinguish mule deer and bighorn sheep fecal pellets and its application to detecting bighorn sheep colonization events in California

Bighorn sheep (Ovis canadensis) habitat frequently is geographically discontinuous and the metapopulation model fits this species well. Consequently, extinction-colonization dynamics are important and need to be monitored. Much of the conservation history of bighorn sheep, however, was based on a theory that natural colonization is not part of the biology of this species. That theory is not supported by a growing body of evidence that natural colonization of vacant habitat does occur in this species. Here I present a simple PCR test that distinguishes bighorn sheep fecal pellets from those of mule deer (Odocoileus hemionus) as a useful tool in documenting bighorn sheep occupancy where mule deer are present. I include examples from California of applications of this method to investigate potential colonization events.

Concrete for Living Walls: Current Status and a New Design Recommendation

Concrete may be a promising material for application in living walls, broadening existing vertical greening systems and, most importantly, reducing installation costs. This study presents the concept of layered living concrete (LLC) wall panels that were developed and field-tested over the past 3 years. Simultaneously with long-term field observations, several laboratory studies on the selection of a rational concrete mix composition were carried out. Based on field data, the results of laboratory tests, and numerical simulations, a new LLC wall panel design was proposed. The new panel design retains the previous idea of a layered structure suitable for natural colonization by plants, but also adds improved material characteristics, rational dimensions, the economical use of water, and, potentially, the ability to hasten the greening of vertical surfaces.

Brazilian Clinical Strains of Actinobacillus pleuropneumoniae and Pasteurella multocida: Capsular Diversity, Antimicrobial Susceptibility (In Vitro) and Proof of Concept for Prevention of Natural Colonization by Multi-Doses Protocol of Tildipirosin.

One hundred Actinobacillus pleuropneumoniae (App) and sixty Pasteurella multocida subsp. multocida serogroup A (PmA) isolates were recovered from porcine pneumonic lungs collected from eight central or southern states of Brazil between 2014 and 2018 (App) or between 2017 and 2021 (PmA). A. pleuropneumoniae clinical isolates were typed by multiplex PCR and the most prevalent serovars were 8, 7 and 5 (43, 25% and 18%, respectively). In addition, three virulence genes were assessed in P. multocida isolates, all being positive to capA (PmA) and kmt1 genes, all negative to capD and toxA, and most of them (85%) negative to pfhA gene. The susceptibility of both pathogens to tildipirosin was investigated using a broth microdilution assay. The percentage of isolates susceptible to tildipirosin was 95% for App and 73.3% for PmA. The MIC50 values were 0.25 and 1 μg/mL and the MIC90 values were 4 and >64 μg/mL for App and PmA, respectively. Finally, a multiple-dose protocol of tildipirosin was tested in suckling piglets on a farm endemic for both pathogens. Tildipirosin was able to prevent the natural colonization of the tonsils by App and PmA and significantly (p < 0.0001) reduced the burden of Glaesserella parasuis in this tissue. In summary, our results demonstrate that: (i) tildipirosin can be included in the list of antibiotics to control outbreaks of lung disease caused by App regardless of the capsular type, and (ii) in the case of clinical strains of App and PmA that are sensitive to tildipirosin based on susceptibility testing, the use of this antibiotic in eradication programs for A. pleuropneumoniae and P. multocida can be strongly recommended.

Novel insight into the aging process of microplastics: An in-situ study in coastal wetlands

Coastal wetlands, the critical interface between the terrestrial and marine environments, provide a dynamic and unique environment for the aging of microplastics (MPs). Nevertheless, both abiotic and biotic processes that contribute to the aging of MPs in coastal wetlands have been largely neglected. In this study, the aging of MPs was continuously characterized in Hangzhou Bay, a representative coastal wetland in Zhejiang, China. Three-month exposure of polymers in sediment-water interface induced the aging phenomenon with embrittlement and exfoliation, as evidenced by simultaneous observed alternations in crystallinity and functional groups. A first-order kinetic model was fitted to describe the rate and degree of aging quantitatively. As evidenced by the carbonyl index, the residence time of all the examined MPs exhibited significant variance, ranging from 335 to 661 days. These variations might be caused by the selective attachment of plastic-degrading microorganisms (such as Moraxella sp. and Rhodococcus sp.). A positive correlation between the carbonyl index, the number of OTUs in the MP-associated biofilm, and irradiation was observed (p < 0.001), suggesting that the aging process may be co-regulated by natural sunlight and wetland microbial colonization. This study sheds new light on the long-term environmental fate of MPs and their associated ecological risks.

Modelling of drifting object dispersion in linear wave-induced flow

The dispersal of drifting objects like plant propagule under wave action can provide important information for the restoration of the plant community in coastal areas. An improved motion equation of the drift under wave action is established by coordinate transformation under the premise that the buoyancy is always perpendicular to the free surface, considering the variation of submerged height and the pressure gradient. Its numerical solution obtained by the fourth order Runge-Kutta method was compared with the experimental results in the literature, and the rationality of the improved motion equation is then verified. Then the perturbation method and Lagrange tracking principle were used to obtain the average longitudinal drift velocity expressed by Stokes drift velocity. The effect of different forces in the motion equations to the longitudinal diffusive motion was analyzed. Finally, the hydrodynamic coefficients affecting the prediction were analyzed, and the results show that the effects of the added mass coefficient and the drag coefficient are mainly reflected in the region with higher wave heights. The findings may provide certain scientific guidance for the restoration of coastal vegetation ecology through natural colonization.

Ceftiofur treatment of sows results in long-term alterations in the nasal microbiota of the offspring that can be ameliorated by inoculation of nasal colonizers

BackgroundThe nasal microbiota of the piglet is a reservoir for opportunistic pathogens that can cause polyserositis, such as Glaesserella parasuis, Mycoplasma hyorhinis or Streptococcus suis. Antibiotic treatment is a strategy to control these diseases, but it has a detrimental effect on the microbiota. We followed the piglets of 60 sows from birth to 8 weeks of age, to study the effect of ceftiofur on the nasal microbiota and the colonization by pathogens when the treatment was administered to sows or their litters. We also aimed to revert the effect of the antibiotic on the nasal microbiota by the inoculation at birth of nasal colonizers selected from healthy piglets. Nasal swabs were collected at birth, and at 7, 15, 21 and 49 days of age, and were used for pathogen detection by PCR and bacterial culture, 16S rRNA amplicon sequencing and whole shotgun metagenomics. Weights, clinical signs and production parameters were also recorded during the study.ResultsThe composition of the nasal microbiota of piglets changed over time, with a clear increment of Clostridiales at the end of nursery. The administration of ceftiofur induced an unexpected temporary increase in alpha diversity at day 7 mainly due to colonization by environmental taxa. Ceftiofur had a longer impact on the nasal microbiota of piglets when administered to their sows before farrowing than directly to them. This effect was partially reverted by the inoculation of nasal colonizers to newborn piglets and was accompanied by a reduction in the number of animals showing clinical signs (mainly lameness). Both interventions altered the colonization pattern of different strains of the above pathogens. In addition, the prevalence of resistance genes increased over time in all the groups but was significantly higher at weaning when the antibiotic was administered to the sows. Also, ceftiofur treatment induced the selection of more beta-lactams resistance genes when it was administered directly to the piglets.ConclusionsThis study shed light on the effect of the ceftiofur treatment on the piglet nasal microbiota over time and demonstrated for the first time the possibility of modifying the piglets’ nasal microbiota by inoculating natural colonizers of the upper respiratory tract.

Reconstructing the complex colonisation histories of lizards across Mediterranean archipelagos

AbstractAimThe Mediterranean Basin is a global biodiversity hotspot and has one of the longest histories of human–biota interactions. Islands host a large fraction of Mediterranean diversity and endemism, but the relative importance of natural versus human‐mediated colonisation processes in shaping the distribution and genetic structure of Mediterranean island fauna remains poorly understood. Here, we combine population genomics, demographic models and palaeoshoreline reconstructions to establish the island‐colonisation dynamics of wall lizards in Mediterranean archipelagos.LocationFour Mediterranean archipelagos in Italy and Croatia.TaxonThe wall lizard Podarcis siculus.MethodsWe used ddRAD sequencing to genotype 140 lizards from 23 island and mainland populations. Analyses of admixture and site frequency spectra were used to reconstruct population structure, demographic history and variation of gene flow through time. Genomic results were integrated with palaeogeographical reconstructions and were compared to archaeological evidence of human presence on these islands.ResultsAlthough many island populations of this species are assumed to be non‐native, we find that many islands were colonised long before any known human settlements (230,000–12,000 years ago). This natural colonisation most likely occurred through land bridges during glacial marine regression or by over‐sea rafting. On the other hand, islands distant from the continent were often colonised recently, and some of the estimated island colonisation times match historical records of human arrival. We also determine that long‐established island populations generally show lower genetic diversity compared to proximate mainland populations, contrary to recently colonised islands that must have experienced higher rates of post‐colonisation gene flow.Main ConclusionOur integrated approach provides us with the power to accurately quantify the origin, timing and mode of island colonisation. This framework helps to clarify the biogeographical and evolutionary history of island populations, with important implications for conservation and management of island biodiversity.

Natural Regrowth of Mangrove Five Years After a Large-Scale Disturbance

Haiyan, the strongest typhoon on landfall damaged mangrove covers resulting to uprooting, cutting off trunks, total defoliation and eventually death. After five years, Cancabato Bay were monitored for mangal succession and natural regrowth. Basal area was computed using diameter by breast height to estimate recovery. New species reappeared compared to the baseline study of GIZ (2014). Mangroves manifested regrowth and should be given time for natural regeneration and colonization. Natural succession patterns should be considered in human-assisted rehabilitation after disturbance.

Supercomplex systems and constitutional (dis)order: The consequences of the Anthropocene in the law1

Modern law is based in the existence of a constitutional order, guaranteeing the predictability of law. Since we are entering in new geological era, according to the contributions of Earth System Science, defined mostly by complexity and uncertainty, this hegemonic idea of order in law is challenged. This contribution explores how constitutional order was linked to the process of colonization of nature within the capitalist world system and how this process results into a planetary transformation. Discussing the Faustic expectations embedded in the some Anthropocene narratives, associated with the human manipulation of Earth System, the paper considers the profound consequences of the complexity of the intertwined ecosocial global system, and explores how supercomplexity gives way to a legal disorder which resolves itself into conflict and dialog.

Assessing the use of natural colonization to create new forests within temperate agriculturally dominated landscapes

There is a global drive to increase forest cover to protect biodiversity and help combat climate change. Tree planting is widely used to increase forest cover, although there is growing interest in using natural processes. However, predicting the outcome of natural colonization is challenging as it is a highly variable process and evidence is sparse, especially in temperate landscapes. Our study aims to evaluate the spatial and temporal scale over which natural colonization may be an effective approach to create new forests within temperate agriculturally dominated landscapes. We examine the spatial patterns in tree density and height across 90 sites in the United Kingdom that attempted to use natural colonization to create new forests between 1994 and 2004. Tree density and tree height were measured using light detecting and ranging point cloud data, an increasingly common technique for surveying forest characteristics. This research shows that natural colonization is a highly variable, relatively slow process in temperate agricultural landscapes, spatially restricted to a narrow fringe around existing forests and trees (105 m; 95% CI: 70–174 m, within approximately 19 years), although exact distances vary with former land use. This suggests that in some circumstances natural colonization will need to be assisted and supplemented with tree planting to increase forest cover, especially in areas away from a seed source. In reality, a blend of approaches will be needed to meet policy and land manager objectives, increase forest cover and tackle the biodiversity and climate crises.

Population genetics of recent natural colonization by the bull-headed shrike (Lanius bucephalus; Aves) suggests the importance of recurrent immigration on remote islands

Abstract Founder effects and recurrent immigration are two major factors that might potentially contribute to genetic differentiation and population persistence in the early stage of remote island colonization. However, their relative importance remains controversial. By conducting population genetic analyses of four remote island populations of the bull-headed shrike (Lanius bucephalus) established naturally within several decades in Japan, we examined the contributions of founder effects and recurrent immigration to these island populations. Based on the standard genetic indices and population structure analyses using 15 microsatellite loci, we suggested island-specific scenarios of colonization. Notably, the founder effect strongly influenced genetic differentiation in the population on the most remote oceanic island, Chichi-jima Island, which, however, became extinct 20 years after colonization, possibly owing to a lack of recurrent immigration. In contrast, another oceanic island, Minami-Daito Island, was probably subjected to multiple recurrent immigration events from the mainland, which obscured any genetic differentiation previously established by the founder effect. Temporal samples collected over 8 years on this island confidently supported this scenario. Underlying the island-specific scenarios of colonization, we provide evidence that recurrent immigration strongly affected the population persistence, overwhelming the initial founder effects. We argue for the importance of recurrent immigration even in colonization of highly remote islands.

Endophytic Beauveria bassiana induces biosynthesis of flavonoids in oilseed rape following both seed inoculation and natural colonization.

Cultivation of oilseed rape Brassica napus is pesticide-intensive, and alternative plant protection strategies are needed because both pesticide resistance and legislation narrow the range of effective chemical pesticides. The entomopathogenic fungus Beauveria bassiana is used as a biocontrol agent against various insect pests, but little is known about its endophytic potential and role in plant protection for oilseed rape. First, we studied whether B. bassiana can establish as an endophyte in oilseed rape, following seed inoculation. To evaluate the plant protection potential of endophytic B. bassiana on oilseed rape, we next examined its ability to induce plant metabolite biosynthesis. In another experiment, we tested the effect of seed inoculation on seedling survival in a semi-field experiment. Beauveria bassiana endophytically colonized oilseed rape following seed inoculation, and, in addition, natural colonization was also recorded. Maximum colonization rate was 40%, and generally increased with inoculation time. Seed inoculation did not affect the germination probability or growth of oilseed rape, but B. bassiana inoculated seeds germinated more slowly compared to controls. Endophytic colonization of B. bassiana induced biosynthesis of several flavonoids in oilseed rape leaves under controlled conditions. In the experiment conducted in semi-field conditions, inoculated seedlings had slightly higher mortality compared to control seedlings. Beauveria bassiana showed endophytic potential on oilseed rape via both natural colonization and seed inoculation, and it induced the biosynthesis of flavonoids. However, its use as an endophyte for plant protection against pests or pathogens for oilseed rape remains unclear. © 2023 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Effects of root plates, pits and mounds following windthrow events on the biodiversity of plants and lichens in Central European mountain forests

Windthrows lead to the formation of a characteristic land microtopography and they provide new habitats and substrata for species. The present study is novel, because it concerns different taxonomic groups of organisms and effects of various variables on species composition, richness and abundance within root plate-pit-mound (RPM) complexes. For the first time we studied lichens, liverworts, mosses and vascular plants on RPM complexes across major vegetation types, from deciduous to coniferous forests. We analyzed 164 RPM complexes which included in total 268 components: 115 root plates, 80 pits and 73 mounds. Besides the vegetation types and component types the following variables were also considered: dimensions (area, size), age (decomposition degree), species of the uprooted tree, shade, and elevation. The research revealed that root plates, pits and mounds had their own indicator species, but the key factors for the species composition were the species of uprooted trees and the vegetation type. Furthermore, the type of component and the vegetation type were also the most main variables determining the richness and abundance of the studied taxonomic groups of species. Analyses revealed the greatest species richness and abundance of lichens on root plates, and the highest richness of liverworts and vascular plants in pits. The type of component had no effect on the species richness of mosses. However, they were most abundant on mounds. The response of individual groups to the vegetation types and other variables varied. Findings from the study indicated that the presence of uprooted trees with different features (size and age, species of uprooted tree) is important for the biodiversity of mountain forests. We recommend leaving uprooted trees in forest communities for natural colonization, preferably large objects. Moreover, in managed forests a mosaic of forest communities that mimic the structure of natural unmanaged forests should be supported.

Immune responses following neonatal vaccination with conserved F4 fragment of VtaA proteins from virulent Glaesserella parasuis adjuvanted with CAF®01 or CDA

Glaesserella parasuis is a Gram-negative bacterium that colonizes the upper airways of swine, capable of causing a systemic infection called Glässer’s disease. This disease is more frequent in young post-weaning piglets. Current treatments against G. parasuis infection are based on the use of antimicrobials or inactivated vaccines, which promote limited cross-protection against different serovars. For this reason, there is an interest in developing novel subunit vaccines with the capacity to confer effective protection against different virulent strains. Herein, we characterize the immunogenicity and the potential benefits of neonatal immunization with two different vaccine formulations based on the F4 polypeptide, a conserved immunogenic protein fragment from the virulence-associated trimeric autotransporters of virulent G. parasuis strains. With this purpose, we immunized two groups of piglets with F4 combined with cationic adjuvant CAF®01 or cyclic dinucleotide CDA. Piglets immunized with a commercial bacterin and non-immunized animals served as control groups. The vaccinated piglets received two doses of vaccine, at 14 days old and 21 days later. The immune response induced against the F4 polypeptide varied depending on the adjuvant used. Piglets vaccinated with the F4+CDA vaccine developed specific anti-F4 IgGs, biased towards the induction of IgG1 responses, whereas no anti-F4 IgGs were de novo induced after immunization with the CAF®01 vaccine. Piglets immunized with both formulations displayed balanced memory T-cell responses, evidenced upon in vitro re-stimulation of peripheral blood mononuclear cells with F4. Interestingly, pigs immunized with F4+CAF®01 controlled more efficiently a natural nasal colonization by a virulent serovar 4 G. parasuis that spontaneously occurred during the experimental procedure. According to the results, the immunogenicity and the protection afforded by F4 depend on the adjuvant used. F4 may represent a candidate to consider for a Glässer‘s disease vaccine and could contribute to a better understanding of the mechanisms involved in protection against virulent G. parasuis colonization.

Squamates, rodents, and birds from Holocene deposits of the Illa Grossa Island (Columbretes Islands, Castellón, Spain): an unexpected diverse assemblage

The Columbretes Islands (eastern Spain) comprise a volcanic archipelago 50 km off the eastern Spanish coast. Illa Grossa is the main island of the archipelago. After the settlement of humans during the mid-19th century, strong modifications in biodiversity took place, including the reduction of the non-flying vertebrate fauna to a single taxon, Podarcis liolepis atratus (Boscá, 1916). The study of the fossils yielded at the COLT site (dated by AMC between 2780-2724 cal BP and 2600-2492 cal BP) from a loess deposit located at the south of Illa Grossa Island shows that this fauna was more diverse in the past. The assemblage was dominated by squamate reptiles, belonging to two lizards (Chalcides bedriagai (Boscá, 1880) and cf. Podarcis Wagler, 1830) and one snake (Vipera cf. latastei Boscá, 1878). The record in the late Holocene of Columbretes of Vipera cf. latastei settles the existing controversy about the identity of snakes in the archipelago. Regarding Chalcides bedriagai, this is the first record of its presence in the Columbretes Islands. Other faunal elements recovered from the site include at least six species of undetermined birds (passeriforms and procellariforms) and the anthropophilic Mus cf. musculus Linnaeus, 1758. The origin of the squamates taxa from Columbretes palaeontological site (COLT) is still unclear, but was probably related to the Late Pleistocene-Greenlandian (11 700 to 8300 BP, Early Holocene) sea transgression, natural passive colonization, and/or human introduction. We hypothesize implications for the conservation and management of the Natural Reserve.

Drivers of tree colonization, species richness, and structural variation during the initial three decades of natural forest colonization in abandoned agricultural soils

Natural colonization of abandoned agricultural soils has been proposed as a cost-effective strategy for simultaneously mitigating the climate crisis, restoring ecological integrity, and promoting the re-establishment of native biodiversity. The success and speed at which forests develop in the presence of land-use legacies are highly variable, and empirical knowledge on the drivers of the initial phase of temperate forest colonization in a range of site conditions is lacking. We analyzed and compared drivers of the initial three decades of natural forest colonization in 33 afforestation sites laid out between 1990 and 2018, in Denmark. We analyzed how the age of colonization, size of the area, soil type, topography, and abundance of neighboring woody vegetation influenced woody vegetation cover, species richness, and variation in vegetation structure of the colonizing woody species. We found that woody vegetation cover, species richness, and structural variation increased significantly with time since abandonment. Woody vegetation cover was significantly higher in sites with abundant neighboring vegetation and on loamy soils compared to sand or clay. Woody vegetation cover tended to be higher in sites with sloping terrain. Species richness increased significantly in sites with loamy or clay soil and sites with sloping terrain. After accounting for the other drivers, none of the elements were influenced by the size of the area. The results suggest that time since abandonment, neighboring mature woody vegetation, soil and topography are key drivers of the different elements of colonization. This knowledge can be used to prioritize areas for natural colonization and to improve active afforestation methods, especially in new forests where the main focus is on nature quality and biodiversity.

The interacting influences of competition, composition and diversity determine successional community change

Abstract Community change is one of the few constants in nature, and the balance of mechanisms influencing this change is central for understanding the structure and functioning of communities and ecosystems. Newly established communities undergo succession and can change in diversity and composition as local environmental change, interspecific interactions and immigration play out over time. Understanding the influence of initial conditions and priority effects (long‐term consequences of the initial community composition and species identity) on community change is critically important for both evaluating ecological theory and predicting restoration outcomes. Here I evaluate how initial experimental conditions in 2012, like initial sown species richness, phylogenetic diversity and early biomass production, along with priority effects caused by the identity of sown species, influence subsequent herbaceous plant community composition and the number of colonizing species after 9 years of uninterrupted natural colonization. I found that sown phylogenetic diversity (measured as mean pairwise distance) indirectly affected the number of colonizing species by increasing biomass production early on and plots with more biomass in 2012 were colonized by fewer species. Individual species influenced the number of colonizing species with taller species reducing the number of colonists and some shorter species subject to a greater number of colonists. Synthesis. Taken together, these results indicate that initial composition influences the number of colonizing species via community‐wide competition. These findings suggest that restoration outcomes can be greatly influenced by decisions about sown species composition and early management practices.

The turnover of plant-frugivore interactions along plant range expansion: consequences for natural colonization processes.

Plant-animal mutualisms such as seed dispersal are key interactions for sustaining plant range shifts. It remains elusive whether the organization of interactions with seed dispersers is reconfigured along the expansion landscape template and, if so, whether its effects accelerate or slow colonization. Here we analyse plant-frugivore interactions in a scenario of rapid population expansion of a Mediterranean juniper. We combined network analyses with field surveys, sampling interactions between individual plants and frugivores by DNA-barcoding and phototrapping over two seasons. We assess the role of intrinsic and extrinsic intraspecific variability in shaping interactions and we estimate the individual plant contributions to the seed rain. The whole interaction network was highly structured, with a distinct set of modules including individual plants and frugivore species arranged concordantly along the expansion gradient. The modular configuration was partially shaped by individual neighbourhood context (density and fecundity) and phenotypic traits (cone size). Interaction reconfiguration resulted in a higher and more uneven propagule contribution, with most effective dispersers having a prominent role at the colonization front stand, where a distinct subset of early arriving plants dominated the seed rain. Our study offers new insights into the key role of mutualistic interactions in colonization scenarios by promoting fast plant expansion processes.