Abstract. Ecological interaction networks are a valuable approach to understanding plant-pollinator interactions at the community level. Highly structured daily activity patterns are a feature of the biology of many flower visitors, particularly provisioning female bees, which often visit different floral sources at different times. Such temporal structure implies that presence/absence and relative abundance of specific flower-visitor interactions (links) in interaction networks may be highly sensitive to the daily timing of data collection. Further, relative timing of interactions is central to their possible role in competition or facilitation of seed set among coflowering plants sharing pollinators. To date, however, no study has examined the network impacts of daily temporal variation in visitor activity at a community scale. Here we use temporally structured sampling to examine the consequences of daily activity patterns upon network properties using fully quantified flower-visitor interaction data for a Kenyan savanna habitat. Interactions were sampled at four sequential three-hour time intervals between 06:00 and 18:00, across multiple seasonal time points for two sampling sites. In all data sets the richness and relative abundance of links depended critically on when during the day visitation was observed. Permutation-based null modeling revealed significant temporal structure across daily time intervals at three of the four seasonal time points, driven primarily by patterns in bee activity. This sensitivity of network structure shows the need to consider daily time in network sampling design, both to maximize the probability of sampling links relevant to plant reproductive success and to facilitate appropriate interpretation of interspecific relationships. Our data also suggest that daily structuring at a community level could reduce indirect competitive interactions when coflowering plants share pollinators, as is commonly observed during flowering in highly seasonal habitats.
Ant-plant protection symbioses, in which plants provide food and/or shelter for ants in exchange for protection from herbivory, are model systems for understanding the ecology of mutualism. While interactions between ants, host plants, and herbivores have been intensively studied, we know little about how plant-plant interactions influence the dynamics of these mutualisms, despite strong evidence that plants compete for resources, that hosting ants can be costly, and that host-plant provisioning to ants can therefore be constrained by resource availability. We used field experiments in a semiarid Kenyan savanna to examine interactions between the ant-plant Acacia drepanolobium, neighboring grasses, and two species of symbiotic acacia-ants with divergent behaviors: Crematogaster mimosae, an aggressive symbiont that imposes high costs to host trees via consumption of extrafloral nectar, and Tetraponera penzigi, a less-protective symbiont that imposes lower costs because it does not consume nectar. We hypothesized that by competing with acacias for resources, neighboring grasses (1) reduce hosts' ability to support costly C. mimosae, while having little or no effect on the ability of hosts to support low-cost T. penzigi, and (2) reduce sapling growth rates irrespective of ant occupant. We factorially manipulated the presence/absence of grasses and the identity of ant occupants on saplings and evaluated effects on colony survivorship and sapling growth rates over 40 weeks. Contrary to prediction, the high-cost/high-reward nectar-dependent mutualist C. mimosae had higher colony-survival rates on saplings with grass neighbors present. Grasses appear to have indirectly facilitated the survival of C. mimosae by reducing water stress on host plants; soils under saplings shaded by grasses had higher moisture content, and these saplings produced more active nectaries than grass-removal saplings. Consistent with prediction, survival of low-cost/low-reward T. penzigi did not differ significantly between grass-removal treatments. Saplings occupied by low-cost/low-reward T. penzigi grew 100% more on average than saplings occupied by high-cost/high-reward C. mimosae, demonstrating that mutualist-partner identity strongly and differentially influences demographic rates of young plants. In contrast, contrary to prediction, grass neighbors had no significant net impact on sapling growth rates. Our results suggest that neighboring plants can exert strong and counterintuitive effects on ant-plant protection symbioses, highlighting the need to integrate plant-plant interactions into our understanding of these mutualisms.
Experimental studies assessing the impact of demographic changes on aggression and inter‐group competitive outcomes in communities of social species are rare. This gap in our knowledge is important, not only because social species are foundational elements of many terrestrial ecosystems, but because interference competition among social groups often involves decision‐like processes influenced by demographic and environmental contexts. In East Africa, the symbiotic ant Crematogaster mimosae is a co‐dominant competitor that engages in high‐mortality, intra‐ and interspecific battles for sole possession of host trees. We manipulated worker density on C. mimosae Acacia host trees, and the colony's opportunity to expand onto neighboring trees to identify how these factors influenced colony‐level aggression, expansion success, and longer‐term survivorship. Worker density on focal trees was increased through translocation of domatia‐bearing branches, and was decreased using partial tree fumigations. We examined impacts of density manipulations on aggression and immediate expansion success under two different risk scenarios. We tied focal trees to either an experimentally emptied‐tree (low‐risk treatment), or to a C. nigriceps‐occupied tree (high‐risk treatment). Expansion success onto emptied neighbor trees was 100% for controls and increased‐density colonies, but only 50% for decreased‐density colonies, despite the fact that host trees are a limiting resource in this system. Success expanding onto trees occupied by a heterospecific competitor reached 36%, 40%, and 79% in decreased, control, and increased‐density trees, respectively. Our results show that changes in worker density due to disturbances or inter‐group battles have the potential to disrupt competitive hierarchies. Worker density manipulations also affected longer‐term colony persistence. Behavioral and genetic data revealed that 12 months after expansions 100% of the decreased‐density colonies, and 25% of control and increased‐density colonies, had been supplanted by neighboring opportunistic conspecifics. Intraspecific aggression may have lower costs in C. mimosae because aggressive colonies can incorporate workers or queens from defeated competitors. The unexpectedly high frequency of conflicts between conspecific C. mimosae, in combination with behaviors decreasing the cost of intraspecific competition relative to interspecific conflict, may create opportunities for the survival of subordinate competitors in this ant–plant system.
We isolated 11 polymorphic microsatellite markers from Acacia mellifera, a savannah woodland tree in sub-Saharan Africa and southern Arabia. The loci were screened for polymorphism using 48 Kenyan individuals. Allelic diversity ranged from three to 19 per locus and the polymorphic information content varied from 0.287 to 0.893. These loci will be useful in studies of genetic structure, gene flow and breeding systems.
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