Positive biotic interactions such as mutualism, commensalism and facilitation are ubiquitous in nature, but historically have received considerably less research attention than negative interactions such as competition, predation and parasitism. The paucity of research on positive interactions is particularly evident in stream ecosystems and in vertebrate communities. Stream fishes clearly provide an ideal system for advancing research on positive interactions. Many minnows (Cyprinidae) of eastern North America engage in a potentially mutualistic reproductive interaction known as nest association, in which individuals of one species (nest associates) spawn in nests constructed by host species. In nest association, hosts provide unsilted gravel substrate for spawning nest associates, and increased parental care to associate broods. High associate: host egg ratios can create a dilution effect, reducing the probability that host eggs will be preyed upon by egg predators. Nest associative interactions are common, but are relatively understudied compared to other interactions among stream fishes.

The goals of this study were to apply general ecological models to this novel system to (a) gain new insight into the mechanisms structuring nest associative stream fish communities, and (b) to use inference from stream fish communities to potentially expand and improve the general ecological models. These goals required completion of three objectives, including (1) examining the influence of abiotic and biotic contexts on reproductive behavior and fitness outcomes between a cyprinid host and associate, using the biological markets model to generate predictions; (2) examining the utility of the nest web framework (previously only used for cavity nesting vertebrate communities) and the stress gradient hypothesis (previously applied almost exclusively to plant communities) for predicting which associate species spawn on nests built by various nest building species, and the consequences of these choices, respectively; and (3) using two-species occupancy modeling to determine the relative influence of biotic interactions and habitat covariates on the co-occurrence of a host and two nest associates.

To accomplish these goals, I conducted a large-scale experiment to manipulate presence of mutualists (Nocomis leptocephalus, host; Chrosomus oreas, associate), egg predators (biotic context) and habitat quality (abiotic context). I conducted behavioral nest observations and conducted repeated stream fish stream fish community surveys to collect demographic data. I constructed a nest web from observational data, and implemented structural equation modeling through an information-theoretic framework to identify nest web plausibility across a large spatial extent. I tested some predictions of the stress gradient hypothesis by regressing juveniles-per-nest and a metric of cyprinid community structure on a composite measure of physical stress (scaled gradients of catchment-scale agricultural land use and catchment area). I used two-species occupancy modeling to model co-occurrence of N. leptocephalus hosts and two associates, C. oreas and Clinostomus funduloides, and used an information-theoretic framework to compare hypotheses representing the importance of biotic interactions, habitat covariates or both at determining species co-occurrence.

Results corroborated some (but not all) model predictions, and identified room for improvement in each of the general models. Nest associative spawning by C. oreas was not context dependent; C. oreas did not spawn in the absence of a reproductively active male N. leptocephalus at any treatment level. However, the net fitness outcome of host and associate species was mutualistic, and the interaction outcome switched from commensalistic to mutualistic with abiotic context. N. leptocephalus reproductive success was improved by C. oreas presence in less-silted habitats, but not in heavily-silted habitats. This is most likely because broods were subject to predation in both habitat types, but were also negatively affected by siltation in silted habitats. Accordingly, egg dilution by associates was not sufficient to support a mutualistic relationship in less favorable habitats. Results suggest that the biological markets model may be a useful tool for predicting fitness outcomes of nest associative mutualism, but may not be as useful for predicting the behavioral outcomes of obligate mutualisms. Future applications of the biological markets model should carefully consider species traits, specifically the degree to which trading behavior is obligate for participants. Future work with this model will yield more insight by considering highly facultative associates.

Nest webs constructed from nest observational data suggested an interaction topology in which strong (nearly-obligate) associates relied most frequently on N. leptocephalus nests, and less frequently on nests constructed by Campostoma anomalum. Weak (facultative) associates were seldom associated with nests constructed by either species, and probably spawned before hosts began nesting activity. Structural equation models corroborated this topology throughout the New River basin, although some less-supported model evidence specified some nest association by weak associates. Juveniles-per-nest of strong associates responded positively to physical stress, while this metric for other cyprinid reproductive groups showed no relationship. Proportional representation of Nocomis and strong associates also increased predictably with physical stress. This study suggests that the nest web framework can be informative to systems outside the ones for which it was developed; future studies may be able to use this framework to better understand the role of habitat-modifying species in communities other than cavity nesting terrestrial vertebrates and nest associative stream fishes. This work extended the nest web framework by (a) modeling the outcomes of interactions instead of the interactions themselves, and (b) by using structural equation modeling to test nest web predictions with an information-theoretic framework. This study also suggests that the stress gradient hypothesis can be useful for understanding interaction dynamics in vertebrate communities; this represents the first direct evidence that this model can be used in vertebrate communities. Further, I demonstrate that the stress gradient hypothesis may be extended to predict community structure. However, more research in a diversity of systems will be needed to determine the extent to which this can be applied.

This study provides some of the first evidence of large-scale positive co-occurrence patterns in vertebrates. However, the precise roles of habitat covariates and biotic interactions were species-specific. Occupancy results suggest that co-occurrence between N. leptocephalus and nest associate C. funduloides is driven only by reproductive behavioral interactions. Alternatively, evidence suggests that co-occurrence between N. leptocephalus and C. oreas is driven by both nest association and habitat covariates. That two-species occupancy modeling can be a useful tool for comparing difficult-to-test hypotheses involving biotic interactions at large spatial scales. This study represents the first quantitative, multi-scale treatment of positive interactions in stream ecosystems.

This study demonstrates that applying general ecological models to stream fish communities can yield new insights about both the study system and the models themselves. While models of negative interactions, food webs and dispersal have been applied to stream fishes, we stand to gain much ground by also considering positive biotic interactions. In doing so, stream fish ecologists will also be able to contribute to the advancement of general ecology, and thus raise awareness for these understudied ecosystems and taxa.