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The strength of Nocomis nest association contributes to patterns of rarity and commonness among New River, Virginia cyprinids.


Freshwater fishes exhibit diverse modes of reproduction, life history strategies and habitat requirements (Balon, 1975; Winemiller and Rose, 1992). Nest association represents one unique reproductive mode. This occurs when individuals of one species (the associate) use nests made by other species (the host) for reproduction (Johnston and Page, 1992). In North America, nest associates typically belong to the family Cyprinidae (36 known species; Johnston, 1999) and use nests constructed by other cyprinids or centrarchids (Johnston and Page, 1992). Probably the most common nest associations in North America occur between chubs (Nocomis spp.) and several other genera of minnows. Fishes use nest association to increase brood survival (Wisenden, 1999), as nest associates benefit by receiving parental care (Johnston, 1994a; Shao, 1997) and better substrate conditions (Cooper, 1980; Vives, 1990; Fletcher, 1993). Additionally, hosts can receive decreased probability of egg predation through a dilution effect (McKaye and McKaye, 1977; Johnston, 1994b). Because both hosts and associates may benefit from the interaction, nest association may reflect a mutualistic relationship (Johnston, 1994b) that strongly influences stream fish community structure (Walser et al., 2000; Hitt and Roberts, 2011).

Most nest associates do not solely rely on nest association for reproduction. Facultative associates often revert to alternate, more ancestral spawning modes. Predominantly, these species are broadcast open substratum lithophils (i.e., requiring clean gravel substrate, sensu Balon, 1975). However, some associates can construct their own nests (e.g., Campostoma and Luxilus spp.) or spawn in natural crevices (e.g., red shiner Cyprinella lutrensis). Although well documented, the degree to which most associates must use nest association with Nocomis remains unknown. Some species represent obligate nest associates whose reproductive success heavily depends on the spawning activities of the host (Wallin, 1992; Black, 2007). Although many facultative associates probably occur (Johnston and Page, 1992), little attention has been given to differences in nest association strength among species, or the implications of these differences for population dynamics. Grouping species based on nest association strength could provide a framework for understanding the importance of this interaction.

Species traits provide an ideal basis for grouping fishes to simplify the process of testing ecological hypotheses (Frimpong and Angermeier, 2010). Ecological, biological, and ethological traits have been used to delineate groups of fish easily relatable to environmental requirements (Balon, 1975; Growns, 2004). Ethological and ecological traits may be important to quantifying the strength of nest association with Nocomis. For example, traits such as spawning substrate preference, the ability to construct nests, and the provision of parental care provide information about a species' propensity to utilize nest association or alternate spawning modes. By carefully selecting behavioral traits from independent studies, groups or gradients of nest association strength may be delineated among species.

In general, reproductive modes have been shown to influence rarity (or commonness), extinction proneness (Kunin, 1997) and patterns of imperilment of freshwater fishes in particular (Johnston, 1999). However, the current paradigm of conservation largely centers on single species, and little attention goes to interspecific interactions aside from competition and predation. In the broader field of applied ecology, conservation efforts increasingly recognize mutualisms as a priority (Bronstein, 2009; Kiers et al., 2010). Because of its strong influence on the reproductive success of both hosts and associates, nest association provides a lens through which the importance of interspecific interactions to the conservation of North American cyprinids can be examined.

Rarity, in spite of its importance in conservation biology, is an attribute of species that is complex to define. Rabinowitz (1981) proposed that species be considered rare based on three dimensions: geographic range, habitat breadth, and local abundance. In this framework, a species that exhibits at least one form of rarity is considered rare. Models that include the facilitative effects of mutualism into niche theory predict that the distribution and habitat requirements (e.g., spawning temperature) of obligate nest associates must be nested within those of the host (Bruno et al., 2003). For instance, a species that fully relies on Nocomis nests for spawning should not maintain reproducing populations in systems where Nocomis is absent. Additionally, abundances of associates should be directly linked to that of their host (Wallin, 1992; Walser et al., 2000; Hitt and Roberts, 2011). Because the distributional and habitat requirements of obligate nest associates should be more closely related to that of their host, obligate nest associates should be more likely to exhibit rarity (by distribution and habitat breadth) than weak associates.

This study sought to: (1) develop and use a methodology for identifying the strength of nest association (with Nocomis) among 11 cyprinids occurring in the New River basin, Virginia; (2) evaluate the groups based on geographic and spawning temperature range overlap with Nocomis; and (3) identify the relationship between the strength of nest association and rarity. We hypothesized that the geographic distributions and spawning temperature ranges of strong nest associates significantly overlap with that of Nocomis, whereas those of weak associates do not. We further hypothesized that strong nest associates exhibit rarity along at least two dimensions (distribution and habitat breadth) and that weaker nest associates are relatively common. We did not expect strong nest associates of Nocomis to exhibit rarity based on local abundance because the benefits of nest association are widely reported to confer high local abundances on associates (Walser et al., 2000; Hitt and Roberts, 2011).


To achieve our objectives, we needed to delineate groups of species based on their tendency to utilize nest association. To do so, we synthesized life-history information into matrices of ecological and ethological trait similarity. We then used ordination techniques to identify species groups and correlated traits to ordination axes to interpret groupings. We evaluated the groups for their dependence on Nocomis by comparing spawning temperature and range overlap. These groupings served as the basis for relating spawning modes to rarity.


This study focuses on the middle New River basin of southwestern Virginia. The system represents an ideal location for studying nest association because many of the cyprinids occurring in the basin utilize nest association as a reproductive mode. The 28 cyprinids inhabiting the New River basin in Virginia include 11 documented associates of Nocomis. These include central stoneroller Campostoma anomalum, mountain redbelly dace Chrosomus oreas, rosyside dace Clinostomus funduloides, crescent shiner Luxilus cerasinus, white shiner L. albeolus, rosefin shiner Lythrurus ardens, swallowtail shiner Notropis procne, rosyface shiner N. rubellus, saffron shiner N. rubricroceus, blacknose dace Rhinichthys atratulus and longnose dace R. cataractae. Additionally, the basin-wide ubiquity of two Nocomis species (hosts), bluehead chub N. leptocephalus (abundant in tributaries) and bigmouth chub N. platyrynchus (abundant in mainstem and large tributaries), make nest association possible throughout the drainage (Jenkins and Burkhead, 1994).


We performed an extensive literature review to obtain ecological and ethological reproductive trait information for the species listed above. First, we searched secondary literature (species account books) throughout the geographic range of each species listed above for unique records of reproductive traits. Next, we reviewed each primary study cited in these secondary sources. To ensure no studies were overlooked, we performed an internet-based literature search using the ISI Web of Knowledge database. Searches included year (Jan. 1969 through Sep. 2011), and search terms included each species' scientific and common names. We included former scientific names for recently renamed species (e.g., Phoxinus and Chrosomus oreas, and Luxilus and Notropis albeolus).

Selected traits provided insight into the breadth of spawning modes the substrate use. We grouped behavioral spawning traits into three types: broadcasting eggs over open substrate (if so, substrate type is indicated) or in natural depressions, spawning over conspecific nests/constructing own nests, and spawning on nests constructed by other species (nest association). Reports of nest association were classified by host genus (Nocomis or other). We also collected information on the provision of parental care to brood because we expected this trait to be inversely related to nest association strength. We identified geographic ranges of all 11 focal nest associate species, as well as all Nocomis species (some associates occur within the ranges of multiple Nocomis species). We compared the location of each study with range overlap with Nocomis and classified it as "sympatric" if it occurred within the range of Nocomis, or otherwise "allopatric." This approach allowed us to identify 10 unique behavioral reproductive traits (Table 1).

To create a matrix of reproductive traits, we calculated the proportion of studies in which a specific trait was observed. We used binary coding to describe reproductive traits as either present (score of "1") or absent (score of "0") for each independent published record of each species. We then divided the sum of all observations of a trait by the number of studies we gathered for that species to calculate a proportion. Because some studies report multiple spawning modes for the same species [e.g., Jenkins and Burkhead (1994) report both broadcast spawning and nest association for mountain redbelly dace], it was not a requirement that observed trait frequencies sum to 1.0 for any given species (Table 2).


Any comparative trait analysis can be confounded by phylogenetic relatedness among species. Due to common ancestry and limited time for evolutionary divergence, closely related species inherently share traits. As a result, trait similarities depend upon phylogenetic distance. This effect often produces misleadingly similar relationships among species and must be accounted for (Fisher and Owens, 2004). To account phylogenetic relatedness among species, we used phylogenetic eigenvector regression (PVR; Diniz-Filho et al., 1998). We chose to use PVR because it accommodates variability among data types and better handles small sample sizes (Diniz-Filho et al., 1998; Olden et al., 2008) than other methods such as independent contrasts (Felsenstein, 1985).

To perform PVR, we first required a phylogeny. Selecting an appropriate phylogeny can be contentious because Cypriniformes represents one of the most complex teleost orders (Mayden et al., 2008; Britz and Conway, 2011), and phylogenetic hypotheses of Cyprinidae continue to be updated (Johnston and Page, 1992; Simons et al., 2003; Gaubert et al., 2009). We sought the most recent phylogeny that includes all focal genera of this study, and thus chose the supertree hypothesis proposed by Gaubert et al. (2009). This phylogeny mostly delineates genera but does distinguish a few species, specifically rosyface shiner in this analysis (Gaubert et al., 2009). Most of the remaining Notropis separate into three groups (Notropis 1, 2, and 3, each representing divergent suites of species). We did not assign other Notropis species to these groups because the phylogenetic topology does not distinguish evolutionary distance among them. Therefore, all distances among Notropis species used in this study (other than rosyface shiner) were equal.

We created a matrix of phylogenetic relatedness by enumerating the number of nodes separating each pair genera. Congeners (excluding rosyface shiner from other Notropis) received zero values. We then subtracted each distance value from the largest distance value and added one to prevent zero values. This created a matrix of phylogenetic similarity in which the most closely related species displayed highest values (Table 3). When performing PVR, the use of actual genetic distances estimated by molecular techniques is ideal. However, a uniform measure of actual genetic distances among cyprinid genera is unavailable, and the node counting method is an appropriate approximation of distance (Olden et al., 2008).

To calculate a matrix of phylogenetically independent traits, we first ordinated the phylogenetic similarity matrix using nonmetric multidimensional scaling (NMS; Legendre and Legendre, 1998). We then extracted eigenvectors from the first two dimensions. We used these eigenvectors as explanatory variables in multiple linear regression models for each trait. Residuals from these models represented variability in traits that could not be explained by phylogenetic relatedness among species. To identify groups of species that represent nest association strength, we first calculated a matrix of trait similarity among species based on the phylogenetically independent traits. We used Gower's index of similarity, which suits this dataset better than the more commonly used Jaccard's index because it can incorporate negative residual values into similarity calculations (Zar, 1999).

We then ordinated the phylogenetically independent trait similarity matrix using NMS, extracted eigenvectors from the first two dimensions, and plotted the data to examine the location of species and potential groupings. To interpret trait relationships to ordination axes, we calculated Pearson correlation between each raw trait vector and the two ordination axes. Significant correlations (P < 0.10) were used to interpret axes. Groupings based on nest association strength were initially screened by visual interpretations of the ordination plot. To avoid bias in visual interpretations, we numerically ranked species by their score on eigenvectors on both axes and averaged the two rankings. All statistical analyses were performed in SAS 9.2.


We evaluated group dependence on Nocomis by analyzing spawning temperature and geographic range overlap of each nest associate with Nocomis under the hypothesis that strong nest associates should have greater geographic and spawning temperature overlap with Nocomis than weak associates. We calculated each species' geographic range throughout North America using maps in Page and Burr (1991). These maps were digitized and georeferenced in ArcGIS 9.2, as reported in detail in Frimpong and Angermeier (2009). Although more detailed maps may be available (e.g., those published by NatureServe), they provide no better resolution than those in Page and Burr (1991) at the spatial scale of interest. A union of ranges for all Nocomis species was created from digital maps. We calculated the area of each species' range, which was overlaid with the range of Nocomis to obtain the area of intersection. This same process was performed to estimate spawning temperature overlap. We calculated range and temperature overlap using the equation:

proportion contained = (A [intersection] B)/B (1)

where A = geographic or temperature range of Nocomis, and B = area or temperature range of nest associate. We used a one tailed t-test ([alpha] = 0.05) to test for significant differences in range overlap with Nocomis between the strong and weak nest associates.


To examine the relationship between nest association strength and rarity, we used Rabinowitz's (1981) definition of rarity as implemented by Pritt and Frimpong (2010) for fishes of the United States. In their classification, Pritt and Frimpong (2010) distinguished 399 species of freshwater fish as rare or common by (1) geographic extent, based on Page and Burr (1991) range maps, (2) habitat specificity, based on documented habitat associations (Frimpong and Angermeier, 2009), and (3) local abundance, based on U.S. Geological Survey and U.S. Environmental Protection Agency national fish monitoring databases. The classification results in eight groups of species, one of which displays commonness across all three dimensions of rarity and seven groups that display rarity in one or more dimensions. Species classified as rare on any dimension met the definition of rare, and those that did not exhibit rarity on any dimension met the definition of common. We tested for a relationship between nest association strength and rarity by performing a 2 x 2 cross classification of species' nest association strength and rare/common status using Fisher's exact test (Zar, 1999), with a null hypothesis of independence.


We identified 40 original references that reported spawning mode and/or spawning temperatures. The number of studies per species ranged from 2 (white shiner) to 17 (rosyface shiner) with a median of 6 (Appendix 1). Two nest associates, crescent shiner and white shiner, spawned only on Nocomis' nests, whereas the remaining eight nest associates use alternative spawning modes (Table 2).

Ordinations of trait similarities produced easily interpretable groupings. Most axis scores were correlated with traits associated with weak nest association (with Nocomis) strength, producing a tightly clustered group of strong nest associates in the center of the plot, near zero on both axes. Easily distinguishable from the strong group, weaker nest associates were located on the outer fringes of the plot, driven there by various traits associated with independent spawning (Fig. la).

Traits that correlated strongest and positively with dimension 1 of the multidimensional scaling included building nest in sympatry with Nocomis (BNS) and building nest in allopatry (BNA) (r = 0.67 for both) and association with Nocomis in sympatry (NNS) (r = 0.58). Negative correlation with dimension 1 include broadcast spawning in sympatry with Nocomis (BS) (r = -0.60), spawning on sand (SAND) (r = -0.55), spawning in natural depressions in allopatry (NDA) (r = -0.53), and broadcast spawning in allopatry (BA) (r = -0.53). Therefore on dimension 1, the ability to build nest (whether in sympatry and or allopatry) and broadcast spawning (often associated with sand substrates) drove species in opposite directions, away from the center of the plot. Traits that correlated strongest with dimension 2 include nest association with other species' nest in sympatry with Nocomis (NOS) (r = -0.62), BS (r = -0.54), and NDA (r = 0.59) (Table 4; Fig. 1a). Thus, on dimension 2, association with other species' nests in sympatry with Nocomis or broadcast spawning and spawning in natural depressions in allopatry also drove species in opposite directions, away from the center. These patterns of correlation support the interpretation that the strongest Nocomis nest associates occur near the center of the plot.

Strong associates included rosyside dace, rosefin shiner, mountain redbelly dace, crescent shiner, and white shiner (Fig. 1a). Weak associates included longnose dace, blacknose dace, spottail shiner, and central stoneroller (Fig. la). Rosyface shiner and saffron shiner occurred on the upper and lower fringes of the strong group, primarily due to records of broadcast spawning in sympatry with Nocomis and association with other species' nest in sympatry with Nocomis, respectively. Placement of these two species in either group solely based on visual plot interpretation was undesirable.

Because strong nest associates clearly grouped near the center of the ordination plot (dimension 1 and 2 [approximately equal to] 0, 0), a plot of the absolute values of each species' scores on dimension 1 and 2 provided a more easily interpretable visualization of nest association strength groupings (Fig. 1b). Based on this plot, a simple vertical line on dimension 1 (score = 1.0) delineated the two groups. Rosyface and saffron shiners remained in the strong group but are justifiably considered as moderate associates based on greater deviations from the core group on dimension 2.


Averaged ranking based on scores on the two dimensions supported our visual interpretations of ordination plots. When rankings were averaged between the two dimensions, rosyface and saffron shiners occurred towards the weaker end of the strong group (Table 5). Based on these rankings and the visual interpretations of ordination plots, we concluded that rosyface and saffron shiners represent strong nest associates of Nocomis but probably use nest association in a more facultative manner than the other five species in the strong group.


Analysis of the geographic range overlap between nest associates and Nocomis among groups showed that strong associates had larger geographic range overlap with Nocomis (mean = 92%, SE = 0.03) than the weak group (mean = 37%, SE = 0.11) (P = 0.0002). Spawning temperature overlap with Nocomis was similar between strong (mean = 92%, SE = 0.05) and weak (mean = 94%, SE = 0.04) groups.

Six of seven strong nest associates met the classification as rare based on at least one dimension of rarity, whereas weak nest associates came out as common across all three dimensions of rarity (Table 6). Fisher's exact test revealed that strong nest associates are much more likely to exhibit rarity than weak associates (P = 0.0152).


The methods in this study prove to be a reliable approach for grouping stream fishes to determine their nest association strength. We identified a group of species that, based on reproductive trait similarities, probably are strong nest associates of Nocomis. Members of this group occur mostly (and sometimes entirely) within the range of Nocomis. Thus, strong nest associates clearly depended more on Nocomis nests for spawning than do those in the weaker group. Unless they can associate with alternative hosts, reproduction of these species will be difficult without Nocomis species. Over large spatial scales, it appears dispersal of strong nest associates may be limited to systems where their hosts are present. In mutualisms such as nest association, dispersal of associates is often dependent upon that of the host (Cushman and Beattie, 1991).

Spawning temperature overlap with Nocomis did not differ between the two groups, suggesting that for the species used in this study, spawning temperature overlap is an insensitive indicator of nest association strength. This observation may be an artifact of the level of detail at which most life-history studies operate. Most studies report the range of temperatures at which spawning observations occur but not individual observations. Yet, reports often indicate that nest associates spawn synchronously with Nocomis (Cooper, 1980; Maurikis et al., 1998). Further, Nocomis spawns between 12.5 (Jenkins and Burkhead, 1994) and 26 C (Vives, 1990). Since this encompasses much of the available breeding season for cyprinids, it may explain the lack of observable differences in spawning temperature between the groups.

For the purposes of this study, we present nest association as a binary condition (strong or weak). In nature, however, nest association, like all mutualisms, may be a continuum (Bronstein, 2009). When examining specific spawning behaviors that drive rosyface and saffron shiners away from the strong group, one may conclude that these species represent a moderate group of nest association strength. For instance, rosyface shiner exhibits the least amount of range overlap with Nocomis and sometimes can use broadcast spawning in sympatry with Nocomis. Obviously, this species cannot be a strong associate throughout its entire range because it persists in the absence of Nocomis. However, a plurality of the records for this species documents nest association with Nocomis.

This observation would be easily explainable if the study areas for each spawning mode observation represented a nonrandom distribution. However, this did not occur, as many reports of spawning mode duality often came from the same article (e.g., Reed, 1957; Miller, 1964). Accordingly, rosyface shiner probably represents an opportunistic nest associate that uses differing spawning modes based on local conditions. Saffron shiner diverges from the strong nest associate group for different reasons. Our groupings are based on nest association strength with Nocomis, and saffron shiner was driven away from the strong group by records of nest association with Semotilus. Semotilus nests (pit-ridge) exhibit similar construction and maintenance to the mound nests built by Nocomis. This does not negate the strength of this species' tendency for nest association. Instead, it reflects upon the flexibility of the species' host choices throughout its range.

Knowledge of the plasticity of reproductive modes remains vital to quantifying species-level responses to environmental variability (Gaston, 1994; Kunin, 1997; Johnston, 1999). Five of the seven species we identified as strong nest associates also use the more ancestral reproductive mode of broadcast spawning (Johnston and Page, 1992). Despite this knowledge, virtually no research has investigated the contexts in which a particular spawning mode may be favored. At small spatiotemporal scales, the relative necessity for nest association may vary based on local biotic or abiotic conditions. For instance, parental care provided by hosts greatly reduces the risk of egg predation on associate broods (Johnston, 1994a; Shao, 1997).

In a given spawning season, high abundances of egg predators may cue potential associates to seek out host nests to avoid egg predation. Further, in heavily silted systems, Nocomis nests are often the only source of clean gravel substrate (Vives, 1990; Fletcher, 1993). In these situations, pressure for a species to use nest association may be greater than in less silted conditions. Reports of dual spawning modes could be attributable to differences in contexts between instances of observation, especially for species like rosyface shiner which are widely distributed and more likely to be subjected to encounter habitat heterogeneity throughout its range. Another potentially influential biotic factor may lie in the presence or abundances of different hosts. In light of saffron shiner's observed nest association strength, a study of the dependency of an associate on any given host, or simply on the process itself, would be a useful contribution to this field of study.

It is possible that the variable amount of studies for each species introduced bias into the analyses by not allowing for the observation of potential alternative reproductive modes. However, many of the species with lower numbers of studies were also more narrowly distributed, which reduces the risk that alternative modes of reproduction are known elsewhere. The necessity for describing spawning modes (and other life-history characteristics) throughout species' ranges is clearly presented: without basic life-history knowledge, more complex trait-based ecological analyses may be misguided or unachievable. As research programs continue to address higher-order issues, researchers must also strive to fill basic knowledge gaps of species' life-histories.


Nest association strength strongly related to species' rarity, particularly rarity in geographic extent (four of seven strong nest associates). Only two strong nest associates met rare considerations in terms of habitat breadth. Rosefin shiner did not meet rarity criteria by range extent (although its range is contained mostly within that of Nocomis) but instead by local abundance (Pritt and Frimpong, 2010). Increased reproductive success through nest association may explain high local abundances of both hosts and associates (Wallin, 1992; Walser, 2000; Herrington and Popp, 2004). In most New River tributaries, bluehead chubs and their associates are some of the most locally abundant species found in community samples (Jenkins and Burkhead, 1994; Hitt and Roberts, 2011).

This study presents a previously overlooked conservation aspect. Many of the strong nest associates we identified exhibit narrow geographic distributions, and their typically high local abundances depend on their host. Accordingly, we posit that if abundances of Nocomis were to be drastically reduced (or if Nocomis was extirpated) in the ranges of geographically rare nest associates, populations of the associates would experience significant decreases in abundance. If such a scenario arose, then species currently regarded as relatively ubiquitous within their ranges (regardless of extent) would rapidly exhibit imperiled characteristics. Of course, this is inconsiderate of an associate's ability to revert to broadcast spawning; a strong associate may not necessarily be an obligate associate. Possibly, in the absence of hosts, many of these species could use more ancestral forms of reproduction (Johnston and Page, 1992). We encourage future research to follow that of Wallin (1992) and Black (2007) in empirically quantifying nest association strength.

Since nest association strength is associated with limited geographic distribution (rarity), and rarity is closely associated with imperilment, then why are so few nest associates considered imperiled? Based on a recently updated list of imperiled fishes (Jelks et al., 2008), Pritt and Frimpong (2010) observed that rarity by local population size is a strong determinant of species' imperilment status. Few nest associates or mound building hosts exhibit rarity based on local population size, although several exceptions exist (Andersen, 2002; Mammoliti, 2002; Hamed and Alsopp, 2005). Instream habitat degradation affects stream fishes which construct nests and provide parental care to broods less than those with simpler spawning modes (Berkman and Rabeni, 1987; Johnston and Page, 1992; Peoples et al., 2011). Further, mound construction by Nocomis seems to counteract some negative effects of siltation on reproductive success (Vives, 1990; Peoples et al., 2011). Accordingly, species that have evolved to use (or exploit other species') complex reproductive strategies may possess an evolutionary advantage to persistence as the landscape continues to be influenced by anthropogenic degradation (Johnston and Page, 1992). Emerging research has shown that Nocomis and its associates possess a higher likelihood than other species to colonize and persist in habitats that have undergone significant agricultural disturbance (Hitt and Roberts, 2011).

Examining nest association strength provides a new direction for relating reproductive modes to the conservation of cyprinids whose modes of reproduction are poorly documented. Nest association should assume increasing significance in the future of conservation planning for North American cyprinids. When designing conservation plans for North American cyprinids, attention must be given to spawning modes (Johnston, 1999) and the conservation of mutualisms (Bronstein, 2009) so that natural assemblages, and thus native species, persist.
APPENDIX 1.--References used for reproductive trait review and

Blacknose dace            Central stoneroller

Bartnik, 1970           Etnier and Starnes, 1993
Becker, 1983            Hankinson, 1919

Jenkins and Burkhead,   Jenkins and Burkhead,
  1994                    1994
Menhinick, 1991         Johnston, 1991

Mettee et al., 1996     Johnston, 1994
Schwartz, 1958          Lachner, 1952
Tarter, 1969            Menhinick, 1991
Werner, 2004            Miller, 1962a
                        Miller, 1964
                        Raney, 1947a
                        Reed, 1958
                        Reighard, 1943
                        Smith, 1935
                        Werner, 2004
Crescent shiner         Longnose dace
Hambrick, 1977          Bartnik, 1970
Hankinson, 1919         Bartnik, 1972
Lachner, 1952           Copper, 1980
Maurakis, and           Etnier and Starnes, 1993
Woolcott, 1993
Menhinick, 1991         McPhail and Lindsey,
Raney, 1947a            Menhinick, 1991
                        Werner, 2004
Bluehead chub           Saffron shiner
Maurakis et al., 1992   Copper, 1980
Wallin, 1989              Jenkins and Burkhead,
Wallin, 1992            Menhinick, 1991

Blacknose dace                Rosyside dace

Bartnik, 1970           Etnier and Starnes, 1993
Becker, 1983            Jenkins and Burkhead,
Jenkins and Burkhead,   Johnston, 1991
Menhinick, 1991         Johnston, 1994
Mettee et al., 1996     Lachner, 1952
Schwartz, 1958          Maurakis and Woolcott,
Tarter, 1969            Menhinick, 1991
Werner, 2004            Ross, 2001
Crescent shiner         Mountain redbelly dace
Hambrick, 1977          Hambrick, 1977
Hankinson, 1919         Jenkins and Burkhead,
Lachner, 1952           Lachner, 1952
Maurakis, and           Maurakis and Woolcott,
Woolcott, 1993            1992
Menhinick, 1991         Menhinick, 1991
Raney, 1947a            Raney, 1947a
Bluehead chub           Swallowtail shiner
Maurakis et al., 1992   Jenkins and Burkhead,
Wallin, 1989            Menhinick, 1991
Wallin, 1992            Raney, 1947a
                        Raney, 1947b
                        Werner, 2004

Blacknose dace              Rosyface shiner

Bartnik, 1970           Becker, 1983
Becker, 1983            Copper, 1980
Jenkins and Burkhead,   Hankinson, 1932
Menhinick, 1991         Jenkins and Burkhead,
Mettee et al., 1996     Lachner, 1952
Schwartz, 1958          Menhinick, 1991
Tarter, 1969            Miller, 1962a
Werner, 2004            Miller, 1962b
                        Miller, 1964
                        Pfeiffer, 1955
                        Pflieger, 1975
                        Raney, 1940
                        Raney, 1947a
                        Reed, 1957
                        Reighard, 1943
                        Werner, 2004
Crescent shiner         Rosefin shiner
Hambrick, 1977          Etnier and Starnes, 1993
Hankinson, 1919         Jenkins and Burkhead,
Lachner, 1952           Johnston, 1991
Maurakis, and           Menhinick, 1991
Woolcott, 1993
Menhinick, 1991         Raney, 1947a
Raney, 1947a            Yokely, 1974
Bluehead chub           White shiner
Maurakis et al., 1992   Jenkins and Burkhead,
Wallin, 1989            Maurakis, and Woolcott,
Wallin, 1992

Acknowledgments.--The Virginia Agricultural Experiment Station funded this research.


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Department of Fish and Wildlife Conservation, Virginia Polytechnic Institute and State University, Blacksburg 24061

(1) Present address: Department of Biological Sciences, University of North Texas, Denton 76203

(2) Present address: Department of Environmental Sciences, University of Toledo, Lake Erie Center, Oregon, Ohio 43616

(3) Corresponding author: Telephone: (540) 231-6880; e-mail:
TABLE 1.--Reproductive traits obtained from literature review and
coding used for trait  similarity analysis

Reproductive Trait                                        Code

Builds own nest in sympatry with Nocomis                  BNS
Builds own nest in allopatry with Nocomis                 BNA
Broadcast spawning on open substrate in sympatry with     BS
Broadcast spawning on open substrate in allopatry with    BA
Spawns in natural depressions in sympatry with Nocomis    NDS
Spawns in natural depressions in allopatry with Nocomis   NDA
Associates with Nocomis nests in sympatry                 NNS
Associates with other species nests in sympatry with      NOS
Provides parental care                                    PC
Spawns on sand substrate                                  SAND

TABLE 2.--Frequency at which reproductive traits are observed
over species (1) ranges, based on the  number of records found for
each species. See Table 1 for trait codes

                      Reproductive trait

Species   BNS    BNA    BS     BA     NDS    NDA    NNS

BNDA      0      0      0.50   0.17   0      0.17   0
CESR      0.54   0.08   0      0      0      0      0.54
CRSH      0      0      0      0      0      0      1
LNDA      0      0      0      0.33   0      0.33   0.33
MRBD      0      0      0      0      0      0      0.83
RFSH      0      0      0      0      0      0      0.75
RSDA      0      0      0      0      0      0      0.71
RYSH      0      0      0.31   0      0.08   0.08   0.77
SASH      0      0      0.50   0      0      0      1
STSH      0      0      0.67   0      0      0      0.33
WHSH      0      0      0      0      0      0      1

          Reproductive trait

Species   NOS    PC     SAND

BNDA      0.17   0.17   0.33
CESR      0.15   0.08   0.08
CRSH      0      0      0
LNDA      0      0.33   0
MRBD      0.33   0      0
RFSH      0.50   0      0
RSDA      0.57   0      0
RYSH      0.24   0      0
SASH      1      0.50   0
STSH      0.33   0.33   0.67
WHSH      0      n      0

(1) Species: (BNDA) blacknose dace, (CESR) central stoneroller,
(CRSH) crescent shiner, (LNDA) longnose dace, (MRBD) mountain
redbelly dace, (RFSH) rosefin shiner, (RSDA) rosyside dace,
(RYSH) rosyface shiner, (SASH) saffron shiner, (STSH) swallowtail
shiner, (WHSH) white shiner

TABLE 3.--Phylogenetic similarity among nest associates of
Nocomis occurring in the New River basin, Virginia. Similarities
are based on the number of nodes separating two genera in the
supertree hypothesis proposed by Gaubert et al. (2009). See Table
2 for species codes


BNDA      21
CESR      13     21
CRSH       9      8     21
LNDA      21     13      9     21
MRBD       7      6      4      7     21
RFSH      10      9     19     10      5     21
RSDA      12     12     10     12     11     11     21
RYSH       6      5     17      6      1     17     11     21
SASH       7      6     18      7      2     17      8     19     21
STSH       7      6     18      7      2     17      8     19     21
WHSH       9      8     21      9      4     20     10     17     18

Species   STSH   WHSH

STSH      21
WHSH      18     21

TABLE 4--Correlations of raw reproductive trait frequencies with
ordination axes. References used to interpret axis relationships
are in bold. See Table 1 for trait codes

            Axis 1            Axis 2

Trait     r        P        r        P

BNS      0.67#   0.025#   -0.17    0.628
BNA      0.67#   0.025#   -0.17    0.628
BS      -0.60#   0.053#   -0.54#   0.086#
BA      -0.53    0.095     0.48    0.134
NDS      0.02    0.963     0.48    0.139
NDA     -0.53    0.095     0.59#   0.055#
NNS      0.58    0.059    -0.02    0.963
NOS     -0.07    0.836    -0.62#   0.042#
PC      -0.48    0.139    -0.44    0.171
SAND    -0.55#   0.081#   -0.49    0.125

Note: References used to interpret axis relationships are indicated
with #.

TABLE 5.--Raw and absolute values of dimension scores derived
from an ordination of a phylogenetically independent trait
similarity matrix of eleven nest associates of Nocomis occurring
in the New River basin, Virginia. Ranks are based on absolute
values of scores, and arranged from strongest (1) to weakest
(11). See Table 2 for species codes

          Dimension 1   Dimension 2   Dimension 1 score
Species   score (raw)   score (raw)   (absolute value)

RSDA         0.3148       -0.0424          0.3148
RFSH         0.3178       -0.0235          0.3178
MRBD        -0.4089        0.0484          0.4089
SASH        -0.0333       -1.3501          0.0333
RYSH         0.0589        1.1939          0.0589
CRSH         0.9117        0.2707          0.9117
WHSH         0.9117        0.2707          0.9117
BNDA        -1.6659       -0.1873          1.6659
STSH        -1.6227       -1.1629          1.6227
CESR         2.4667       -0.4138          2.4667
LNDA        -1.2509        1.3964          1.2509

          Dimension 2 score                     Average
Species   (absolute value)    Rank 1   Rank 2    rank

RSDA           0.0424            3        2       2.5
RFSH           0.0235            4        1       2.5
MRBD           0.0484            5        3       4.0
SASH           1.3501            1       10       5.5
RYSH           1.1939            2        9       5.5
CRSH           0.2707            6        6       6.0
WHSH           0.2707            7        5       6.0
BNDA           0.1873           10        4       7.0
STSH           1.1629            9        8       8.5
CESR           0.4138           11        7       9.0
LNDA           1.3964            8       11       9.5

TABLE 6.--Sorted average rank of strong and weak associates,
their rarity tendencies (after Pritt and Frimpong, 2010), and
geographic range and spawning temperature overlap with Nocomis.
The dashed line delineates strong and weak nest associates. See
Table 2 for species codes

             strength        Rare        Rare       Rare by
Species   (Average rank)   by range   by habitat   abundance

RSDA           2.5            No          Yes         No
RFSH           2.5            No          No          Yes
MRBD           4.0            Yes         No          No
SASH           5.5            Yes         No          No
RYSH           5.5            No          No          No
CRSH           6.0            Yes         No          No
WHSH           6.0            Yes         Yes         No

BNDA           7.0            No          No          No
STSH           8.5            No          No          No
CESR           9.0            No          No          No
LNDA           9.5            No          No          No

              Rare                        Spawning
          (Rabinowitz,    Geographic     temperature
Species      1981)       range overlap     overlap

RSDA          Yes             1              1
RFSH          Yes             0.78           0.75
MRBD          Yes             0.86           1
SASH          Yes             0.86           0.98
RYSH          No              0.98           1
CRSH          Yes             1              0.69
WHSH          Yes             1              1

BNDA          No             0.24            1
STSH          No             0.12            1
CESR          No             0.56            0.92
LNDA          No              3              0.85
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Date:Jul 1, 2012
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