Goodeidae

Goodeidae Through the Late Eocene and Oligocene, reorganization and drainage expansion within the Madrean River (Galloway et al. 2011, Snedden and Galloway 2019) could have allowed goodeids to disperse along the front of the Sierra Madre Occidental to the southern Great Basin (Fig. 7). Divergence of Great Basin Empetrichthyinae from Goodeinae of the Trans-Mexican Volcanic Belt and Sierra Madre 23.3–12.9 Mya (Fig. 3, node 6; Table 2) agrees with break-up of this drainage at the end of the Oligocene (Snedden and Galloway 2019). The great distance between northern (Empetrichthyinae) and southern (Goodeinae) goodeids is a biogeographical anomaly (Webb 2020). Parenti (1981) proposed that regional desiccation eliminated intervening populations (Grant and Riddle 1995, Webb et al. 2004, Miller et al. 2005, Webb 2020), but living Empetrichthyinae occupy the driest region of North America (i.e. aridity is associated with survival). Fragmentation of the Madrean River might have been an additional cause of extirpations (Fagan et al. 2002), and climatic cooling might have reduced habitat suitability at higher elevations and latitudes, perhaps explaining why Empetrichthyinae are associated with warm-water springs. Loss of goodeids from the Gulf of México drainage could reflect Late Miocene integration of the ancestral Río Nazas with the Río Grande, which facilitated an influx of fishes (Hoagstrom and Osborne 2021). Characodon Günther, 1866 persisted only in remote basins († Characodon garmani, Mayran-Parras Basin; Characodon audax Smith & Miller, 1986 – Characodon lateralis Günther, 1866, Río Tunal) (Beltrán-López et al. 2021), potentially protected from invading fishes. It is also possible that remnant goodeid populations were present in the uninhabited region before European settlement, but disappeared owing to habitat destruction, dewatering, or invasive species associated with early European contact. Because this scenario agrees with reconstructed hydrography, it is unnecessary to invoke the popular hypothesis (Miller and Smith 1986, Minckley et al. 1986, Grant and Riddle 1995, Miller et al. 2005, Webb 2020) that northward drift of the Pacific Plate created the gap between Empetrichthyinae and Goodeidae. Furthermore,thehypothesisthatGoodeidaeusedawesternroute from the Great Basin into central México (Pérez-Rodríguez et al. 2015) is unnecessary because goodeids dispersing inland from the Gulf of México via the Madrean River would have reached central México en route to the Great Basin (Fig. 7), not from the Great Basin. Separation of Empetrichthyinae from Goodeinae is attributable to vicariance during fragmentation of the Madrean River drainage, with Empetrichthyinae representing a relict lineage that has persisted in the Mojave Desert after reaching the region when the Madrean River extended from the Gulf Coast to the Basin and Range geomorphic province. Goodeinae represent lineages descending from populations further downstream that persisted within the Sierra Madre Occidental and Trans-Mexican Volcanic Belt after separation from the Gulf of México drainage. Within Goodeinae, divergence of CharacodontiniIllyodontini from crown Goodeinae 16.8–10.3 Mya (Fig. 3, node 7; Table 2) appears to have been an east–west divergence, possibly between ancestral ríos Nazas and Aguanaval (Fig. 8). Reactivation of the San Marcos Fault 14.0–5.0 Mya and contemporary volcanism on the borders of the Mesa Central (Aranda-Gómez et al. 2005, 2007, Chávez-Cabello et al. 2007, Nieto-Samaniego et al. 2007) are likely to have contributed to drainage fragmentation. Positioning of crown Goodeinae on the Trans-Mexican Volcanic Belt throughout its tectonic evolution, which initiated ~20 Mya and lasted 17 Myr (Ferrari et al. 2012), ensured that goodeids experienced frequent drainage reorganizations and shifting hydrographic barriers (Barbour 1973, Domínguez‐Domínguez et al. 2010, Pérez-Rodríguez et al. 2015). Barrier displacement via stream capture was probably the main mode of range expansion and diversification (Webb et al. 2004, Domínguez‐Domínguez et al. 2010, Beltrán-López et al. 2021).