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Upon reaching the surf, hatchlings swim toward favorable offshore habitat (Putman et al. 2010). Thereafter, those in the oceanic (surface pelagic) life stage for 1-3 yrs (Ramirez 2019; Ramirez et al. 2020) are dispersed throughout the GoM by surface circulation, and some are also swept through the Florida Straits into the western North Atlantic Ocean (NAO) by the Loop Current, then northward by the Gulf Stream, as well as eastward via the North Atlantic Gyre to European waters (Carr 1963, 1967, 1980, 1986; Hendrickson 1980; Ogren 1989; Collard & Ogren 1990; Marquez-M. 1994; Brongersma 1995; Hildebrand 1995; Marquez M. 2001; Putman et al. 2010, 2013, 2019; NMFS et al. 2011; Witherington et al. 2012; NMFS & USFWS 2015; Putman & Mansfield 2015; Putman et al. 2019; Botterell et al. In press). They also orient and swim in patterns that can alter their direction of travel, and this behavior appears to promote their numerically greater retention within the GoM (Putman & Mansfield 2015). In experiments of swimming performance of captive-reared oceanic stage Kemp’s ridleys, individuals swam against artificially produced currents (Stabenau et al. 1992).

Hendrickson (1980) explained that when the Atlantic Ocean and Pacific Ocean systems were connected, westward flow of the Atlantic North Equatorial Current presumably entered the Pacific system. Kemp’s ridley did not exist as a species at that time, but it diverged later from olive ridley (L. olivacea) after emergence of the Panamanian isthmus (Bowen et al. 1991; Bowen & Karl 2007). With emergence of the Panamanian isthmus, flows of the Loop Current and Gulf Stream became more powerful (Schmidt 2007), and increased the potential for transporting oceanic stage sea turtles from the GoM to the NAO (Hendrickson 1980). Hendrickson (1980) considered this to be “a gigantic leak” that continually extracted an important fraction of the Kemp’s ridley population from the GoM. Putman et al. (2013) used dispersal modeling to predict that 5.1 - 28.4% of oceanic stage Kemp’s ridleys were transported annually from the GoM into the NAO during 2003-2010. Thus, “emigration” of significant portions of oceanic stage Kemp’s ridleys from the GoM to the NAO is a natural annual occurrence. The Atlantic coast of North America is considered important developmental habitat for Kemp’s ridleys (TMTRT 1984; Collard and Ogren
1990; TKRRT 1992; Schmid & Witzell 1997; Witzell 1998; Schmid & Woodhead 2000; Morreale & Standora 2005; Ramirez 2019; Ramirez et al. 2020). Evidence that Kemp’s ridleys reproduce in the NAO is provided by rare nesting events from Florida to New York (Shigetomo 2014; Caillouet et al. 2016a; Rafferty et al. 2019). Although the fate of hatchlings from such nestings is unknown (ibid.), it is unlikely that they survive within the NAO (Ramirez, M.D., pers. comm.). Kemp’s ridleys that do not return to the GoM from the NAO cannot contribute to population growth and recovery within the GoM. However, GoM and NAO ecosystems both provide important developmental habitat in the context of ecological roles of Kemp’s ridley as predator, prey, carrion, scavenger, etc. (Lovich et al. 2018; Caillouet 2019).

Neritic Kemp’s ridleys seasonally migrate and forage in waters within the 50-m depth contour on continental shelves of the GoM and western NAO (NMFS et al. 2011; NMFS & USFWS 2015; Hart et al. 2018; Ramirez et al. 2020). However, the continental shelf along the western NAO coast is generally narrower than that along the GoM coast. Neritic Kemp’s ridleys in the western NAO exhibit seasonal migrations up and down the U.S. east coast (Carr 1980; Hendrickson 1980; Henwood and Ogren 1987; Ogren 1989; Collard & Ogren 1990; Renaud 1995; Gitschlag 1996; Marquez M. 2001; Bernardo & Plotkin 2007; Pritchard 2007; Putman et al. 2010, 2019; NMFS et al. 2011; Bovery & Wyneken 2013; Putman & Mansfield 2015). Water depths <100 m occur between the shoreline and 2 km to the east along Florida’s southeast coast, so the southward migration corridor for neritic Kemp’s ridleys within 50 m is the narrowest there (Land & Paull 2000; Fig. 3 in Pernas et al. 2001; Correa et al. 2012). Southward flow on the western flank of the Florida Current was characterized by Soloviev et al. (2017).
Until recently (Ramirez 2019; Ramirez et al. 2020), emigration and immigration have generally been ignored in age-structured modeling of the Kemp’s ridley population (Heppell et al. 2007). According to Heppell et al. (2007), “Simply stated, population growth occurs when births exceed deaths and/or immigration exceeds emigration. We can ignore the latter for Kemp’s because we have data for the entire species.”

Yet, limited numbers of neritic stage Kemp’s ridleys have been documented to have returned to the GoM from the NAO (Márquez-M. et al. 1995; Heppell et al. 1996; Heppell & Crowder 1998; TEWG 1998, 2000; Heppell et al. 2005, 2007; Crowder & Heppell 2011; NMFS et al. 2011; Bovery & Wyneken 2013; NMFS & USFWS 2015; Gallaway et al. 2016a,b; Kocmoud et al. 2019). Numbers of neritic stage Kemp’s ridleys that remain in, survive to maturity and reproduce within the GoM are likely higher than those that mature and reproduce in the NAO or after returning to the GoM. Migration distances to western GoM nesting beaches are greater from the NAO than from within the GoM. Therefore, it is likely that much higher numbers of each year-class (cohort) of Kemp’s ridleys enter the NAO in the oceanic life stage (Putman et al. 2010, 2013, 2019) than return to the GoM in any life stage or at any age. Fewer than 20 neritic Kemp’s ridleys have been documented to have returned from the NAO to the GoM, although most were reported nesting at Tamaulipas (Schmid 1995; Witzell 1998; Schmid & Woodhead 2000; Renaud & Williams 2005; Schmid & Witzell 2005; Bovery & Wyneken 2013; Caillouet et al. 2015). This paucity of evidence that Kemp’s ridleys return to the GoM from the NAO could result from inadequate numbers tagged and satellite-tracked in the NAO, or inadequate focus of monitoring NAO-tagged and NAO-tracked individuals around the Florida Straits and southwest Florida (Ramirez, M.D., pers. comm.). Yet, numbers of neritic Kemp’s ridleys documented as having been caught or found stranded, with or without tags or transmitters, along the coasts of eastern North America and the GoM have been adequate for evaluations of somatic growth and seasonal migrations (Snover et al. 2007; NMFS et al. 2011; Bovery & Wyneken 2013; NMFS & USFWS 2015). Returns to western GoM nesting beaches of Kemp’s ridley tagged or fitted with transmitters and tracked within the GoM dwarf those for conspecifics tagged or fitted with transmitters in the NAO (ibid.). Unless substantially more evidence is amassed to the contrary, it will continue to appear that most Kemp’s ridleys that nest in the Gulf of Mexico have not spent time in the NAO. The dearth of evidence to the contrary begs the question of whether Kemp’s ridley immigration from NAO to GoM has had or can have a measurable effect on growth of the population within the GoM. Additional tagging and tracking of neritic immature and adult Kemp’s ridleys on the continental shelf along the southeast coast of Florida and Florida Straits may be necessary to resolve questions regarding the magnitude of their immigration from NAO to GoM (Ramirez, M.D., pers. comm.).

Avens et al. (2017) recommended integration of skeletochronology with stable isotope and trace element analyses to increase understanding of long-term changes in trophic ecology and movements of Kemp’s ridleys between GoM and NAO foraging habitats and the relative contributions of these regions to the reproductive population. Ramirez (2019) used complementary lead isotopes (208Pb:206Pb) in humeri collected from dead-stranded Kemp’s ridleys to distinguish, with exceptional accuracy (94.1%), those found in the NAO from those found in the GoM. Perhaps this isotopic tracer methodology could be used to identify Kemp’s ridleys that spent time in the NAO then returned to the GoM (Ramirez, M.D., pers. comm.). Ramirez (2019) also used a spatially explicit, age-structured matrix model to examine habitat-specific demographic rates and variable ontogenetic shifts (oceanic to neritic). Although his model simulations showed that NAO to GoM transition influenced the population during its 1990-2009 period of rapid growth, they suggested that Kemp’s ridleys in the western NAO were not strong contributors to growth of the Kemp’s ridley population during that period, and were unlikely to influence population recovery time, even under the most extreme scenario evaluated that was based on annual emigration of 30% of Kemp’s ridleys in the oceanic stage from the GoM to the NAO, and no immigration to the GoM of survivors in the NAO before age 7.

Over the years, annual numbers of Kemp’s ridleys found stranded alive or dead from natural and anthropogenic causes have increased along GoM and NAO shores (TMTRT 1984; Meylan & Sadove 1986; Witherington & Ehrhart 1989; CSTC 1990; Tomás et al. 2003; Still et al. 2005; Witt et al. 2007; Insacco & Spadola 2010;
NMFS et al. 2011; Caillouet et al. 2015; Nicolau et al. 2016; Avens & Dell’Amico 2018; Liu et al. 2018; Griffin et al. 2019; Innis et al. 2019). Those related to shrimp trawling and cold-stunning have received considerable attention (CSTC 1990; NMFS & USFWS 2015). Substantial effort and resources have been expended toward rescue, resuscitation, rehabilitation and release of live-stranded Kemp’s ridleys. For US rehabilitation programs, Innis et al. (2019) reported that at least 5,137 Kemp’s ridleys found stranded alive were rehabilitated and released during 1997-2016, and 61 were rehabilitated and released by those programs before 1997. They have contributed in various ways to Kemp’s ridley conservation (NMFS et al. 2011; Innis et al. 2019), but their contribution to population growth and recovery has not been evaluated (Caillouet et al. 2015; Caillouet 2016b). It is noteworthy that oceanic stage Kemp’s ridleys that enter Cape Cod Bay apparently are unable to navigate and swim their way out of it before temperatures drop and they become cold-stunned and stranded (Still et al. 2005; Liu et al. 2018; Griffin et al. 2019). This warrants further investigation in the context of their navigational and swimming abilities.

Heppell et al. (1996) and Heppell & Crowder (1998) set the precedent for using age-structured modeling to estimate the probability that a given Kemp’s ridley conservation intervention contributed to Kemp’s ridley population growth, and the use and improvement of age-structured models for such purposes has continued since then. We therefore recommend that age-structured modeling be used to determine the contribution made by Kemp’s ridleys in the NAO to population growth in the GoM (Caillouet et al. 2015; Caillouet 2016b). The negative differences between annual numbers that “emigrate” in the oceanic stage from the GoM to the NAO and those that return to the GoM in any life stage should be included in age-structured modeling of the population in the GoM. They represent natural losses to the population in the GoM, and could be treated as natural mortality. If these differences were included in modeling, estimated total mortalities in the GoM would thereby be increased, and the proportion of total mortality represented by anthropogenic mortality would thereby be decreased. Results could have implications for Kemp’s ridley conservation actions and interventions going forward. Therefore, emigration and immigration of Kemp’s ridleys between the GoM to the NAO should no longer be ignored in age-structured modeling of the Kemp’s ridley population in the GoM. However, as climate warms and sea level rises (NMFS et al. 2011; Pike 2013a,b), Kemp’s ridleys in the NAO may become more important to population recovery and resilience if they adapt to such changes and establish new nesting colonies along the NAO coast (Caillouet 2012; Bevan et al. 2019; Butler 2019).

Acknowledgements. We thank Nathan F. Putman and Matthew D. Ramirez for their reviews of the manuscript.

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