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Migration rates among Beringian Dallia populations. Migration rates in individuals per generation since the time of divergence (t) between populations of Dallia estimated by Isolation with Migration analyses. Approximate geographic distribution of populations defined by clustering algorithms are outlined with black dashed lines. Arrows reflect the magnitude of migration rate. Current coastlines and international borders (yellow lines) are overlaid onto a map of -110 meter sea level (dark green layer) corresponding to the last glacial maximum available from the US National Geophysical Data Center
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Pleistocene climatic instability had profound and diverse effects on the distribution and abundance of Arctic organisms revealed by variation in phylogeographic patterns documented in extant Arctic populations. To better understand the effects of geography and paleoclimate on Beringian freshwater fishes, we examined genetic variability in the genus...
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... of migration rates between populations in the IM comparisons were in general asymmetrical and ranged between 0.08 and 2.42 individuals per generation on aver- age since the time of divergence between each population pair (calculated by (mean population θ x mean migration rate m)/2) (Fig 4). The migration rate from the coastal Alaska population to the other three populations was consistently smaller than the reverse. ...
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... In contrast, Beringia is an area where early Paleoindian fishing adaptations could be observed, and it occupies a key geographic position in current peopling of the Americas models. Beringia was mostly ice-free during the last glacial period and was a major refugium for a variety of fish taxa (12)(13)(14), and open-air loess sites with excellent organic preservation have the potential to preserve delicate fish remains. Very little is known about early fishing in Beringia, and reports are generally limited to the presence/absence of fish with no further taxonomic specificity (15,16). ...
While freshwater and anadromous fish have been critical economic resources for late prehistoric and modern Native Americans, the origin and development of fishing is not well understood. We document the earliest known human use of freshwater and anadromous fish in North America by 13,000 and 11,800 years ago, respectively, from primary anthropogenic contexts in central Alaska (eastern Beringia). Fish use appears conditioned by broad climatic factors, as all occurrences but one are within the Younger Dryas chronozone. Earlier Bølling-Allerød and later early Holocene components, while exhibiting similar organic preservation, did not yield evidence of fishing, suggesting that this was a response to changing environmental factors, perhaps reductions in higher ranked resources such as large terrestrial mammals. Late Pleistocene and recent Indigenous peoples harvested similar fish taxa in the region (salmon, burbot, whitefish, and pike). We characterize late Pleistocene fishing in interior Beringia as an important element of broad-spectrum foraging rather than the intensive communal fishing and storage common among recent peoples.
... Several strictly freshwater species, particularly Dallia pectoralis Bean, 1880, Catostomus catostomus Forster, 1773, Coregonus pidschian Gmelin, 1789, Cottus cognatus Richardson, 1836, Thymallus arcticus Pallas, 1776, and Prosopium coulterii (Eigenmann et Eigenmann 1892), have been known to have trans-Beringian distribution, which is evidently defined by the water net transformations and a land bridge between North America and Asia during the cold ages. Several detailed phylogeographic reconstructions denote a specific colonization history with different resettlement pathways during different times for each of the abovementioned species [5][6][7][8][9][10]. However, the over-arching concept of the freshwater fauna formation and timing of resettlement is still lacking for the Beringian area. ...
The pygmy whitefish Prosopium coulterii (C. H. Eigenmann & R. S. Eigenmann, 1892) is a freshwater fish with a highly disjunct distribution ranging from the middle part of North America to Chukotka. There is still no consensus regarding its phylogeny and dispersal history due to limited information from the Chukotkan part of the range. We investigated 22 lakes over Chukotka and found a much broader distribution than it was previously thought. Pygmy whitefish was found to be a common species in the lakes that belong to rivers draining into the Arctic. Cytochrome B, cytochrome oxidase subunit 1, and ATP synthase F0 subunit 6 mitochondrial sites were analyzed from 25 samples to reconstruct the phylogenetic history of pygmy whitefish. Two haplogroups belonging to the east and west Chukotkan ranges were identified; both groups are closely related to Alaskan pigmy whitefish and distant from the Cascadia-Mackenzie (Peace) populations. Combining the distribution patterns, phylogenetic network topology, and the contemporary knowledge on the glaciation history of the region, we suggest a possible colonization pathway over Beringia region and beyond it. The basic biological characteristics (fork length, number of gill rakers, and pyloric caeca, age structure, and feeding) are also presented to characterize the populations over the investigated range.
... Some fishes and small mammals, on the other hand, do provide evidence for intra-Beringian refugia, reflecting fundamental differences in dispersal abilities (e.g. Campbell, Takebayashi & L opez, 2015;Cook et al., 2017). Marine organisms such as sea stars and sea cucumbers show the expected geographically orthogonal trans-Beringian divergences (i.e. ...
Beringia is a biogeographically dynamic region that extends from northeastern Asia into northwestern North America. This region has affected avian divergence and speciation in three important ways: (i) by serving as a route for intercontinental colonisation between Asia and the Americas; (ii) by cyclically splitting (and often reuniting) populations, subspecies, and species between these continents; and (iii) by providing isolated refugia through glacial cycles. The effects of these processes can be seen in taxonomic splits of shallow to increasing depths and in the presence of regional endemics. We review the taxa involved in the latter two processes (splitting-reuniting and isolation), with a focus on three research topics: avian diversity, time estimates of the generation of that diversity, and the regions within Beringia that might have been especially important. We find that these processes have generated substantial amounts of avian diversity, including 49 pairs of avian subspecies or species whose breeding distributions largely replace one another across the divide between the Old World and the New World in Beringia, and 103 avian species and subspecies endemic to this region. Among endemics, about one in three is recognised as a full biological species. Endemic taxa in the orders Charadriiformes (shorebirds, alcids, gulls, and terns) and Passeriformes (perching birds) are particularly well represented, although they show very different levels of diversity through evolutionary time. Endemic Beringian Charadriiformes have a 1.31:1 ratio of species to subspecies. In Passeriformes, endemic taxa have a 0.09:1 species-to-subspecies ratio, suggesting that passerine (and thus terrestrial) endemism might be more prone to long-term extinction in this region, although such 'losses' could occur through their being reconnected with wider continental populations during favourable climatic cycles (e.g. subspecies reintegration with other populations). Genetic evidence suggests that most Beringian avian taxa originated over the past 3 million years, confirming the importance of Quaternary processes. There seems to be no obvious clustering in their formation through time, although there might be temporal gaps with lower rates of diversity generation. For at least 62 species, taxonomically undifferentiated populations occupy this region, providing ample potential for future evolutionary diversification.
... The BLB remained predominantly ice-free during glacial advances across its Eurasian and North American extensions, acting for many taxa as a glacial refugium and as an important source and route of post-glacial colonization. Examples can be found among plants (Brubaker et al., 2005), insects (Elias & Crocker, 2008), fish (Campbell et al., 2015), and mammals (Anijalg et al., 2018;Heintzman et al., 2016), including humans (Watson, 2017). Yet, habitat and climate heterogeneity across the BLB shaped species ranges and promoted population structure (Dale Guthrie, 2001;Elias & Crocker, 2008;Kuzmina et al., 2011;Vershinina et al., 2021). ...
Glacial and interglacial periods throughout the Pleistocene have been substantial drivers of change in species distributions. Earlier analyses suggested that modern grey wolves (Canis lupus) trace their origin to a single Late Pleistocene Beringian population that expanded east and westwards, starting ca. 25,000 years ago (ya). Here, we examined the demographic and phylogeographic histories of extant populations around the Bering Strait with wolves from two inland regions of the Russian Far East (RFE) and one coastal and two inland regions of North‐western North America (NNA), genotyped for 91,327 single nucleotide polymorphisms. Our results indicated that RFE and NNA wolves had a common ancestry until ca. 34,400 ya, suggesting that these populations diverged before the previously proposed expansion out of Beringia. Coastal and inland NNA populations diverged ca. 16,000 ya, concordant with the minimum proposed date for the ecological viability of the migration route along the Pacific Northwest coast. Demographic reconstructions for inland RFE and NNA populations reveal spatial and temporal synchrony, with large historical effective population sizes that declined throughout the Pleistocene, possibly reflecting the influence of broad‐scale climatic changes across continents. In contrast, coastal NNA wolves displayed a consistently lower effective population size than the inland populations. Differences between the demographic history of inland and coastal wolves may have been driven by multiple ecological factors, including historical gene flow patterns, natural landscape fragmentation, and more recent anthropogenic disturbance.
... The Alaska blackfish (Dallia pectoralis) is a facultative air-breather endemic to the tundra wetlands of western and northern mainland Alaska, the Bering Sea Islands and Eastern Siberia (Campbell and Lopéz, 2014;Campbell et al., 2015). Unlike most extant air-breathing fishes, which reside in tropical freshwaters and are rarely, if ever, restricted from accessing atmospheric oxygen for prolonged time periods, the Alaska blackfish is forcibly submerged by ice and snow cover for many months in winter, which not only precludes air-breathing, but also results in aquatic hypoxia. ...
The Alaska blackfish (Dallia pectoralis) is a facultative air-breather endemic to northern latitudes where it remains active in winter under ice cover in cold hypoxic waters. To understand the changes in cellular Ca²⁺ cycling that allow the heart to function in cold hypoxic water, we acclimated Alaska blackfish to cold (5 °C) normoxia or cold hypoxia (2.1–4.2 kPa; no air access) for 5–8 weeks. We then assessed the impact of the acclimation conditions on intracellular Ca²⁺ transients (Δ[Ca²⁺]i) of isolated ventricular myocytes and contractile performance of isometrically-contracting ventricular strips. Measurements were obtained at various contractile frequencies (0.2–0.6 Hz) in normoxia, during acute exposure to hypoxia, and reoxygenation at 5 °C. The results show that hypoxia-acclimated Alaska blackfish compensate against the depressive effects of hypoxia on excitation-contraction coupling by remodelling cellular Δ[Ca²⁺]i to maintain ventricular contractility. When measured at 0.2 Hz in normoxia, hypoxia-acclimated ventricular myocytes had a 3.8-fold larger Δ[Ca²⁺]i peak amplitude with a 4.1-fold faster rate of rise, compared to normoxia-acclimated ventricular myocytes. At the tissue level, maximal developed force was 2.1-fold greater in preparations from hypoxia-acclimated animals. However, maximal attainable contraction frequencies in hypoxia were lower in hypoxia-acclimated myocytes and strips than preparations from normoxic animals. Moreover, the inability of hypoxia-acclimated ventricular myocytes and strips to contract at high frequency persisted upon reoxygenation. Overall, the findings indicate that hypoxia alters aspects of Alaska blackfish cardiac myocyte Ca²⁺ cycling, and that there may be consequences for heart rate elevation during hypoxia, which may impact cardiac output in vivo.
... Previous studies have suggested that genetic diversity in western North American wildlife populations is the result of recurrent cycles of range shifts caused by expansion and contraction, isolation, and potential introgression that, in combination with habitat heterogeneity, potentially explains the observed intraspecific diversity and local adaption (Campbell et al., 2015;Galbreath et al., 2011;Shafer et al., 2010). There have been many genetic and genomic studies which have reconstructed the evolutionary history of wildlife in relation to glacial cycles in North America, particularly from the last glacial maximum (LGM; 26.5,000-19,000 years ago [kya]; Clark et al., 2009), and events since then. ...
Pleistocene glacial cycles influenced the diversification of high‐latitude wildlife species through recurrent periods of range contraction, isolation, divergence, and expansion from refugia and subsequent admixture of refugial populations. We investigate population size changes and the introgressive history of caribou (Rangifer tarandus) in western Canada using 33 whole genome sequences coupled with larger‐scale mitochondrial data. We found that a major population expansion of caribou occurred starting around 110 kya, the start of the last glacial period. Additionally, we found effective population sizes of some caribou reaching ~700,000 to 1,000,000 individuals, one of the highest recorded historical effective population sizes for any mammal species thus far. Mitochondrial analyses dated introgression events prior to the LGM dating to 20‐30 kya and even more ancient at 60 kya, coinciding with colder periods with extensive ice coverage, further demonstrating the importance of glacial cycles and events prior to the LGM in shaping demographic history. Reconstructing the origins and differential introgressive history has implications for predictions on species responses under climate change. Our results have implications for other whole genome analyses using Pairwise Sequentially Markovian Coalescent (PSMC) analyses, as well as highlighting the need to investigate pre‐LGM demographic patterns to fully reconstruct the origin of species diversity, especially for high‐latitude species.
... On the other hand, there existed several aquatic refugia in Beringia during the Pleistocene Glacial Maximum, which promoted the genetic structuring within freshwater organisms (Campbell, Takebayashi, & López, 2015) and, potentially, may be viewed as a driver of speciation processes. Therefore, we find it probable that the Alaskan species L. perpolita is not conspecific with K. kamtschatica and may represent a Nearctic vicariate taxon sister to the latter. ...
The taxonomic position of three nominal species of lymnaeid snails placed by Kruglov & Starobogatov (1993) into the subgenus Lymnaea ( Pacifimyxas) Kruglov & Starobogatov, 1985, has been re-assessed based on a molecular genetic study of topotypic specimens and an examination of the type series and other materials a vailable. It has been shown that the two species, Lymnaea (Pacifimyxas) magadanensis Kruglov & Starobogatov, 1985 and Lymnaea (Pacifimyxas) streletzkajae Kruglov & Starobogatov, 1985, are identical with the species Kamtschaticana kamtschatica (Middendorff, 1850) and must be treated as its junior synonyms. Hence, Pacifimyxas
becomes a junior synonym of Kamtschaticana Kruglov & Starobogatov, 1984. The taxonomic identity of the third species of Pacifimyxas, Lymnaea (Pacifimyxas) perpolita, remains obscure, and this species is considered here as taxon inquirendum. Two other
nominal species, Lymnaea aberrans (Westerlund, 1897) and Lymnaea middendorffi (W. Dybowski, 1904), have been synonymized with K. kamtschatica based on morphological and geographical evidence. The lectotype of Limnaea peregra var. middendorffi is
designated. The actual level of species richness in the Beringian freshwater malacofauna may be 20–25% lower than it was determined on the basis of the traditional system. Some implications of this outcome for the biogeography of the Beringian freshwater fauna are discussed.
... 18. Dallia admirabilis Chereshnev, 1980 -Wonderful blackfish According to Mecklenburg et al. (2002) and Campbell et al. (2015), this species is a synonym of Dallia pectoralis Bean, 1880; which agreed to the work of Campbell et al. (2015), the latter authors stress the similarity of the morphology of this species with some specimens from Alaska identified as D. pectoralis. Nevertheless, prior to relevant studies with a large number of samples throughout the known area for all members of the Dallia genus, the status of these populations remains questionable. ...
... 18. Dallia admirabilis Chereshnev, 1980 -Wonderful blackfish According to Mecklenburg et al. (2002) and Campbell et al. (2015), this species is a synonym of Dallia pectoralis Bean, 1880; which agreed to the work of Campbell et al. (2015), the latter authors stress the similarity of the morphology of this species with some specimens from Alaska identified as D. pectoralis. Nevertheless, prior to relevant studies with a large number of samples throughout the known area for all members of the Dallia genus, the status of these populations remains questionable. ...
In the present paper, the history of fish faunistic and taxonomic knowledge is analyzed, and the freshwater and brackish water fish diversity in the territory of the Russian Federation and adjacent areas is examined. An overview of higher taxa and species is presented (3 classes, 26 orders, 100 families, and 317 genera), including a total of 719 native fish species (plus 36 introduced species, and also 16 species not yet recoded for Russian waters, but expected in the future, and 20 species that were previously removed from the ichthyofauna of Russia due to taxonomic changes; total number in all categories 791 taxa). The Russian water includes freshwater 353, brackish 329, diadromous 82, and amphidromous 27 species. A total of 103 endemic species in the native ichthyofauna adds to an endemism rate of 14,3%. This study significantly increases the total number of fishes ecologically related to fresh and brackish waters of Russia. The native freshwater and brackish water ichthyofauna of the Russian Federation consists of approximately 4% of the global fish species. The fish diversity in selected river systems and lakes is also discussed, with respect to ecoregions, latitude and longitude, and compared with the fish faunas of Europe and North America (north of 50 o N).
... Acclimation to low temperature results in a higher heart rate, shortened contraction duration, increased contractile rates, increased rate of SR Ca 2+ sequestration and heightened adrenergic sensitivity than at warmer temperatures (Aho and Vornanen, 1999;Aho and Vornanen, 2001;Keen et al., 1993;Tiitu and Vornanen, 2002). The Alaska blackfish (Dallia pectoralis) is an air-breathing fish species that resides in Western Alaska and Eastern Siberia (Campbell and Lopéz, 2014;Campbell et al., 2015), but the overwintering strategy of ...
... Indeed, the findings are consistent with the habitat conditions and ecology of the fish. Alaska blackfish are native to the tundra wetlands of western and northern Alaska (Campbell and Lopéz, 2014;Campbell et al., 2015;Ostdiek and Roland, 1959). In these regions there are many ponds and lakes of varying depths and dissolved oxygen levels that are interconnected by shallow drainage ditches (Haynes et al., 2014;Leppi et al., 2016;Ostdiek and Roland, 1959). ...
The air-breathing Alaska blackfish (Dallia pectoralis) experiences aquatic hypoxia, but restricted air-access in winter due to ice-cover. To lend insight into its overwintering strategy, we examined the effects of thermal acclimation (15 °C vs. 5 °C), acute temperature change (to 10 °C), increased pacing frequency, inhibition of sarcoplasmic reticulum (SR) Ca2+ release and uptake and adrenaline (1000 nmol l-1) on the contractile performance of isometrically-contracting, electrically-paced ventricular strips. At routine pacing frequencies, maximal developed force (Fmax) was equivalent at 5 °C (2.1 ± 0.2 mN mm-2) and 15 °C (2.2 ± 0.3 mN mm-2), whereas contraction durations were 2.2- to 2.4-times longer and contraction rates 2.4- to 3.5-times slower at 5 °C. Maximum contraction frequency was reduced by decreased temperature, being 0.91 ± 0.04 Hz at 15 °C, 0.35 ± 0.02 Hz at 5 °C and equivalent between acclimation groups at 10 °C (~0.8 Hz). 15 °C and 5 °C strips were insensitive to SR inhibition at routine stimulation frequencies, but SR function supported high contraction rates at 10 °C and 15 °C. Adrenaline shortened T0.5R and increased relaxation rate by 18-40% at 15 °C, whereas at 5 °C, adrenaline augmented Fmax by 15-25%, in addition to increasing contraction kinetics by 22-82% and decreasing contraction duration by 20%. Overall, the results reveal that ventricular contractility is suppressed in cold-acclimated Alaska blackfish largely by acute and perhaps direct effects of decreased temperature, which effectively preconditions the tissue for low energy supply during winter hypoxia. Additionally, the level of cardiac performance associated with maintained activity in winter is supported by enhanced inotropic responsiveness to adrenaline at 5 °C.
... For example, water bodies in region 2 flow into Norton Sound, while water bodies in region 3 flow into Bristol Bay and the distance between these coastal regions may also serve as a geographic barrier between populations (this hypothetical barrier line is drawn in Fig. 1);(2) no evidence has been found of localized life history patterns for C. sardinella in which individuals have limited migration over their lifetime that limits gene flow; (3) these patterns of differentiation are inherited from ancient C. sardinella populations that were separated by glacial refugia. A recent study of Alaska blackfish Dallia (Bean 1880 species found similar genetic structuring across Alaska, suggesting the maintenance of genetic diversity formed during the Pleistocene, when glacial sub-refugia existed across Beringia, allowing for genetic differentiation (Campbell et al., 2015). The research identified up to five genetic clusters of Dallia spp. with groups found in the Arctic coastal plain north of the Brooks Range, the upstream Kuskokwim and Yukon River basins, the Seward Peninsula, the Alaska coast downstream of the Kuskokwim and Yukon Rivers and in south-west Alaska and another distinct coastal Alaska group. ...
... Dallia spp. are strictly freshwater fishes and therefore have limited mobility across the landscape and the genetic structuring as seen in the Campbell et al. (2015) study is better understood when considering this particular life history trait. In comparison, C. sardinella are relatively mobile across the landscape, with many populations probably accessing fresh and marine waters at different stages of their life cycles. ...
This study presents the first detailed analysis of the mitochondrial DNA diversity of least cisco Coregonus sardinella in Alaska using a 678 bp segment of the control region (D-loop) of the mitochondrial genome. Findings suggest that the history of C. sardinella in Alaska differs from that of other species of Coregonus present in the state and surrounding regions. The examined populations of C. sardinella are genetically diverse across Alaska. Sixty-eight distinct mitochondrial haplotypes were identified among 305 individuals sampled from nine locations. The haplotype minimum spanning network and phylogeny showed a modest level of geographic segregation among haplotypes, suggesting high levels of on-going or recent connectivity among distant populations. Observed ΦST values and the results of homogeneity and AMOVAs indicate incipient genetic differentiation between aggregations in three broad regional groups. Sites north of the Brooks Range formed one group, sites in the Yukon and Selawik Rivers formed a second group and sites south of the Yukon drainage formed the third group. Overall, the sequence data showed that a large proportion of mtDNA genetic variation in C. sardinella is shared across Alaska, but this variation is not homogeneously distributed across all regions and for all haplotype groups.
