Fig. 6: DIYABC Scenario 9 - Events and marine isotope stage (MIS) assignment
According to our results from the ABC, the calibrated times of demographic events can be assigned to marine isotope stages (MIS; Railsback 2006) (Fig. 6). The first larch colonization (t4, population N1a) has been recorded at around 224,000 years BP, which is equivalent to MIS7 and specified as a moderate interglacial. This suggests that the climate was favorable for larch establishment. The subsequent ancient and severe bottleneck (t4-db, population N1a to population N1) can be dated to 161,750 years BP, corresponding to MIS6, when the penultimate glacial period occurred, which was a more severe glaciation than the LGM. These harsh climate conditions may have caused loss of diversity and a decrease in population size. The population N1a may have been threatened with extinction, but after the bottleneck event, the population size increased again. The subsequent split event (t3) is registered at 32,000 years BP within MIS2, containing the LGM. This period of moderate glaciation did not cause a significant decrease in population size. The following split event (t2) and the admixture event (t1) can be dated at 11,700 years BP and 4,175 years BP, respectively, both in MIS1, thus in the Holocene. The split events and the higher population size indicate better conditions such as greater fitness of pollen or faster growth.
The ABC leads to the assumption that the common ancestors of today’s larch populations (N1 and N2) must have been present in northeast Siberia long before the last glacial. However, the exact timing of both recent and ancient historical events should be considered a rough estimate because of possible biases, as it is difficult to estimate the average generation time of long-lived trees. Because the generation time is likely to be greater than 25 years during colder climate stages, the inferred divergence time is most probably underestimated. Hence, we can deduce that larch populations must have survived in isolated refugia during the last glacial. This hypothesis is also supported by similar findings in other genetic studies that suggest the presence of several refugia during Pleistocene glacial intervals (Polezhaeva et al. 2010; Semerikov et al. 2013). Both pollen and macrofossil evidence indicates the survival of Larix in northern regions throughout the LGM in multiple and often isolated refugia (Khatab et al. 2008; Binney et al. 2009; Müller et al. 2010). Furthermore, we detected a genetic differentiation that serves as an indicator of the mentioned long-term isolation of the recent populations within geographically disconnected refugia (Tóth et al. 2019). For Alaska, there are also indications of a possible in situ persistence of larches during the LGM (Napier et al. 2020). Binney et al. (2017) mention that northern Eurasia is topographically complex so it is likely that the wide range of local climates provided conditions for refugial populations to persist. In the region of the Eastern Yakutian cluster, the sheltered valleys of the Verkhoyansk Mountains (Tarasov et al. 2009) or the Tschuch’ye Lake area in Eastern Yakutia with its deep protected valleys (Lozhkin et al. 2018) could have provided shelter for the persistence of Larix during the LGM.
Our ABC shows that the populations that persisted during the LGM in northern refugia have genetically contributed to post-LGM recolonization. This is also corroborated by other studies that suggest that these populations were established well before the LGM from a single source population (Western Siberia) with probably a small effective size and low recent gene flow (Ma et al. 2020; Semerikov et al. 2013). The population in Chukotka probably originated earlier and the population in Eastern Yakutia subsequently emerged in the course of an admixture event of the populations from Western Yakutia and Chukotka. However, our results contradict the conclusions of Schulte et al. (2022b), who state that L. sibirica had to recolonize northern areas from refugia in the south in the postglacial. It is possible thatL. sibirica retreated locally from the region of the lake investigated in their study during the LGM but survived the LGM in other northern areas. In general, the existing refugia are likely to have strongly assisted the colonization of more northerly areas of the forest-tundra ecotone since they provide a seed source and shelter for recruitment of larch regeneration (Kharuk et al. 2013). During range expansion and reconnection of refugia or at contact areas, hybrids form and the tendency for this process to occur is common in many forest tree species and is known for Siberian larch species (Semerikov et al. 2007).
4.3 Absence of northern refugia could possibly explain the current treeline migration lag
The refugial populations may have served as a starting point for rapid colonization of the areas north of the treeline in the early Holocene (Tarasov et al. 2009; Epp et al. 2018). This explains the existence ofLarix in the far north at that time (Bigelow 2003), although the migration rates at the treelines were slow. Today the initial situation is different. The climate cooling during the Little Ice Age (LIA; extending from the 16th to the 19thcenturies) negatively impacted tree population densities and caused range contraction, while the enhanced recruitment in the twentieth century has not been of sufficient magnitude to compensate for this range contraction (MacDonald et al. 2008). As a result, there are currently no refugial populations in northern Siberia, as was most likely the case in the early Holocene. This fact could possibly explain why the current treeline advance is lagging behind climate warming. However, if individual trees establish themselves in the tundra area ahead of the treeline in the future, they could be the initial spark for rapid dispersal of the boreal coniferous forest. If the progressive forest expansion keeps pace with climate change in the future, as various studies assume (MacDonald et al. 2008; Pearson et al. 2013; Kruse and Herzschuh 2022), the habitats of tundra are threatened and could recede or disappear completely. This knowledge can be implemented in simulation models such as LAVESI (Larix vegetation simulator) (Kruse et al. 2016; 2019).
5 Conclusions and outlook
We inferred spatial distribution patterns of the genetic variability of Siberian larches by GBS. The data are best explained by three and four genetic groups. However, from an ecological point of view, a differentiation of five to six clusters has the potential to reveal admixture regions and not just the main areas in the given region but even, for example, the emergence of two clusters belonging to the same species but differing from one another. According to Bobrov’s taxonomic system (1972), the four statistically verified main clusters match well with the expected distinction into the three Siberian larch speciesL. sibirica , L. gmelinii, and L. cajanderi from Western to Eastern Eurasia. The most eastern cluster is in Chukotka and seems to be another aggregation of L. cajanderi . Furthermore, the geographical barriers correspond to the habitat zones of the different species.
Our aim was to answer the question of whether refugia existed in northern areas during the LGM and to get an idea of the temporal classification concerning possible demographic events. Altogether the ABC supports a scenario whereby the present Siberian larch populations have survived the LGM in refugia in the north, rather than migrating in the postglacial from the south. The presence of northern LGM refugia may explain the early existence of larches in the far north in the Holocene and their dominance until today. In contrast to the past situation, there are no northern refugia today, which could delay the treeline advance to the north despite climate warming.
The results of this study provide a better understanding on how refugial populations contribute to the treeline migration of Siberian larches. Furthermore, cluster analysis could be used to search for possible refugial populations on a small scale for conservation purposes. Additionally, more complex scenarios, complementary to the most probable scenario detected in the present study, could be analyzed using the ABC method.
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