Southernmost Distribution Limit Research Articles

The northern wheatear is reducing its distribution in its southernmost European range and moving to higher altitudes

Under the current pattern of climate change, mountain bird populations are generally shifting their ranges to higher elevations, tracking their climatic optima. Nevertheless, space limitations at high altitudes constrain mountain species' resilience to climatic change, making them particularly vulnerable. In extreme cases, the climatic niche of some species can move beyond mountaintops, ultimately driving such species to extinction. This study presents the case of the northern wheatear Oenanthe oenanthe in mainland Spain and compares its breeding distribution from 2003 to 2022. Spain, where the species mostly occupies mountain areas, represents its southernmost distribution limit in Europe. We built environmental favourability models using information from the two most recent Spanish bird atlases and a set of climatic, topographic, human activity and lithological variables to determine the factors affecting the occurrence of the species. The influence of climate compared to all other factors was obtained using variation partitioning analysis. The northern wheatear has suffered a strong reduction (67%) in occupied areas and also in favourability throughout mainland Spain (especially in the south) where climate change may have far‐reaching consequences, including local extinctions. Climate explained more than 90% of the variation in the model obtained for 2022. Interestingly, the occupied areas were, on average, 100 m higher in 2022 than in 2003. If the effects of climate warming persist, the northern wheatear will likely disappear in its southernmost distribution limit, being the Sierra de las Nieves National Park population the most vulnerable in mainland Spain. For this reason, it is necessary to implement monitoring programmes for northern wheatear populations, with priority for mountain systems wherein they already occupy higher elevations, as well as further studies on other mountain species.

The genus Boreomysis G.O. Sars, 1869 (Peracarida, Mysida, Mysidae) in western Mexico.

Two species of the genus Boreomysis, B. arctica (Kryer, 1861) and B. californica Ortmann, 1894, were collected in western Mexico. In the eastern Pacific, Boreomysis arctica had previously been recorded from the Bering Sea to southern California. New records in Mexico extend its southernmost distribution limit to 172533 N, off SW Mexico, including samples collected off the west coast of the Baja California Peninsula and the Gulf of California (25 specimens in 12 samples). In the eastern Pacific, B. californica has been reported from Alaska to Peru, with its type locality in the Central Gulf of California. During this study, it was by far the most frequently species collected (343 specimens in 58 samples) off the west coast of the Baja California Peninsula in the Gulf of California, and off SW Mexico (south to 165800 N).

Modeling and updating the occurrence of Aedes aegypti in its southern limit of distribution in South America

Aedes aegypti is the main vector of arboviruses in the world. This mosquito species is distributed from tropical to temperate regions. In Argentina, it has been reported in 20 out of 23 provinces and reaches its southernmost distribution in the world. Its distribution and persistence are affected by meteorological, demographic and environmental factors, such as temperature, precipitation, and population. The aim of this study was to update and model the occurrence of Aedes aegypti in its southern limit of distribution in Argentina. To this end, a total of 37 sites were inspected in La Pampa and Río Negro provinces. Generalized Linear Models were used to explain the occurrence of Aedes aegypti based on meteorological, environmental and demographic variables. Aedes aegypti was found in 11 cities of La Pampa province where it had not been previously reported, but was not found in any of the cities evaluated in Río Negro province. The averaged model explaining the occurrence of Aedes aegypti included the minimum temperature, precipitation and interactions between maximum temperature and precipitation as explanatory variables. Although precipitation was statistically significant, other factors such as minimum temperature are also important in modeling the occurrence of Aedes aegypti in its southernmost distribution limit.

Delineating vulnerability to drought using a process-based growth model in Pyrenean silver fir forests

Assessing tree growth patterns and deviations from expected climate baselines across wide environmental gradients is fundamental to determine forest vulnerability to drought. This need is particularly compelling for the southernmost limit of the tree species distribution where hot droughts often trigger forest dieback processes. This is the case of some silver fir (Abies alba) populations located in southwestern Europe (Spanish Pyrenees) which present ongoing dieback processes since the 1980s. We sampled 21 silver fir stands showing different dieback intensity, assessed using defoliation levels, quantified their growth patterns and characterized their responses to climate. Then, we assessed growth deviations from climatic predictions using the process-based Vaganov-Shashkin (VS) growth model. The forests showing most intense dieback, i.e. highest defoliation levels, were mainly located in low-elevation sites of the western Pyrenees. Trees in these stands displayed the lowest growth rates and the highest year-to-year variability in growth and their growth was limited by late-summer evaporative demand. In eastern and central Pyrenees, we detected a mild growth limitation by low soil moisture during the late growing season and positive growth recovery in recent years with respect to a climate baseline. Decreasing growth trajectories were the most common pattern, while rising trends were common in stands with low dieback in eastern and central Pyrenees. Our results portend systematic spatial variability of growth trends across the Pyrenean silver fir populations forming the south-western distribution limit of the species in Europe. Decoupling of growth between eastern and western populations observed in the recent decades suggests contrasting vulnerability to climate change, and more importantly, the decoupling of growth patterns in western clusters could be used as an early-warning signal of impending dieback. Consequently, we foresee future dieback events to have more detrimental effects in the western compared with the eastern Pyrenees.

Predator-prey collapses at the edge of predator distribution: the case of clupeids and common guillemots (Uria aalge) in NW Iberia

The spatial structuring of seabird populations makes individual colonies very dependent on regional factors. That is especially the case in small edge populations located far from large colonies. We analysed retrospectively the poorly known collapse, some 50 years ago (around 1962-1973), of a relatively small population of breeding common guillemots (Uria aalge) located at their southernmost limit of distribution in Europe (NW Iberia). We assumed that guillemots behaved locally as facultative specialists in small pelagic fish due to the occurrence of a strong clupeid fishery, and we studied the association between the guillemot collapse and annual regional landings of sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus), used as a proxy of total stock size. The overall relationship between ln-transformed guillemot counts (May-June) and ln-transformed sardine landings (May-October) was stronger than the fit on untransformed variables (r2 =0.52 vs. 0.27), indicating an exponential relationship between the non-transformed variables. This relationship was somewhat stronger and linear after the collapse, when only a few tens of guillemots remaining (r2 =0.60). A strong regime shift in sardine landings was detected in 1968 and also in anchovy landings in 1969. The overall relationship between guillemot numbers and anchovy landings was linear and strong (r2 =0.72) but completely dependent on the large 1960s estimate of guillemots. However, no relationship was found between guillemot numbers and anchovy landings (April-June) after the guillemot collapse. The most likely period for the guillemot collapse was therefore 1968-1970, as seabird colonies are known to collapse immediately after their staple prey crashes. Local guillemot colonies were not subsequently rescued by immigration and have remained empty or almost empty until present, showing the high sensitivity of edge populations to environmental variability at the regional scale.

Revision of the genus Eotrechus Kirkaldy (Hemiptera, Heteroptera, Gerridae), with descriptions of six new species

Species of the Asian genus Eotrechus Kirkaldy, 1902 (Hemiptera, Heteroptera, Gerromorpha, Gerridae, Eotrechinae) are atypical water striders because they live on the near-vertical rock surfaces of waterfalls. The taxonomy of the genus is revised and six new species are described: E. steinerisp. nov., E. kerberossp. nov., E. thaisp. nov., E. konkakinhsp. nov., E. boukalisp. nov., and E. andersenisp. nov. Supplementary descriptions of the apterous female and macropterous male of E. fuscus Basu, Chandra & Venkatesan, 2017 are provided. Additional and updated distribution data of species of Eotrechus are reported. Eotrechus fuscus is reported from Myanmar for the first time; E. brevipes is reported from Nepal for the first time; E. hygropetricus Andersen, 1982 is reported from southern Thailand for the first time, which extends the southernmost limit of the known distribution of the genus. A revised key to the species of Eotrechus is also provided.

Seagrasses of West Africa: New Discoveries, Distribution Limits and Prospects for Management

The onset of a major seagrass initiative in West Africa enabled important seagrass discoveries in several countries, in one of the least documented seagrass regions in the world. Four seagrass species occur in western Africa, Cymodocea nodosa, Halodule wrightii, Ruppia maritima and Zostera noltei. An area of about 62,108 ha of seagrasses was documented in the studied region comprising seven countries: Mauritania, Senegal, The Gambia, Guinea Bissau, Guinea, Sierra Leone and Cabo Verde. Extensive meadows of Zostera noltei were recorded for the first time at Saloum Delta, Senegal, which represents the new southernmost distribution limit of this species. This paper also describes the seagrass morphology for some study areas and explores the main stressors to seagrasses as well as conservation initiatives to protect these newly documented meadows in West Africa. The produced information and maps serve as a starting point for researchers and managers to monitor temporal and spatial changes in the meadows’ extent, health and condition as an efficient management tool.

Does environmental adaptation or dispersal history explain the geographical distribution of Ixodes ricinus and Ixodes persulcatus ticks in Finland?

In Finland, the distribution area of the taiga tick, Ixodes persulcatus (Schulze, 1930), is nested within a broader area of distribution of a congeneric species, the sheep tick, Ixodes ricinus (Linnaeus, 1758) (Acari: Ixodidae). We assess whether distinct environmental adaptations or dispersal history provides a more parsimonious explanation for the differences in the distributions of the two common and medically important ixodids in Finland. We used an innovative spatially constrained randomization procedure to analyze whether crowdsourced occurrence data points of the two tick species had statistically different associations with any of the 28 environmental variables. Using points of presence in a region of species co-occurrence, we built Maxent models to examine whether environmental factors or dispersal history could explain the absence of I. persulcatus in a part of the range of I. ricinus in Finland. Five environmental variables-number of inhabitants, road length, elevation above sea level, proportion of barren bedrock and boulders, and proportion of unsorted glacial deposits-were significant at p ≤ .05, indicating greater between-species difference in original than in the randomized data. Of these variables, only the optimum value for unsorted glacial deposits was higher for I. persulcatus than for I. ricinus. Maxent models also predicted high relative habitat suitability (suitability >80%) for I. persulcatus south of its current, sharply bounded distribution range, suggesting that the species has not fulfilled its distribution potential in Finland. The two most common and medically relevant ixodids in Finland may colonize habitats with different environmental conditions. On the contrary, the recent establishment and ongoing dispersion of I. persulcatus in Fennoscandia rather than environmental conditions cause the southernmost distribution limit of the species in Finland.

Weak genetic differentiation but strong climate-induced selective pressure toward the rear edge of mountain pine in north-eastern Spain

Local differentiation at distribution limits may influence species' adaptive capacity to environmental changes. However, drivers, such gene flow and local selection, are still poorly understood. We focus on the role played by range limits in mountain forests to test the hypothesis that relict tree populations are subjected to genetic differentiation and local adaptation. Two alpine treelines of mountain pine (Pinus uncinata Ram. ex DC) were investigated in the Spanish Pyrenees. Further, an isolated relict population forming the species' southernmost distribution limit in north-eastern Spain was also investigated. Using genotyping by sequencing, a genetic matrix conformed by single nucleotide polymorphisms (SNPs) was obtained. This matrix was used to perform genotype-environment and genotype-phenotype associations, as well as to model risk of non-adaptedness. Increasing climate seasonality appears as an essential element in the interpretation of SNPs subjected to selective pressures. Genetic differentiations were overall weak. The differences in leaf mass area and radial growth rate, as well as the identification of several SNPs subjected to selective pressures, exceeded neutral predictions of differentiation among populations. Despite genetic drift might prevail in the isolated population, the Fst values (0.060 and 0.066) showed a moderate genetic drift and Nm values (3.939 and 3.555) indicate the presence of gene flow between the relict population and both treelines. Nonetheless, the SNPs subjected to selection pressures provide evidences of possible selection in treeline ecotones. Persistence in range boundaries seems to involve several selective pressures in species' traits, which were significantly related to enhanced drought seasonality at the limit of P. uncinata distribution range. We conclude that gene flow is unlikely to constrain adaptation in the P. uncinata rear edge, although this species shows vulnerability to future climate change scenarios involving warmer and drier conditions.

Diversity and Distribution of Fiddler Crabs (Crustacea: Brachyura: Ocypodidae) in Vietnam.

Based on recently collected material and records in the literature, 14 species of fiddler crabs (Crustacea: Ocypodidae: Gelasiminae) are reported from Vietnam. DNA barcoding analyses using the mitochondrial gene COI (cytochrome c oxidase subunit I) was performed to identify examined materials and their precise distributional range. Thirteen species-level taxa are identified and, with the exception of Galsimus borealis and G. vocans, have minimum interspcific divergences of at least 7.27%. The identified species include seven species of Tubuca Bott, 1973, three of Austruca Bott, 1973 and three of Gelasimus Latreille, 1817, and one Paraleptuca Bott, 1973. Two new records of Vietnam are herein reported: Tubuca rhizophorae and T. dussumieri. The southernmost distribution limits of East Asian G. borealis, T. acuta and T. arcuata are in northern Vietnam, A. lactea in central Vietnam, whereas northernmost limit of Southeast Asian T. rhizophorae and T. forcipata in southern Vietnam. A dichotomous key to identify the 14 Vietnamese species is provided.

Climate-Change Impacts on the Southernmost Mediterranean Arctic-Alpine Plant Populations

Human-induced climate- and land-use change have been affecting biogeographical and biodiversity patterns for the past two centuries all over the globe, resulting in increased extinction and biotic homogenization rates. High mountain ecosystems are more sensitive to these changes, which have led to physiological and phenological shifts, as well as to ecosystem processes’ deformation. Glacial relicts, such as arctic-alpine taxa, are sensitive indicators of the effects of global warming and their rear-edge populations could include warm-adapted genotypes that might prove—conservation-wise—useful in an era of unprecedented climate regimes. Despite the ongoing thermophilization in European and Mediterranean summits, it still remains unknown how past and future climate-change might affect the distributional patterns of the glacial relict, arctic-alpine taxa occurring in Greece, their European southernmost distributional limit. Using species distribution models, we investigated the impacts of past and future climate changes on the arctic-alpine taxa occurring in Greece and identified the areas comprising arctic-alpine biodiversity hotspots in Greece. Most of these species will be faced with severe range reductions in the near future, despite their innate resilience to a multitude of threats, while the species richness hotspots will experience both altitudinal and latitudinal shifts. Being long-lived perennials means that there might be an extinction-debt present in these taxa, and a prolonged stability phase could be masking the deleterious effects of climate change on them. Several ex situ conservation measures (e.g., seed collection, population augmentation) should be taken to preserve the southernmost populations of these rare arctic-alpine taxa and a better understanding of their population genetics is urgently needed.

Projections of surface air temperature required to sustain permafrost and importance of adaptation to climate change in the Daisetsu Mountains, Japan

Permafrost is known to occur in high mountainous areas such as the Daisetsu Mountains in Japan, which are located at the southernmost limit of the permafrost distribution in the world. In this study, areas with climatic conditions suitable for sustaining permafrost in the Daisetsu Mountains are projected using bias-corrected and downscaled climate model outputs and statistical relationships between surface air temperatures and permafrost areas. Using freezing and thawing indices, the size of the area in the Daisetsu Mountains where climatic conditions were suitable for permafrost were estimated to be approximately 150 km2 in 2010. Under the RCP8.5 scenario, this area is projected to decrease to about 30 km2 by 2050 and it is projected to disappear by around 2070. Under the RCP2.6 scenario, the area is projected to decrease to approximately 20 km2 by 2100. The degradation of mountain permafrost could potentially affect the stability of trekking trails due to slope displacement, and it may also have deleterious effects on current alpine ecosystems. It is therefore important to accurately monitor changes in the mountain ecosystem environment and to implement measures to adapt to an environment that is projected to change significantly in the future.

Genetic Diversity and Structure of Rear Edge Populations of Sorbus aucuparia (Rosaceae) in the Hyrcanian Forest.

Sorbus aucuparia (Rosaceae) is a small tree species widely distributed in Eurasia. The Hyrcanian forest is the southernmost distribution limit of this species. Severe habitat degradation and inadequate human interventions have endangered the long-term survival of this species in this region, and it is necessary to develop and apply appropriate management methods to prevent the loss of its genetic diversity. In this study, we used 10 SSR markers in order to evaluate the genetic diversity of this taxon. Leaf samples were collected from five known populations of S. aucuparia throughout its distribution area in the Hyrcanian forest. Expected heterozygosity ranged from 0.61 (ASH) to 0.73, and according to the M-ratio, all populations showed a significant reduction in effective population size, indicating a genetic bottleneck. Global FST was not statistically significant and attained the same values with and without excluding null alleles (ENA) correction (FST = 0.12). Bayesian analysis performed with STRUCTURE defined two genetic clusters among the five known populations, while the results of discriminant analysis of principal components (DAPC) identified three distinct groups. The average proportion of migrants was 22. In general, the gene flow was asymmetrical, with the biggest differences between immigration and emigration in Barzekoh and Asbehriseh. The Mantel test showed that there was no significant correlation between genetic distance (FST) and geographic distance in S. aucuparia. The best pathway for theoretical gene flow is located across the coast of the Caspian Sea and significant spatial autocorrelation was observed in only one population. In order to reduce the extinction risk of very small and scattered populations of S. aucuparia in the Hyrcanian forest, it is very important to establish and/or enhance the connectivity through habitat restoration or genetic exchange.

A sliver of the past: The decimation of the genetic diversity of the Mexican wolf.

The endangered Mexican wolf (Canis lupus baileyi) is known to carry exceedingly low levels of genetic diversity. This could be (i) the result of long-term evolutionary patterns as they exist at the southernmost limit of the species distribution at a relatively reduced effective size, or (ii) due to rapid population decline caused by human persecution over the last century. If the former, purifying selection is expected to have minimized the impact of inbreeding. If the latter, rapid and recent declines in genetic diversity may have resulted in severe fitness consequences. To differentiate these hypotheses, we conducted comparative whole-genome analyses of five historical Mexican wolves (1907-1917) and 18 contemporary Mexican and grey wolves from North America and Eurasia. Based on whole-genome data, historical and modern Mexican wolves together form a discrete unit. Moreover, we found that modern Mexican wolves have reduced genetic diversity and increased inbreeding relative to the historical population, which was widespread across the southwestern United States and not restricted to Mexico as previously assumed. Finally, although Mexican wolves have evolved in sympatry with coyotes (C. latrans), we observed lower introgression between historical Mexican wolves and coyotes than with modern Mexican wolves, despite similarities in body size. Taken together, our data show that recent population declines probably caused the reduced level of genetic diversity, but not the observed differentiation of the Mexican wolves from other North American wolves.

Living on the edge: reproductive cycle of a boreal barnacle at its southernmost distribution limit

Living on the edge: reproductive cycle of a boreal barnacle at its southernmost distribution limit

Phylogeography of Otaria flavescens (Carnivora: Pinnipedia): unravelling genetic connectivity at the southernmost limit of its distribution

Abstract The Pleistocene glacial period shaped the current genetic structure of numerous species. The last glacial dynamics has been proposed to have split the South American sea lion, Otaria flavescens, into two Evolutionarily Significant Units (ESUs), one on each side of the continent. However, previous studies have not provided genetic information on colonies found along 3000 km of coastline of the southernmost limit of the species distribution, where gene flow could occur. We conducted an exhaustive phylogeographical analysis of O. flavescens using a mtDNA marker, including, for the first time, data from colonies living south of latitude 45° S, in the Argentinian provinces of Santa Cruz and Tierra del Fuego. Our results indicated the presence of five Conservation Units across the distribution range of O. flavescens and suggest that the Patagonian population must have expanded about 150 000 BP. We found evidence for gene flow across the entire species range, supporting a scenario of secondary contact in Tierra del Fuego where representatives of the oldest lineages coexist. The presence of gene flow between oceans leads us to reject the assumption of complete reciprocal monophyly for mtDNA between the presumed ESUs, suggesting that the species constitutes a single Evolutionarily Significant Unit.

Pelagic amphipods (Crustacea: Amphipoda: Hyperiidea) in western Mexico. 3. Family Lestrigonidae.

Thirteen species belonging to five genera of pelagic amphipods of the family Lestrigonidae were collected during a deep-water survey off western Mexico: four species of Hyperietta, two species of Hyperioides, five species of Lestrigonus, and one species each of Phronimopsis and Themistella. This represents 93% of all known species of Lestrigonidae world-wide. A total of 1092 specimens were obtained in 40 sampling localities in the Gulf of California (35) and off southwestern Mexico (5). Although sampling methodology was rather heterogeneous, two species, Hyperioides sibaginis and Lestrigonus bengalensis, were particularly abundant (336 and 562 specimens) and frequent (present in 24 and 31 localities) in the samples, representing 82% of the total catch. The southernmost distribution limit of Hyperietta stebbingi is extended to off SW Mexico. Of the 13 species that were collected during this survey, 10 and 8 species co-occurred at two sampling localities in the SE Gulf of California.

Factors controlling the buildup of humus and particulate organic matter in European beech and Scots pine stands at their southernmost distribution limits (Moncayo Massif, Spain)

Forests of European beech (Fagus sylvatica) and Scots pine (Pinus sylvestris) in the Iberian Peninsula contain some of the southernmost localities of these species and thus may display highly sensitive responses to environmental variations as a result of subjection to particular climate stresses. In this study, we investigated the effects of elevation and dominant tree species (beech vs. pine) on the buildup of organic matter (OM) and nutrients within the humus (O layers) and particulate organic matter (POM) of the topsoil (Ah layer) in centennial Scots pine forests compared to that in natural beech forests along an altitudinal gradient from 1100 to 1600 m.a.s.l. in the Moncayo Massif (northern Spain). The contents of nonstructural, hemicellulose, cellulose, lignin-like, soluble ash and free lipid fractions and of C and major nutrients (N, P, S, K, and Ca) were analyzed and related to the buildup of OM within individual layers, which we used as an inverse surrogate for decomposition rates. The stocks of the studied variables in the humus and the POM were examined in relation to elevation and tree species. The highest storage values were found in the Scots pine sites at higher elevations in the form of thick layers of strongly decomposed organic matter (OH layers) and in the POM. Climate, as it related to elevation, was found to be the major driver of the early and intermediate stages of decomposition (represented by humus) but showed little effect on POM, which was more strongly affected by the tree species. The buildup of OM has its largest correlation (negative) with the content of Ca (which was higher under beech vegetation and decreased with increasing elevation), suggesting an effect of Ca in improving habitat conditions for biological activity.

Biological traits of the spot prawn Pandalus platyceros Brandt, 1851 (Decapoda, Caridea) at its southernmost distribution limit

Abstract The spot prawn fishery, targeting Pandalus platyceros, has failed to prove commercially viable on the Mexican Pacific coast. A survey was carried out off the northwestern coast of Baja California, at depths of 140-180 m. The objective was to describe the biological traits of the species to test the hypothesis that warmer water stimulates higher growth parameters. An average model was estimated that anticipated an inverted exponential growth curve, consequently the growth parameters of the Von Bertalanffy model were obtained as , carapace length (CL) and . Being a protandric hermaphrodite, the transition size was estimated at 40.8 mm CL. The results challenged the hypothesis that P. platyceros grows faster in temperate environments than in cold environments.

Thermal responses of two sub-Antarctic notothenioid fishes, the black southern cod Patagonotothen tessellata (Richardson, 1845) and the Magellan plunderfish Harpagifer bispinis (Forster, 1801), from southern South America

The Notothenioidei are a typical example of stenothermal fishes since most species have evolved and lived in Antarctic waters, where the water temperature is low and stable. This fact enabled them to evolve physiological characteristics related to cold. Nevertheless, some species came out of Antarctic waters a few million years ago and coped with more variable thermal regimes. This work aims to determine the thermal tolerance and preference of two sub-Antarctic notothenioid species found in Southern South America, Patagonotothen tessellata and Harpagifer bispinis, adding valuable information about thermal adaptation mechanisms. Experiments were conducted after exposing their juveniles for three weeks at 4, 7, 10 and 12 °C. Their thermal tolerance limits were established using the Critical Thermal Methodology and their acute thermal preferenda, employing a horizontal thermal gradient tank. Fishes acclimated to different exposure temperatures had small to intermediate thermal tolerance polygons (P. tessellata: 593.85°C2, H. bispinis: 475.40 °C2) and positive relationships between preferred and acclimation temperatures. The Final Temperature Preferenda were estimated to be 14.25 °C for P. tessellata and 13.05 °C for H. bispinis, allowing to characterize them as cold eurythermal species, with P. tessellata more tolerant to heat and H. bispinis more tolerant to cold. Their different thermal sensitivities are in agreement with their different thermal histories and distributions. In a climate change context, the increase of sea surface temperatures is likely to reduce the northern boundaries of their distributions. Conversely, it can potentially enhance both species’ performances at their southernmost distribution limits since those environments are cooler than their maximum thermal tolerances.