Range Of Species Research Articles

Cover Image

COVER PHOTO: A mother toad‐headed agama (Phrynocephalus vlangalii) with her children. P. vlangalii is a species of viviparous agamid lizard endemic to the Qinghai‐Tibet Plateau in China. This lizard lives in burrows at high elevations of 2,000 to 4,600 meters. They need to adapt to a series of harsh environments, including low oxygen levels, low temperatures, and strong ultraviolet radiation. Jiang et al. (this issue; Article: e4414; doi: 10.1002/ecy.4414) collected active and resting body temperatures of P. vlangalii and conducted physiological experiments and mechanistic modeling to assess how resting status determines physiological constraints within a species' distribution range and how climate change drives changes in these physiological constraints. Photo credit: Zhong‐Wen Jiang. image

Assessing the climatic niche changes and global invasion risk of Solanum elaeagnifolium in relation to human activities

As an invasive plant, Solanum elaeagnifolium has posed a serious threat to agriculture and natural ecosystems worldwide. In order to better manage and limit its spread, we established niche models by combining distribution information and climate data from the native and invasive ranges of S. elaeagnifolium to analyze its niche changes during its colonization. Additionally, we evaluated its global invasion risk. Our results showed that the distribution of S. elaeagnifolium is affected by temperature, precipitation, altitude, and human activities. Solanum elaeagnifolium exhibits different degrees of niche conservatism and niche shift in different invasion ranges.During the global invasion of S. elaeagnifolium, both the niche shift and conservatism were observed, however, niche shift was particularly significant due to the presence of unoccupied niches (niche unfilling). Solanum elaeagnifolium generally occupied a relatively stable niche. However, a notable expansion was observed primarily in Europe and China. In Australia and Africa, its niche largely remains a subset of its native niche. Compared to the niche observed in its native range, its realized niche in China and Europe has shifted toward lower temperature and higher precipitation levels. Conversely, in Africa, the niche has shifted toward lower precipitation levels, while in Australia, it has shifted toward higher temperature. Our model predicted that S. elaeagnifolium has high invasion potential in many countries and regions. The populations of S. elaeagnifolium in China and Africa have reached the adapted stage, while the populations in Australia and Europe are currently in the stabilization stage. In addition, our research suggests that the potential distribution of S. elaeagnifolium will expand further in the future as the climate warms. All in all, our study suggests that S. elaeagnifolium has high potential to invade globally. Due to its high invasive potential, global surveillance and preventive measures are necessary to address its spread.

Assessing the Impacts of Adaptation to Native-Range Habitats and Contemporary Founder Effects on Genetic Diversity in an Invasive Fish.

Species invading non-native habitats can cause irreversible environmental damage and economic harm. Yet, how introduced species become widespread invaders remains poorly understood. Adaptation within native-range habitats and rapid adaptation to new environments may both influence invasion success. Here, we examine these hypotheses using 7058 SNPs from 36 native, 40 introduced and 19 farmed populations of tench, a fish native to Eurasia. We examined genetic structure among these populations and accounted for long-term evolutionary history within the native range to assess whether introduced populations exhibited lower genetic diversity than native populations. Subsequent to infer genotype-environment correlations within native-range habitats, we assessed whether adaptation to native environments may have shaped the success of some introduced populations. At the broad scale, two glacial refugia contributed to the ancestry and genomic diversity of tench. However, native, introduced and farmed populations of admixed origin exhibited up to 10-fold more genetic diversity (i.e., observed heterozygosity, expected heterozygosity and allelic richness) compared to populations with predominantly single-source ancestry. The effects of introduction to a new location were also apparent as introduced populations exhibited fewer private alleles (mean = 9.9 and 18.9 private alleles in introduced and native populations, respectively) and higher population-specific Fst compared to native populations, highlighting their distinctiveness relative to the pool of allelic frequencies across tench populations. Finally, introduced populations with varying levels of genetic variation and similar genetic compositions have become established and persisted under strikingly different climatic and ecological conditions. Our results suggest that lack of prior adaptation and low genetic variation may not consistently hinder the success of introduced populations for species with a demonstrated ability to expand their native range.

Characterization of the reproductive strategy of invasive Round Goby (Neogobius melanostomus) in the Upper DanubeRiver.

Originating from the Black and Caspian seas, the Round Goby (Neogobius melanostomus) has become one of the most successful invaders of freshwater ecosystems. In this study, we provide a characterization of the reproductive strategy of an established population of Round Gobies in the Upper Danube river including sex ratio, fluctuations of gonadosomatic index (GSI), analysis of timing of spawning as well as of clutch and egg size. We compare these results to other studies from the native and invaded range. In the Danube, the Round Goby population was found to be female dominated, however fluctuations in magnitude of female bias were observed between months. Monitoring of the population across 1.5 years revealed that GSI was highest from April to June, while lowest values were observed in August and September. Using time-series analysis, a delayed effect of temperature on GSI was found for females and males, while a quicker response of GSI levels to photoperiod and discharge was observed for females. GSI increased with body size for females and eggs were found to be significantly larger in May, however clutch sizes did not differ between months. Results of a literature review revealed great differences in timing and length of spawning season as well as sex ratio between populations throughout the distribution range, which can probably be explained by climatic and photoperiodic conditions together with the time since invasion and the high plasticity of Round Gobies.

Morphological and molecular characterisation of Mordellistenapeloponnesensis Batten, 1980 (Coleoptera, Mordellidae), with first records from Italy and Turkey.

Mordellistenapeloponnesensis Batten, 1980, previously known from Cyprus and Greece, is reported from Italy and Turkey for the first time. The species is redescribed based on type specimens and additional material from its entire known distributional range. Eighteen DNA barcoding sequences of M.peloponnesensis from Greece, Cyprus, and Italy were generated, and genetic variability across the sampling localities was examined. Three mitochondrial haplotypes were detected within M.peloponnesensis. Specimens from mainland Italy share the same haplotype as those from Rhodes and Cyprus, whereas Sardinian specimens exhibit a distinct haplotype. The third haplotype is represented by one specimen from Cyprus. The DNA barcoding sequences of M.peloponnesensis were compared with those of the morphologically allied M.gemellata Schilsky, 1898, and M.pyrenaea Ermisch, 1966, to reveal the phylogenetic relationships between the species.

Ni0.25Cu0.5Sn0.25 Nanometallic Glasses As Highly Efficient Catalyst for Electrochemical Nitrate Reduction to Ammonia

AbstractElectrochemical nitrate reduction to ammonia (NRA) is a promising approach for alleviating energy crisis and water pollution. Current NRA catalysts are challenged to simultaneously improve the rate of the adsorption and desorption processes to further increase the total activity due to the Brønsted−Evans−Polanyi (BEP) relationships. Herein, a two‐step Joule heating method is utilized for the preparation of Ni0.25Cu0.5Sn0.25 nanometallic glass containing synergistic catalytic sites to simultaneously enhance the adsorption and desorption processes. Kelvin probe force microscopy reveals a pronounced oscillatory behavior in the surface potential of Ni0.25Cu0.5Sn0.25 nanometallic glass, which is an important feature of the synergistic catalytic site, and an empirical formula is proposed to quantitatively characterize its oscillatory characteristic. In situ electrochemical Raman spectroscopy indicates the promotion of nickel and tin atoms for nitrate adsorption and ammonia desorption processes, respectively. DFT calculations demonstrated that Ni0.25Cu0.5Sn0.25 presents a wide range of adsorption energy distributions to favor the multisite synergistic catalysis. The present work provides new ideas for the design and understanding of highly active NRA catalysts.

Topology optimization of regenerative cooling structures under high Reynolds number flow with variable thermo-physical properties

In the design of thermal protection system for hypersonic vehicles, the efficient convective heat transfer within regenerative cooling channels plays a critical role in maintaining their thermal performance. This study focuses on the topology optimization (TO) of the cooling channels that operate at high Reynolds number and have variable thermo-physical properties. A novel approach is introduced to enhance the density-based method by incorporating an effective artificial force correction and tabulating the temperature-dependent thermal properties. Several topology-optimized cooling layouts were generated to minimize the maximum temperature within the design domain while considering different power dissipation constraints. This confirms the validity of the proposed lumped correction term in momentum equation accounting both viscous and convective body forces. Regarding the thermal performance of the optimized layouts, the staggered arrangement of cellular ribs in the TO channel was found to induce multiple flow separations and promote turbulent mixing, thereby improving heat transfer efficiency and mitigating the thermal acceleration effect. Conjugate heat transfer calculations revealed that the optimal topology-optimized channel achieved a 24.3% improvement in overall thermal performance while being 15.3% lighter than the straight channel design. Furthermore, the optimal topology optimized layout consistently outperformed the straight channel design by 6.6% to 24.3% in terms of the overall thermal performance across a wide range of heat flux distribution and mass flow rate conditions. This investigation highlights the effectiveness of topology optimization in designing regenerative cooling structures featuring high Reynolds number hydrocarbon thermal fluid flow.

The marriage between stable isotope ecology and plant metabolomics - new perspectives for metabolic flux analysis and the interpretation of ecological archives.

Even though they share many thematical overlaps, plant metabolomics and stable isotope ecology have been rather separate fields mainly due to different mass spectrometry demands. New high-resolution bioanalytical mass spectrometers are now not only offering high-throughput metabolite identification but are also suitable for compound- and intramolecular position-specific isotope analysis in the natural isotope abundance range. In plant metabolomics, label-free metabolic pathway and metabolic flux analysis might become possible when applying this new technology. This is because changes in the commitment of substrates to particular metabolic pathways and the activation or deactivation of others alter enzyme-specific isotope effects. This leads to differences in intramolecular and compound-specific isotope compositions. In plant isotope ecology, position-specific isotope analysis in plant archives informed by metabolic pathway analysis could be used to reconstruct and separate environmental impacts on complex metabolic processes. A technology-driven linkage between the two disciplines could allow us to extract information on environment-metabolism interaction from plant archives such as tree rings but also within ecosystems. This would contribute to a holistic understanding of how plants react to environmental drivers, thus also providing helpful information on the trajectories of the vegetation under the conditions to come.

Ovarian traits, spawning pattern and season, length at first maturity, and batch fecundity of Caragobius urolepis (Bleeker, 1852) caught from the Vietnamese Mekong Delta

Caragobius urolepis has a wide range of distribution, from brackish to freshwater water, ranging from India to the Philippines and Fiji, including the Vietnamese Mekong Delta (VMD), where it lives mainly in the coastal regions and plays a vital role in food supply. Our preliminary observation showed that its population tended to decline, but little is known about this species. This study, therefore, was conducted to provide data on the ovarian structure, spawning pattern and season, length at first maturity, and batch fecundity of this fish. The study of 306 specimens caught with trawl nets from April 2022 to March 2023 in two coastal areas of VMD—Dong Hai, Bac Lieu (DHBL), and Dam Doi, Ca Mau (DDCM)—showed that this goby spawned more than once because the oocytes in stage IV ovary showed different stages of development. It could release eggs monthly during a 12-month study, with a peak in July–September since stage IV ovaries were found monthly during the study period and the GSI exhibited a high value in July–September. The fish length at first maturity and batch fecundity varied with the site, as they were 18.8 cm and 3,757-9,187 (6,142±707 SE eggs/female) in DDCM and 20.5 cm and 3,760-11,118 (5,634±750 SE eggs/female) in DHBL. The findings provide valuable foundations for exploiting their resources and serve as a basis for studying artificial reproduction. It is suggested to avoid catching these fish species during the breeding season to ensure sustainable exploitation in the future.

Differential Effects of Chewing Lice on Body Condition across Host Age and Sex in Rough-legged Hawks (Buteo lagopus).

Chewing lice infesting avian hosts can significantly affect host health and fitness. Here, we present quantitative data on host body condition and louse abundance observed from 121 Rough-legged Hawks (Buteo lagopus) sampled across the North American nonbreeding range. Among hawks examined, louse prevalence was 71%, with a mean abundance and intensity of 9.1 and 12.8 lice, respectively. We identified lice as Craspedorrhynchus sp., either Craspedorrhynchus dilatatus or Craspedorrhynchus taurocephalus, dependent on future taxonomic revision of the genus. Female and juvenile hawks had greater louse intensity and prevalence compared with male and adult hawks, respectively. Host body condition, measured as a breast muscle score (keel score), was negatively correlated with louse abundance after controlling for host age and sex. Possible explanations for these patterns include the following: sex-biased louse transfer between adults and nestlings, when female nestlings experience increased transfer loads; body size differences between males and females, when females are larger than males in each life stage; and preening limitations in females and juveniles, when both spend more time hunting and less time preening relative to adult males. Our results corroborate previous studies suggesting that the primary sources of intraspecific variation in louse abundance are host body size and preening limitations.

Damage Index Correlation in Massive Granite-Porous Backfills under Hydromechanical Triaxial Cyclic Loading using Acoustic Emissions and X-ray Computed Tomography

The characterization accuracy of damage indices in massive and porous composite biomaterials under complex osmotic conditions has long been a challenge in engineering materials science. Establishing correlations between damage indices calculated using different methodologies is crucial to validate and improve defects characterizations. In this study, we investigated the damage index correlation of cylindrical massive granite-porous backfill structural composites under both dry and seepage conditions. A series of triaxial cyclic loading and unloading tests were conducted on specimens under hydromechanical conditions with varying osmotic pressures (0, 1, 3, 5, 7, and 9 MPa) and a confining pressure of 9 MPa involving acoustic emission monitoring and phase-based 3D nondestructive CT scanning. The results revealed the AE energy was lower under osmotic conditions compared to dry conditions. The AE Damage Index (), defined by separating tensile and shear microcracking signals, significantly increased from 0.32 to 0.55 with rising osmotic pressure. 3D mesoscopic fracture visualization provided detailed insights into the fractures in granite and backfill interfaces. The number of small-angle cracks increased with osmotic pressure, and the crack inclination distribution range expanded from 63°- 85° to 49°-80°. The average crack angle showed a significant monotonic decrease with increasing osmotic pressure. An Energy Dissipation Damage Index () was defined, and its correlation with the under dry conditions was modeled using a reciprocal function . This novel approach offers insights into the transition points between tensile and shear-dominated failure stages. Furthermore, an exponential function was used to model the correlation between AE Damage Index and Visual Damage Index () under osmotic conditions. The damage indices derived from different methods can be mathematically correlated, providing a comprehensive evaluation of material damage states. The study found the flow transition between different natural-human medias results in different damage mechanisms, which are all obey the correlation law. These damage mechanisms offered constructive recommendations for engineering applications based on varying ratios of osmotic pressure to confining pressure.

Climate change effects on the diversity and distribution of soybean true bugs pests.

Climate change and pests are two major factors in the reduction of global soybean yields. The diversity and geographic distribution of soybean true bug pests vary across soybean production areas worldwide, and climate change impacts are different across species and regions. Therefore, we integrated spatial and temporal predictions at the global scale to predict the impact of global warming on the distribution of 84 soybean true bug pests by the maximum entropy niche model (MaxEnt) under present (1970-2000) and future (2041-2060) scenarios. We produced an ensemble projection of the potential distribution of pests and crop production areas to estimate how and where climate warming will augment the threat of soybean true bug pests to soybean production areas. Our results indicated that Southeast North America, Central South America, Europe and East Asia were the regions with the higher richness of soybean true bug and the most vulnerable areas to invasion threats. Climate change would promote the expansion of the distribution range and facilitate pest movement pole wards, affecting more soybean cultivated areas located in mid-latitudes. Moreover, species with different distribution patterns responded differently to climate change in that large-ranged species tended to increase in occupancy over time, whereas small-ranged species tended to decrease. This result indicates that some pests that have not yet become notable may have the chance to develop into serious pests in the future due to the expansion of their geographical range. Our findings highlight that soybean cultivated regions at mid-latitudes would face general infestations from soybean true bug pests under global warming. These results will further facilitate the formulation of adaptation planning to minimize local environmental impacts in the future. © 2024 Society of Chemical Industry.

Two Palaearctic species of Orthonevra (Diptera: Syrphidae) under the name O. brevicornis

During studying European material of Orthonevra Macquart, 1829 the presence of two widespread species so far interpreted as Orthonevra brevicornis (Loew, 1843) has been discovered. One species occurs from Central Europe into Transcaucasia and agrees with the taxon O. brevicornis of Loew (1843). The second species, O. atlantica Żóralski & Van de Meutter sp. nov., has its main distribution range in the Atlantic parts of Europe and is described here. We provide a redescription of O. brevicornis and designate a neotype from the type locality (Poland, Posen [= Poznań] vicinity). The most important characters for distinguishing males of these two closely related species are in the structures of the terminalia. A diagnosis and identification key to European black-legged Orthonevra species is given as well as new sequences of the cytochrome c oxidase subunit I (COI) of mitochondrial DNA (mtDNA), also known as DNA barcodes, for most of the species of this group. Some new information and changes to the known ranges of Orthonevra species are provided as a result of this revisionary work: O. frontalis (Loew, 1843) is removed from the list of species occurring in Poland; O. gemmula Violovitsh, 1979 is confirmed from Hungary; O. incisa (Loew, 1843) is confirmed from its locus typicus (Poland, Posen vicinity); and O. montana Vujić, 1999 is reported from Poland and Austria for the first time and confirmed for Germany. An explanation for some previous records published as O. plumbago (Loew, 1840) is also given.

Black grouse (Lyrurus tetrix) population status, reasons for decline and potential conservation measures from Western and Central Europe to Fennoscandia: a literature review

The long-term decline of the black grouse population throughout Europe is influenced by many factors that affect populations differently depending on the distribution range, from Central Europe to the Scandinavian countries. Therefore, we analyzed available literature sources to describe the main reasons for the population decline of this species and to suggest conservation measures. In total, 228 pieces of literature from 1955 to 2024 were analyzed for this study. Based on the literature analysis, it is clear that the drivers of population decline differ across the distributional range. In Central Europe, where the population is declining rapidly, habitat loss and forest fragmentation are crucial factors, as is the negative impact of tourism. In Scandinavia, where the population is gradually declining, decreasing breeding success and increasing chick mortality rates are generally considered the main negative factors. However, these factors also affect black grouse populations in Central Europe. It is crucial to acknowledge that a significant proportion of the contributing factors, such as predation and habitat loss, can be attributed to human activities. Therefore, it is necessary to emphasize that environmental protection should work hand in hand with wildlife managers to improve the situation.

Estimates of Effective Number of Breeders Identify Drivers of Decline in Mid-Atlantic Brook Trout Populations.

Brook Trout (Salvelinus fontinalis) populations have experienced marked declines throughout their native range and are presently threatened due to isolation in small habitat fragments, land use changes, and climate change. The existence of numerous, spatially distinct populations poses substantial challenges for monitoring population status (e.g., abundance, recruitment, or occupancy). Genetic monitoring with estimates of effective number of breeders (N b) provides a potentially powerful metric to complement existing population monitoring, assessment, and prioritization. We estimated N b for 71 Brook Trout habitat units in mid-Atlantic region of the United States and obtained a mean N b of 73.2 (range 6.90-493). Our modeling approach tested whether N b estimates were sensitive to differences in habitat size, presence of non-native salmonids, base flow index, temperature, acidic precipitation, and indices of anthropogenic disturbance. We found significant support for three of our hypotheses including the positive influences of available habitat and base flow index and negative effect of temperature. Our results are consistent with presently observed and predicted future impacts of climate change on populations of this cold-water fish. Importantly, these findings support the use of N b in population assessments as an index of relative population status.

Not Everyone Likes Manatees: Fishers' Perceptions Unveil Opportunities and Challenges for Manatee Conservation in the Swamp Complex of Ayapel, Colombia

ABSTRACTGreater Caribbean manatees Trichechus manatus manatus face complex conservation challenges throughout their distribution range, many of which are associated with human factors; and in consequence, the subspecies is considered Endangered (EN) by the International Union for Conservation of Nature (IUCN). Human‐related threats as hunting for meat consumption, collisions with boats, entanglement in fishing nets, pollution and loss of habitat put the Greater Caribbean manatee at risk in Colombia. Studies aiming to describe traditional knowledge and local perceptions provide information that can be useful to design local manatee conservation programmes. We analyse the fishers' perceptions regarding manatees in Ayapel, Córdoba. One hundred semi‐structured interviews were conducted, focusing on personal data, fishing practices, perceptions and knowledge about the manatees. According to the interviews, bycatch is the greatest threat to manatees (56%), followed by pollution (16%), whereas boat collisions (12%), hunting (11%) and drought (5%) are lower ranking threats. Our results demonstrated that participants have a good understanding of manatee biology and ecology. However, a human–manatee conflict was evidenced, as 39% of fishers consider that manatees negatively impact fishing by competition for the fishing resources and/or by damaging fishing gear. We recommend strengthening awareness about the importance of manatee to guarantee an optimal management of this subspecies and its habitat. Fishers should be directly involved in the formulation of management and conservation plans for the subspecies, as their knowledge becomes critical to understanding and mitigating the risks to manatee populations. This work has a clear application to understand the human factor in manatee recovery not only in our study area but also in other locations where manatees share habitat with people, particularly in marginalized human communities.

A Study on the Cutting Characteristics of Bottom Abrasive Grains in Helical Grinding Tools

Helical grinding is crucial for manufacturing small holes in hard-to-machine composite ceramics. This study introduces a geometric model of undeformed chips to analyze the cutting characteristics of abrasive grains on both the bottom and side edges of the tool. It reveals for the first time that the distribution of cutting grains—pure bottom-edge, pure side-edge, and mixed-edge—is influenced by the tool diameter and eccentricity. A novel calculation method for the distribution range (Dp) of pure bottom-edge grains is proposed, demonstrating that using a tool diameter at or below two-thirds of the target hole diameter effectively eliminates pure bottom-edge grains, improving chip evacuation, reducing chip adhesion, and optimizing cutting performance. Experimental validation on small holes in SiCp/Al composites (65% volume fraction) confirmed these findings and provides practical guidance for optimizing cutting parameters and tool design.

New record of Cyanallagma demoiselle Denck, Ehlert & Pinto, 2023 (Odonata, Zygoptera, Coenagrionidae) in southern Brazil

During a field expedition in the municipality of Araquari, state of Santa Catarina, southern Brazil, adults of the damselfly Cyanallagma demoiselle Denck, Ehlert & Pinto 2023 (Odonata, Zygoptera, Coenagrionidae) were collected. This species was recently described and reported to date for the southeastern Atlantic Forest in Brazil. Here, we recorded this species as well as its respective genus for the first time for the state of Santa Catarina, increasing its distribution range of the species to southwards and raising the number of Odonata species recorded for this state to 151.

Why do Eurasian otters eat so few invasive blue crabs?

Invasive species can become key trophic resources for native predators, and thus understanding predator-invasive prey interactions is relevant to assess invasion impacts. The Atlantic blue crab (Callinectes sapidus) has recently invaded coasts and lower stretches of rivers across the Western Palearctic, often establishing very abundant populations. The blue crab is an important prey for a large number of predators within its native range, but predation upon invasive populations is still virtually unknown. The Eurasian otter (Lutra lutra) is a flexible predator, known to incorporate invasive prey into its diet when they become available. We studied otter diet in a small estuary of southern Spain hosting a very abundant blue crab population. The role of the blue crab as otter prey was only anecdotal since, out of 223 spraints (otter faeces) collected between 2021 and 2023, its remains were found in a single one. The avoidance of the blue crab by otters, and also by other predators, seems to be a generalised phenomenon in the invaded range of this crustacean. Predation on invasive blue crabs may require a long-lasting learning process to overcome the crab’s anti-predatory structures and behaviours. Trophic interactions between invasive blue crabs and native predators deserves attention for three main aspects: (i) its implications for the functioning and structure of invaded food webs, (ii) its potential role in limiting the invasion, and (iii) the interest in the generation of new predatory traits among animal populations.

Spatial distribution characteristics of Tibetan Buddhism principal-subordinate monastery systems in the Hehuang region

Tibetan Buddhist monasteries constitute significant cultural heritages of ethnic minorities, evolving into the Principal-Subordinate Monastery System (PSMS) with profound historical and cultural significance. This system exhibits an integrated hierarchical structure in terms of religious dominance, political administration, and cultural diffusion. Existing research primarily focuses on internal humanistic factors such as governance patterns, while there is not any research focused on its patterns and manifestations in spatial distribution. This study aims to elucidate the spatial distribution characteristics of the PSMS in Hehuang region, including the Tibetan Buddhist monasteries from all sects in the Hehuang region since the Song Dynasty It establishes a hierarchical PSMS database based on the affiliation relationships, aimed at storing spatial and property information related to PSMS. The database standardizes the naming and coding of monasteries, and classifies them through hierarchical relationships to ensure data consistency and usability. Finally, the classified and coded monastery data were analyzed by GIS tools to form the PSMS spatial distribution characterisation framework. Results reveal that (1) Monasteries demonstrate notable spatial clustering patterns from both holistic and sectarian perspectives, with density being influenced by the principal monastery’s position. (2) Almost every PSMS exhibits a spatial pattern centered around the principal monastery, with subordinate monasteries clustering within a defined range correlated with the administrative region of the principal monastery. (3) PSMS scale variations are controlled by principal monasteries, exhibiting positive correlations in hierarchical structure, control quantity, and distribution range. (4) The spatial orientation of PSMS correlates with river distribution, while the relationships among mountain ranges require further investigation. These findings provide initial insights into the spatial distribution characteristics of PSMS, confirming the spatial influence of the principal monastery. Besides, this study established an innovative spatial research framework for heritage clusters with multiple types and hierarchies. Thus, this study offers new insights into the spatial distribution of Tibetan Buddhist monastery heritage and presents a framework for further examining the spatial distribution of the Tibetan Buddhist monastery heritage in other regions, as well as other heritage clusters where connected and clustered through religious and cultural ties.