Vascular plant diversity and climate change in the alpine zone of the Snowy Mountains,Australia

This study examines vascular plant species richness along an altitudinal gradient in alpine Australia. Vascular plant composition and soil temperature records were obtained for five summits (from 1729 m to 2114 m a.s.l.) using sampling protocols from the Global Observation Research Initiative in Alpine Environments program. Species richness was examined against altitude, aspect and climatic variables at different spatial scales (10 × 10 cm quadrats, 1 m² quadrats, clusters of 4 * 1 m² quadrats, for the summit area above a line 5 m altitudinally below the summit (the −5 m isoline), for the extended summit down to the −10 m isoline). About 75 taxa (70 species, 5 graminoid genera) were recorded, 9 of which are endemic to the small alpine area of ∼100 km². There were significant linear relationships between species richness and altitude and climatic variables for the top to −5 isolines on the summits. However, there was no consistent pattern for species richness at other spatial scales, altitude, aspect or climatic variables. The proportion of species for the whole summits with localised distributions (local endemics) increased with altitude. Predicted increasing temperatures and reduced snowcover is likely to result in an increase in species richness as shrubs, herbs and introduced weeds become more common at higher altitude. Because Australian alpine areas occur in narrow altitudinal bands with no nival zone, there are no higher altitudinal refuges available for alpine species. Therefore many of these species are likely to be at risk of extinction from climate change.     

Diet selection by European hares (Lepus europaeus) in the alpine zone of the Snowy Mountains, Australia

European hares (Lepus europaeus) are grazers and open grassland specialists that are replaced in mountain areas of their natural range in the northern hemisphere by browsing/intermediate feeding mountain hares (Lepus timidus), but in their introduced range in the southern hemisphere, occupy the alpine zone. We used micro-histological identification of plant fragments and germination of seeds in faecal pellets of L. europaeus from the Snowy Mountains, Australia, to determine diet. We asked whether diet shifted and/or diet breadth expanded in response to seasonally reduced food availability, particularly during winter. If so, did the constraints of food availability in the alpine zone lead to the diet mirroring that of L. timidus in its native alpine habitat. The diet of L. europaeus was dominated by grasses, herbs and shrubs. The main diet items in summer were grasses (70 %) and herbs (28 %). Grasses declined in the diet between summer and autumn when herbs increased to co-dominance, with a further change after establishment of the winter snowpack to a greater preponderance of shrubs (43 % compared with a maximum of 3 % in snow-free months). L. europaeus selected a wider range of plants in winter (59 species compared with 39 in summer) and diet was significantly more variable in winter than in autumn or summer (and in autumn than summer). We concluded that the persistence of L. europaeus in alpine areas of the southern hemisphere is testament to their ability to expand their dietary breadth to occupy mountain climatic zones normally occupied by L. timidus.     

Limited effect of hare grazing and short-term climatic variations on the most common alpine vegetation community in the Snowy Mountains,Australia

Background: In the alpine zone of the Snowy Mountains, grazing by mammals is limited. However, introduced European hare numbers have increased since the 1970s.

Aims: To estimate the density of hares and hence grazing pressure among years. To assess the response of biomass, vegetation height and composition to a cessation of hare grazing.

Methods: We used indices of hare abundance based on spotlighting and counts of hare pellets on a transect. The effect of hare grazing on tall alpine herbfield was assessed by using 15 paired exclosure and control quadrats for six years.

Results: The indices of hare abundance suggested densities similar to those in upland areas of Britain. Grazing did not affect the composition, cover of herbs or graminoids or, for 2010, vegetation height or biomass. Variation in vegetation and hare numbers among years was not correlated with climatic variables. Observations of selective grazing suggested that impacts on vegetation may be localised and restricted to certain species. Prior analyses of hare pellets indicated that hares might spread seed of native and exotic species.

Conclusions: Hares are having no general effect on tall alpine herbfield but may affect certain plant species via selective grazing or by spread of viable seed.     

Pollination biology in the Snowy Mountains of Australia: Comparisons with montane Colorado,USA

Various aspects of the pollination biology of the alpine flora of Kosciusko National Park, NSW, were examined from late December 1983 until the end of March 1984, including flowering phenology, corolla tube lengths, flower colour, ultraviolet reflectance patterns, visitation rates to the flowers and proboscis lengths of the flower-visiting insects. An average of 5.3 species flowered in each of 13, 2 m×2 m montane plots and 5.6 species in the 13 alpine plots. The maximum number in flower simultaneously averaged 4.1 species in the montane and 3.3 in the alpine plots; flowering peaked in mid-January, Corolla tube lengths of the flora averaged 1.73 mm. The most common floral colour was white or predominantly white (40 species), followed by yellow (14 species). Only six of the 38 species (16%) examined had some type of reflectance pattern; the remaining species all absobed ultraviolet. Flies appeared to be the major pollinators. The insects collected in the study area comprised 60 species of Diptera, 33 species of Hymenoptera, and several species each of Lepidoptera and Coleoptera. On average, 14.4% of flowers watched during 379 observation periods (10 min each) were visited. On average, each plant species was visited by 6.4 species of flies, 2.4 species of bees, wasps or sawflies, one species of butterfly or moth and 0.3 species of beetles. Visitation rates increased over the growing season, and were significantly affected by ambient temperature (positively), light levels (positively) and wind speed (negatively). The maximum proboscis length for the 25 most common species of bees was 2.76 mm, but 18 of 51 species of flies had proboscis lengths longer than this. The mean proboscis length for all 25 species of bees was 1.68 mm, and for 51 species of flies was 2.31 mm. Proboscis lengths for flies were positively correlated with the average corolla length for the plant species they visited. For bees, however, the range in proboscis lengths was relatively small and did not show this pattern. There appear to be significant differences between the plant-pollinator community of alpine Australia and other alpine areas where bumblebees are common pollinators. These differences include shorter proboscis and corolla tube lengths, and perhaps an increased diversity and significance of flies as pollinators.     

Linear declines in exotic and native plant species richness along an increasing altitudinal gradient in the Snowy Mountains,Australia

Abstract Declines in plant species richness with increasing altitude are common, but the form of the relationship can vary, with both monotonic decreasing relationships and humped relationship recorded. However, these different richness to altitude relationships may be due to methods that used different plot sizes/areas and survey efforts. To explore native and exotic plant richness along an altitudinal gradient in the Snowy Mountains of Australia, we consistently surveyed plots that were 120 m² in area at 39 sites ranging from 540 to 2020 m. To relate exotic plant richness to disturbance, we surveyed plots at 16 sites along main roads and 23 sites along minor roads and also compared these 39 roadside plots to 120‐m² plots located in undisturbed vegetation adjacent to the roadside (native plant richness was only surveyed in 25 of these 39 adjacent plots). We found a negative linear relationship between total, exotic and native species richness and altitude for plots on the side of main roads (16 sites) and minor roads (23 sites). For adjacent plots negative linear relationships were significant for all measures of species richness except for native species adjacent to major roads. As the pattern occurred for exotics it is less likely to be due to historical constraints on the species pools. The pattern could be influenced by difference in levels of disturbance along the gradient, although any such gradient in disturbance would have to apply to roadside and adjacent plots on major and minor roads. Therefore, it may be due to other factors such as changes in climate along the altitudinal gradient, although additional sampling including direct measures of climatic conditions, soil and disturbance factors would be needed to determine if this was the case.     

Summer frost resistance and freezing patterns measured in situ in leaves of major alpine plant growth forms in relation to their upper distribution boundary

Summer frost resistance and ice nucleation temperatures for 33 alpine plant species were measured in situ to avoid the shortcomings of laboratory tests. Species were selected to investigate the relationship between plant stature and upper distribution boundary, and frost resistance and freezing patterns. The species tested in situ were on average 1.1 K (± 0.2, SE) frost hardier than in laboratory tests. Frost resistance (LT₅₀) ranged from ?4.5 to ?14.6 °C and appeared insufficient to protect against air temperature minima, corroborating reports of natural frost damage. All species tolerated extracellular ice formation (recorded at ?1.9 ± 0.2 °C; E1). Initial frost damage occurred at average temperatures 4.9 K below E1. In 64% of the species a second exotherm (E2) and frost damage were recorded between ?3.7 and ?9.4 °C. In the highest ranging species E2 was not detectable. Frost resistance increased with increasing upper distribution boundary (0.4 K per 100 m), corresponding well with the altitudinal decrease in air temperature minima. No relationship between plant stature and frost resistance was found. Graminoids were significantly frost hardier than other growth forms. Frost survival at high altitudes will depend not only on altitudinal increase in frost resistance but also on freezing avoidance strategies, snow cover protection and a high recuperation capacity.     

Immediate and short‐term responses of bird and mammal assemblages to a subalpine wildfire in the Snowy Mountains,Australia

Abstract Over 35 days in January–February 2003, wildfires burnt across much of the subalpine/alpine landscape of south‐eastern Australia, including about 70% of the land above 1500 m in the Snowy Mountains. At the time of the fire, studies of two subalpine faunal assemblages were being undertaken. The opportunity to resurvey the assemblages was taken in order to examine the immediate impact of fire in an environment where it is uncommon but predicted to occur increasingly with global warming. A study area in the Whites River Valley, where the number of bird species was counted monthly from 1996 to 2003, and weekly in late winter–spring from 2000 to 2003, was burnt in one fire. As well as the expected decrease in the number of individual birds, the fire resulted in an immediate decrease in the number of bird species, unlike in previously studied montane forest, with only the regularly wintering species plus the olive whistler and the ground‐feeding flame robin remaining. During the post‐winter avian immigration, few regular spring migrants appeared on burnt sites despite their nearby presence on the unburnt side of the valley. Five of six small mammal trapping grids were burnt. As with fires at lower altitudes, there was an immediate reduction in mammal numbers on burnt grids following the fire, but in addition, one species, Mastacomys fuscus, declined further in the ensuing 2 months both on burnt and unburnt sites. Numbers of Antechinus swainsonii and Rattus fuscipes stabilized until autumn/winter when there was a further decline due to the unavailability of subnivean space to allow winter foraging, allied with a concentration of fox predation on areas still carrying populations of small mammals.     

Phenology of the environmental weed Achillea millefolium (Asteraceae) along altitudinal and disturbance gradients in the Snowy Mountains, Australia

This study examined the phenology of the weed Achillea millefolium over a growing season in the Snowy Mountains. Vegetative and reproductive characteristics of plants in 1 m² quadrats were compared among sites at four different altitudes (medium and high montane, low and high subalpine) and three types of infrastructure (primary road, secondary road and building, total 12 sites). Altitude, infrastructure and time of year did not affect percentage cover of vegetation. Flowering started earlier and lasted longer in the low montane sites compared to high subalpine sites. The type of infrastructure only affected the number of reproductive structures at the peak of flowering, with A. millefolium growing next to buildings having two to three times more inflorescences per m² than along primary and secondary road verges. At the peak for each reproductive stage, there was an average of 1.47 developing inflorescences, 21 inflorescences in bud, 24 inflorescences in flower, 4 inflorescences setting seed, and 3 releasing seed. Based on the maximum number of inflorescences present at any time at each site, there was an average of 36 inflorescences, giving an estimate potential seed production of 51 400 seed per one m² for A. millefolium in the Snowy Mountains. If the climate changes in the Snowy Mountains as predicted, then it is likely that yarrow will produce more inflorescences and seed in the higher altitude sites.     

Seedling occurrence in alpine treeline conifers: A case study from the central Rocky Mountains,USA

Information concerning the occurrence of very young (1- to 10-year-old) tree seedlings in the alpine treeline ecotone is rare. Seedling occurrence of the dominant conifers Picea engelmannii and Abies lasiocarpa was measured in the treeline ecotone of the Medicine Bow Mountains, Wyoming (central Rocky Mountains, USA), an area composed of elongated tree islands separated by open meadows (ribbon forest) that grade into the closed forest. No seedlings were found on the windward sides of tree islands, while a mean of 0.6 seedlings.m^–2 occurred on the leeward (downward) sides. These values compared to the 4.2 seedlings.m^–2 in the closed forest. In addition, a strong correspondence was found between snowpack depth and seedling abundance, with depths that were either too shallow (< 0.5 m) or too deep (> 1.5 m) associated with fewer or no seedlings. A. lasiocarpa seedlings made up much less of the overall seedling population in the ribbon forest (6 %) than in the closed forest (22 %). Seedling establishment in this portion of the alpine treeline ecotone appears to be occurring at a low rate that differs between the two dominant species and may be strongly influenced by wind-driven snow accumulation patterns.     

Hydatid control in Australia: where it began, what we have achieved and where to from here

Echinococcus granulosus was imported into Australia with domestic livestock about 200 years ago. It spread rapidly through domestic animals and quickly became a public health problem in the new colony. Control was hampered by ignorance of the transmission pattern. The association between metacestodes and tapeworms was not elucidated until 63 years after the arrival of the First Fleet. Australian wildlife were highly susceptible to infection with E. granulosus and wildlife/domestic animal interaction facilated rapid infiltration of wildlife by E. granulosus. The wildlife reservoir has hampered hydatid control campaigns on mainland Australia but successful eradication has been achieved in the island state of Tasmania where there was no wildlife reservoir. The application of a new recombinant vaccine for sheep in control campaigns and the use of praziquantel baits for controlling infection in dingoes around bush campsites and picnic areas is discussed.     

Selective predation on the broad‐toothed rat,Mastacomys fuscus (Rodentia: Muridae), by the introduced red fox,Vulpes vulpes (Carnivora: Canidae), in the Snowy Mountains,Australia

Abstract Since 1981 there has been debate over whether foxes (Vulpes vulpes Linnaeus) selectively prey on the broad‐toothed rat (Mastacomys fuscus Thomas) relative to the bush rat (Rattus fuscipes (Waterhouse)). In the present study, three areas of the argument are examined. (i) In a study of fox diet over 3 years at both alpine and subalpine altitudes, M. fuscus outnumbered R. fuscipes in faecal remains in all seasons, in all years, and at both altitudes. Overall, M. fuscus occurred in scats five times as frequently as did R. fuscipes in the alpine zone and twice as often in the subalpine zone. (ii) Data from population studies of M. fuscus and R. fuscipes showed no evidence that M. fuscus is trap shy; neither the pattern of captures of individuals caught once, twice and so on, nor the proportion of the estimated population of each species captured during trapping sessions was significantly different. (iii) The suitable habitat for M. fuscus within the potential home ranges of foxes contributing to the subalpine fox scat collection constituted approximately 50% of the total area. However, there was no significant difference between the numbers of fox trails encountered in habitat suitable or unsuitable for M. fuscus in 19 paired transects skied in winter, indicating no preferential foraging in either habitat. Selective feeding on M. fuscus was therefore established, but how that choice is exercised was not determined.     

天山高寒草原土壤团聚体结构与养分含量对模拟氮磷沉降的响应

为更好地理解高寒草原土壤团聚体结构及其养分含量对氮(N)、磷(P)沉降增加的响应,于2018年开始依托全球营养网络(Nutrient Network)在巴音布鲁克草原生态系统研究站开展模拟氮磷沉降的短期(<5年)氮磷添加控制实验,设置对照(CK)、N添加、P添加、NP交互添加4个处理。N、P添加量均为10 g m^-2 a^-1。于2021年8月采集植物与土壤样品,采用湿筛法分析土壤水稳定性团聚体组成,测定全土和各粒级团聚体有机碳(SOC)、总氮(TN)、总磷(TP)、速效氮(AN)和速效磷(AP)含量。研究结果表明：(1)巴音布鲁克高寒草原各粒级土壤团聚体比例从低到高依次为：0.053—0.25 mm、<0.053 mm、0.25—2 mm、和>2 mm,以>2 mm团聚体占主导,其比例在45.48%—71.81%之间。(2)N添加显著降低了0—10 cm土壤层>2 mm团聚体比例和团聚体稳定性,而P添加则显著降低了10—20 cm土壤层>2 mm团聚体比例和团聚体稳定性。(3)0—10 cm土壤层各粒级团聚...     

When to come out from a refuge: risk-sensitive and state-dependent decisions in an alpine lizard

Prey often respond to predator presence by increasing theiruse of refuges. However, unfavorable thermal conditions in refugesmight entail physiological costs for an ectothermic prey. Thus,the decision of when to come out from a refuge should be optimizedby considering the expected fitness effects of diminution ofpredation risk with time, but also by considering the cost of theloss of time spent at optimal body temperature maximizing physiological functions.The model of Ydenberg and Dill describes the trade-off betweenrisk and cost for a prey fleeing to a refuge. We present a specialcase of this model to predict how emergence time from the refugein lizards or other ectotherms should vary as a function ofrisk of predation and thermal costs of refuge use. The analysesof the variation in emergence time from a refuge of Lacertamonticola lizards in the field under two different predation risklevels supported the predictions of the model. As predicted,time spent in the refuge was longer when the threat of the initialattack had been higher, and therefore the subsequent diminutionof risk was slower, but only when lizards emerged at the sameplace where they hid. When initial body temperature was high,some lizards decreased emergence time by emerging from a differentplace. In addition, the effects of thermal costs were more relevant inthe high-risk situation. Time spent in the refuge under highrisk increased when thermal conditions of the refuge were moresimilar to thermal conditions outside (i.e., physiological costsof refuge use were lower). We conclude that optimization ofrefuge-use strategies might help lizards cope with changes in predationrisk without incurring excessive physiological costs.     

Allopatric divergence and secondary contact without genetic admixture for Arichanna perimelaina (Lepidoptera: Geometridae), an alpine moth endemic to the Hengduan Mountains

Mountain systems, especially at high altitudes, are an excellent model for determining the mechanisms underlying high species diversity and endemism. Herein, we elucidate the evolutionary history of the alpine moth Arichanna perimelaina (Wehrli), which is endemic to the Hengduan Mountains (HM) region in southwest China, based on three mitochondrial genes and two nuclear genes. Our results revealed six deeply divergent clades that corresponded to populations in different mountain systems in the HM region. Bayesian divergence time estimations suggested a mid‐ to late Pleistocene genetic divergence. The results also showed that the Mt Yulong (YL) region was a refugium and valley corridors established by glaciation during the Pleistocene allowed populations on the separate mountains to migrate. The reproductive isolation among the different clades on contact zone in the YL region may be associated with the asynchronous mating rhythms and/or the divergent mate recognition caused by the ecological source of divergent selection. Allopatric divergence associated with complex topographies and climatic oscillations, regional dispersal via valley corridors and the suitable refugium of the YL region shaped the genetic divergence and distribution pattern of A. perimelaina in the HM region. These findings highlight the essential role of complex terrain and climatic fluctuations in shaping the unique phylogeographic history of a narrow alpine moth, and provide insights into the mechanisms underlying high species richness and endemism in the HM region.     

Three species of fishes from an eutrophic, seasonally alkaline lake are not more tolerant to acute exposure to high pH in the laboratory

A number of different freshwater fish species (perch Perca fluviatilis , roach Rutilus rutilus and rudd Scardinius erythrophthalamus ) from either eutrophic (Slapton Ley, a seasonally alkaline lake) or non-eutrophic waters were compared with respect to their sodium uptake kinetics and tolerance to acute (1 h) exposure to pH 9·5. Further comparisons were made with rainbow trout Oncorhynchus mykiss and brown trout Salmo trutta . The influence of fish size was also investigated in rainbow trout. Exposure to pH 9·5 was found to disrupt sodium balance and inhibit ammonia excretion in all species and sizes of fishes. The origin of fishes did not have a significant effect on the sodium uptake kinetics or the physiological responses to high pH water. The fishes from the eutrophic lake therefore did not appear to have any increased tolerance to acute exposure to alkaline water. In contrast to previous studies there was no inhibition of Na⁺ uptake during exposure to high pH. Indeed in some groups of fish Na⁺ uptake was actually stimulated, as was Na⁺ efflux. These differences are attributed to experimental water composition and interspecific differences in physiology. It was not always possible to size-match fishes of the different species, so rainbow trout were used to assess the effect of body mass (from 2 to 40 g), on Na⁺ uptake kinetics and Na⁺ or ammonia fluxes during alkaline water exposure in rainbow trout. Size had no significant effect on these measurements within this narrow range, which helps validate the comparison between species in this study.     

氮磷添加对昆仑山北坡高山草地N2O排放的影响

氮(N)、磷(P)等养分添加是提高草地生态系统生产力的重要策略, 但其对土壤氧化亚氮(N₂O)排放的影响尚不明确。该研究以南疆昆仑山北坡高山草地为研究对象, 设置氮添加、磷添加、氮磷交互以及不施肥(CK) 4个处理, 采用静态箱-气象色谱法连续监测2017年生长季草地的N₂O排放, 研究不同氮、磷添加处理下的N₂O排放特征, 并利用Pearson相关分析对影响N₂O排放的主要环境因子进行定性识别及定量解析。结果表明: 氮添加处理与氮磷交互处理在施肥后约3周引起显著的N₂O排放峰, 分别为42.3和15.4 g N·hm ^-2·d ^-1。与其他处理相比, 氮添加处理生长季N₂O排放通量显著增加了1.8-3.2倍, 而磷添加以及氮磷交互处理与CK之间没有显著差异。Pearson相关分析结果表明: N₂O排放与微生物生物量碳呈负相关关系, 与溶解性有机碳含量、pH值呈正相关关系, 而与其他环境因子关系不显著。以上结果表明, 与单施氮肥相比, 在该地区草场采用氮磷混施可显著减少N₂O的排放。     

Stromatolite branching in the Neoproterozoic of the Centralian Superbasin, Australia: an investigation into sedimentary and microbial control of stromatolite morphology

The extensive and well-preserved Neoproterozoic Acaciella australica Stromatolite assemblage of Australia is ideal for examining the relative roles of microbial and environmental influences on stromatolite branching and stromatolite macrostructure across a wide geographical area. Detailed sedimentological analyses indicate that the basal hemispheroidal section of bioherms contains abundant sediment. By contrast, the columnar sections of bioherms are composed almost exclusively of micritic laminae. These micritic laminae display little evidence for environmental, especially sedimentary, control over stromatolite morphology. The change from a hemispheroidal morphology to branching morphology is linked to variations in the relative contributions of sediment and framework growth. The shift to columns appears to be closely linked to a decrease in sediment supply that resulted in a more stable environment in which microbially mediated framework growth began to control stromatolite morphology. Branching in the A. australica assemblage stromatolites appears to be caused by shifting sedimentary and microbial control on stromatolite morphology.     

Susceptibility to low-temperature photoinhibition and the acquisition of freezing tolerance in winter and spring wheat: The role of growth temperature and irradiance

Five winter and five spring wheat ( Triticum aestivum L.) cultivars were grown under either control conditions (20°C/250 photosynthetic photon flux density (PPFD) [μmol m⁻² s⁻¹]), high irradiance (20°C/800 PPFD) or at low temperature (either 5°C/250 PPFD or 5°C/50 PPFD). To eliminate any potential bias, the wheat cultivars were arbitrarily chosen without any previous knowledge of their freezing tolerance or photosynthetic competence. We show that the differential susceptibilities to photoinhibition exhibited between spring and winter wheat cultivars, as assessed by chlorophyll fluorescence cannot be explained on the basis of either growth irradiance or low growth temperature per se. The role of excitation pressure is discussed. We assessed the correlation between susceptibility to low-temperature photoinhibition, maximum ribulose 1,5-bisphosphate carboxylase-oxygenase (EC 4.1.1.39) and NADP-dependent malate dehydrogenase (EC 1.1.1.82) activities, chlorophyll and protein concentrations and freezing tolerance determined by electrolyte leakage. Susceptibility to photoinhibition is the only parameter examined that is strongly and negatively correlated with freezing tolerance. We suggest that the assessment of susceptibility to photoinhibition may be a useful predictor of freezing tolerance and field survival of cereals.     

Craniosynostosis: an analysis of the timing, treatment, and complications in 164 consecutive patients

Treatment options for the craniosynostoses vary from conservative observation until completion of growth to radical remodeling in infancy. To further define the timing and type of treatment necessary in these complex disorders, we have retrospectively analyzed all patients operated on for this deformity during the past 12 years. One-hundred and sixty-four patients with craniosynostosis were analyzed and subgrouped into asymmetrical (predominantly unilateral) and symmetrical (bilateral) deformities, in addition to segregation by age and type of procedure performed. This was done recognizing that no deformity, like no normal human face, is truly symmetrical. Results of treatment were categorized on the basis of the need for additional surgery and varied from no refinements necessary (category I) to major reduplication of the initial procedure (category IV). Analysis of the data led us to conclude that excellent results can be expected in the asymmetrical deformities group treated in infancy by a unilateral approach. Similarly, for the mild symmetrical deformities, treatment at this time by bilateral orbital advancement gives satisfactory results in the majority of patients. By contrast, the more severe symmetrical groups treated in childhood have a high incidence of requiring secondary major reconstructions, and consideration should be given to delaying craniofacial surgery until age 7 or older, although earlier cranial surgery may be advisable.