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Flexibility in biological systems is seen as an important driver of macro-ecosystem function and stability. Spatially constrained endosymbiotic settings, however, are less studied, although environmental thresholds of symbiotic corals are linked to the function of their endosymbiotic dinoflagellate communities. Symbiotic flexibility is a hypothesized mechanism that corals may exploit to adapt to climate change. This study explores the flexibility of the coral–Symbiodinium symbiosis through quantification of Symbiodinium ITS2 sequence assemblages in a range of coral species and genera. Sequence assemblages are expressed as an index of flexibility incorporating phylogenetic divergence and relative abundance of Symbiodinium sequences recovered from the host. This comparative analysis reveals profound differences in the flexibility of corals for Symbiodinium, thereby classifying corals as generalists or specifists. Generalists such as Acropora and Pocillopora exhibit high intra- and inter-species flexibility in their Symbiodinium assemblages and are some of the most environmentally sensitive corals. Conversely, specifists such as massive Porites colonies exhibit low flexibility, harbour taxonomically narrow Symbiodinium assemblages, and are environmentally resistant corals. Collectively, these findings challenge the paradigm that symbiotic flexibility enhances holobiont resilience. This underscores the need for a deeper examination of the extent and duration of the functional benefits associated with endosymbiotic diversity and flexibility under environmental stress.
1. Introduction
Variety in endosymbiotic communities provides a setting for interactions between endosymbionts within and among cells that have the potential to influence processes among host cells in both positive and negative directions. The paradigm that flexibility in the form of functional diversity equates to resilience, stability, greater functional range and adaptive potential is heavily grounded in ecological studies [1–3], but has yet to be investigated in detail at both the functional and the genetic level in endosymbioses. Endosymbiotic communities represent extreme examples of such scenarios, with hosts (usually macro-eukaryotes) housing communities of micro-organisms (micro-eukaryotes or bacteria) within the boundaries of their own cell membranes and tissues. The implications of flexibility in spatially constrained, densely populated environments may be particularly profound in driving the structure and function of the community [3]. Cnidarian–dinoflagellate symbioses are provocative subjects for such an investigation because they are taxonomically complex associations, whose stability and functional integrity underpins the persistence of corals and coral reef ecosystems through time.
Coral reefs are often described as sentinel ecosystems for their sensitivity to climate change stressors such as increasing seawater temperatures and ocean acidification as well as local impacts of anthropogenic activities [4,5]. The sensitivity of reef-building corals to these stressors is driven, to a large degree, by physiological constraints imposed by their unions with endosymbiotic dinoflagellates in the genus Symbiodinium [6–8]. The genus Symbiodinium is classified into nine major taxonomic lineages (A–I; [9]) that each contain multiple types. Symbiodinium clades and within-clade types exhibit patterns of association with specific coral taxa, and patterns in the distribution of Symbiodinium types across space and environment indicate differences in their physiological thresholds within the genus [10,11]. Indeed, shifts in the taxonomic composition of the communities of endosymbiotic dinoflagellates harboured by reef-building corals have been implicated as one mechanism that might confer resistance through tolerance, or resilience through adaptation, to changes in the environment [12].
The identification of functional differences in Symbiodinium physiology prompted the introduction of the adaptive bleaching hypothesis (ABH, [12]), which states that 'bleaching provides an opportunity for the host to be repopulated with a different type of partner; frequent stress tends to favour a stress-resistant combination'. It has been further posited that this could be achieved by switching the existing symbiotic community for a new and better-adapted type, or by shuffling the relative proportions of the existing types within the community, increasing the abundance of previously cryptic types better adapted to the stress event [13]. Recent research has emphasized the importance of Symbiodinium shuffling in withstanding changes in global climate, such as temperature-induced bleaching [14,15]. While it has been suggested that the flexibility in Symbiodinium–coral interactions is largely beneficial [13] through improved holobiont thermal resistance [14], there are indications that this may not be the case for all species or temporal scales [7,8,16].
Much of the work characterizing Symbiodinium has focused on dominant Symbiodinium types in hosts, or those that are readily cultured from a host. As a result, the general perception has been that only a few corals are capable of hosting multiple Symbiodinium types, and that corals generally exhibit low endosymbiotic flexibility [17]. However, in recent years and with the application of more sensitive and more inclusive molecular approaches, our understanding of the complexity of cnidarian–Symbiodinium symbiosis has grown significantly [9], and many corals have been identified as hosting multiple types of Symbiodinium [18]. Furthermore, in some coral species, the Symbiodinium communities are now known to vary widely over space and time [19,20]. To date, however, symbiotic flexibility has not been quantitatively compared in a broad range of coral taxa, and the ecological performance of corals has never been linked to the levels of endosymbiotic flexibility of the host.
The goals of this study were to characterize, quantify and compare the flexibility of a range of coral taxa for Symbiodinium and map this trait onto the ecological resilience and biological attributes of the coral host documented in the literature. Flexibility here is defined as the ability to have a varied Symbiodinium sequence assemblage, or sequence community. Importantly, our study uses a comparative approach among samples and taxa, by assessing differences in Symbiodinium sequence assemblages among coral species. These data provide the capacity to link patterns in the Symbiodinium sequence assemblages that represent the endosymbiotic flexibility in corals, with the documented physiological susceptibility of coral holobionts.
2. Material and methods
(a) Sample processing
Corals samples were collected from fringing reef, lagoon reef or forereef habitats around the island of Moorea (figure 1; 1–24 m; electronic supplementary material, tables S1 and S2). DNA was extracted from coral fragments using the Qiagen DNA Easy kit (Qiagen, CA). Following extraction, polymerase chain reaction (PCR) amplification of the nuclear ribosomal partial 5.8S, internal transcribed spacer 2 (ITS2) and partial 28S rDNA regions was carried out using the ITS-DINO forward primer [21] and ITS2rev2 reverse primer [22]. To minimize cross hybridization of the primers with the coral host, touchdown PCR thermocycling conditions (25 µl) were used as follows: (i) 95°C for 10 min; (ii) 25 cycles 94°C for 30 s, 65°C for 30 s (decreasing the annealing temperature 0.5°C for every cycle after cycle 1) and 72°C for 1 min; (iii) 14 cycles of 94°C for 30 s, 52°C for 30 s and 72°C for 1 min; and (iv) final extension of 72°C for 10 min. PCR products were purified using the QIAquick PCR Purification Kit, ligated into pGEM-T Easy vector and transformed using α-select gold efficiency competent cells. Clone libraries were grown overnight on selective LB agar plates containing ampicillin, IPTG (isopropyl-β-d-thiogalactopyranoside, Fermentas, MD) and XGAL. Positive inserts were identified via PCR screening with M13 primers, purified with exonuclease I in shrimp alkaline phosphatase and sequenced on an automated sequencer (BigDye Terminator chemistry) with a target of 10 sequences per sample.
(b) DNA sequence alignment and analysis
Alignment and inspection of raw sequences were completed using Sequencher v. 4.7 and Bioedit v. 5.0.9 [23]. Sequences with single polymorphisms were included only in the downstream analysis if recovered from three or more independent clone libraries (host samples). Symbiodinium sequences with single polymorphisms found in less than three clone libraries were likely due to PCR error, and were converted to the closest sequence in the dataset that occurred greater than three times [11]. Sequences with novel identities as assessed by BLAST results from GenBank were given a new alpha-numeric identifier (sensu [11]), based on the distance from the closest previously documented haplotype (e.g. C15 variant = C15.x). All sequences are available in GenBank under the accession numbers HE578975–HE579042 (table 1).
(c) Statistical analysis
To determine coverage of the clone libraries, and to test for equal coverage across host taxa, coverage estimates were determined following Stat et al. [34], and coverage values were tested with one-way ANOVA among host genera. To compare Symbiodinium sequence assemblages among coral taxa, relative abundance data were square root-transformed, and a Bray–Curtis similarity coefficient [35] was calculated for all samples, ranging from 0 (indicating dissimilarity) to 100 (indicating identical assemblages). Analysis of similarity (ANOSIM) of the Bray–Curtis coefficients was carried out using primer v. 6.0 [36,37], and the two-dimensional ordination of the samples was represented with non-metric multi-dimensional scaling (MDS), or a spatial representation of the relative similarities between Symbiodinium sequence assemblages of coral samples. ANOSIM was used to test the hypotheses that there were no differences in the Symbiodinium sequence assemblages among species or genera.
To describe the Symbiodinium sequence assemblage for each sample, we included both genetic distance and frequency of symbiont occurrence in a single metric, or an index of flexibility (Fl). This index was calculated based on a modification of parasites' specificity for their hosts from parasitological research [38,39]. The index values range from 0 to ∞, where larger values indicate greater flexibility in the coral and the presence of a variety of Symbiodinium sequences that are genetically dissimilar, and lower values indicate lower flexibility in the coral and the presence of one or a few Symbiodinium sequences that are genetically similar. The index is described by the following equation:
In this equation, ωij is equal to the uncorrected pairwise genetic distance between Symbiodinium sequences; pi is equal to the relative abundance of the first Symbiodinium haplotype; pj is equal to the relative abundance of the second Symbiodinium haplotype. The product of ω(pipj) are summed across haplotypes i = 1 … n, and j = 1 … n (i.e. across all possible pairs of Symbiodinium haplotypes within a sample), normalized to the sum of the product of the relative abundance of each pair of haplotypes (pipj). For within-clade comparisons, uncorrected pairwise genetic distances of the ITS2 region were calculated using mega v. 4.0 [40]. Within-clade alignments were based on all non-redundant ITS2 A1, C1 and D1 sequence hits using the Symbiodinium database, SYM-BLAST (http://126.96.36.199/srsantos/symbiodinium/blast/blast_cs.html), in addition to the existing alignments from the current study. Alignments were created using ClustalW [41] as well as further manual alignment, and resulted in alignment of 101, 431 and 53 sequences for clades A, C and D, respectively. Therefore, genetic distances used for the index of flexibility calculations span the range of genetic distance documented within each Symbiodinium clade, and scale the genetic distance of our sequences to appropriate within-clade divergence. Similarly, average uncorrected pairwise genetic distance was calculated for the nine clades of Symbiodinium (A–I, [9]) based on the sequences of a portion of the relatively conserved nuclear ribosomal array (nr28S sequences, D1–D3 region, using the program mega v. 4.0 [40]. As the variation in ITS2 sequences between clades is too great for accurate alignment of sequences of all clades, and owing to the differences between markers (ITS2 versus nr28S), the specificity values are calculated separately within a clade (W) and among clades (A), and summed. Flexibility of the host for its Symbiodinium sequences was calculated for each coral sample. To statistically group coral genera in terms of their flexibility, the Gower % similarity distance measure [42] was used on the column-standardized values of flexibility. Flexibility groupings were calculated by UPGMA clustering [43] of the % similarity matrix among taxa in primer v. 6.0 [36,37].
3. Results
In total, 1240 Symbiodinium ITS2 sequences were recovered from 132 samples representing 34 species, 14 genera and seven families of coral (see the electronic supplementary material, table S1). All 1240 ITS2 sequences were used to report Symbiodinium clade by host (figure 2). Owing to the very low number of sequences recovered from samples of Montastrea curta, this species was excluded from downstream analyses. The average number of sequences recovered from the remaining 128 host samples was 9.6, and a total of 1223 sequences were used in similarity and flexibility calculations and statistical analyses. Coverage of the clone libraries at the level of genus ranged from approximately 54 to 75 per cent and did not differ among genera (F12,115 = 1.01, p > 0.05).
BLAST identity grouped the sequences into 68 different groups, representing 4, 7 and 57 Symbiodinium sequences from clades A, D and C, respectively (table 1 and electronic supplementary material, table S1). These included several commonly reported types (A1, A3, D1, D1a, C15, C45, C1, C1b, C3, C21), in addition to a number of novel ITS2 sequences (table 1). Ninety per cent of the corals sampled hosted one clade of Symbiodinium (n = 101 for C, 11 for D and 7 for A), 9 per cent hosted two (n = 9 for CD, 2 for AC and 1 for AD) and 1 per cent hosted three (n = 1 for ACD; see electronic supplementary material, table S1 for details). The similarities of Symbiodinium sequence assemblages among coral samples were assessed via calculation of Bray–Curtis similarity coefficients. MDS of the similarity coefficients showed separation in the Symbiodinium sequence assemblages among host taxa (figure 3 and electronic supplementary material, figure S1). ANOSIM of the Bray–Curtis coefficients provided support for the MDS ordination with significant differences in the Symbiodinium sequence assemblages among coral hosts at the level of species (R = 0.62, p = 0.001), and genus (R = 0.44, p = 0.001; electronic supplementary material, table S3). The average similarity of Symbiodinium assemblages within a genus ranged from approximately 11 per cent in Leptoseris to 50 per cent in Gardineroseris, and the average dissimilarity between genera ranged from approximately 55 to 100 per cent (see the electronic supplementary material, table S4).
The diversity and genetic divergence of ITS2 sequence assemblages was collapsed into a single metric (index of flexibility) for the 128 coral samples. The distribution of values of flexibility, ranged from 0 to 0.2197, reflecting corals that were only host to a single type (i.e. 0), and those hosting multiple distantly related types of Symbiodinium from the most distantly related clades (i.e. 0.2197), respectively. Overall, seven coral samples hosted one sequence, and 121 corals hosted multiple sequences (see the electronic supplementary material, table S1). Acropora, Leptastrea, Leptoseris, Montipora, Pavona and Pocillopora had the highest values and variance, exhibiting the greatest flexibility with respect to Symbiodinium (figure 4). The high flexibility values for Acropora were driven by six corals each with large values, whereas the high flexibility in Montipora was driven by intersample variance in the presence of a single, but different, clade (A or C; figure 4 and electronic supplementary material, table S1). Cyphastrea, Fungia, Gardineroseris, Herpolitha, Porites and Psammocora had consistently low intersample variance and low flexibility values (figure 4). The patterns of low and high flexibility were supported with UPGMA clustering of percentage similarity in flexibility among genera. The corals clustered into two groups that were less than 26 per cent similar between groups and more than 78 per cent similar within a group. The high flexibility group (i.e. generalists) consisted of Acropora, Leptastrea, Leptoseris, Montipora, Pavona and Pocillopora, and the low flexibility group (i.e. specifists) consisted of Cyphastrea, Fungia, Gardineroseris, Herpolitha, Porites, Pachyseris and Psammocora (see the electronic supplementary material, figure S2).
4. Discussion
Our results demonstrate a large difference in composition of endosymbiotic communities hosted by scleractinian corals from Moorea. Quantitative examination of the flexibility of scleractinian hosts for Symbiodinium sequence assemblages identified two groups of coral genera that can be classified as endosymbiotic 'generalists' (high flexibility) and 'specifists' (low flexibility; figure 4 and electronic supplementary material, figure S2). Notably, the differences in the patterns of ITS2 sequence data, we report here are consistent with results previously published in Stat et al. [11], who examined Symbiodinium sequence assemblages in a range of coral hosts from Johnston Atoll using the ITS2 and the low copy chloroplast 23S in parallel. Here, we use ITS2 sequence assemblages as tools to describe and compare the Symbiodinium communities in corals. Reflecting the known interpretational complexity of this multi-copy nature and intragenomically variable marker [34], we make no inference regarding the relationship between the number of sequence types present in the ITS2 assemblages and the number of independent biological entities in the Symbiodinium communities in these corals. Further, the differences detected using the sequence assemblage approach are also evident when only previously described dominant ITS2 types are examined, or identified to the clade level in our study, as well as within the broader literature ([44]; electronic supplementary material, figure S3). Specifically in our study, Acropora and Pocillopora are symbiotically flexible and host five previously described dominant Symbiodinium types from three different clades, while massive Porites colonies hosts a single type (C15) from a single clade.
To explore the potential functional implications of these very different symbiotic strategies (flexibility versus specificity), and to exploit the comparative opportunity afforded by these groupings, we examined the literature and evaluated other biological traits that associate with generalist and specifist coral hosts, with the goal of assessing the potential physiological and ecological implications of symbiotic generalists versus symbiotic specifists (table 2 and references therein). Generalist corals that are flexible with respect to Symbiodinium exhibit very different biological attributes when compared with corals that are specifists, or inflexible (table 2). Most surprising among these is the stark difference in environmental sensitivity. Many of the corals that exhibit high symbiotic flexibility (generalists), such as the acroporids and pocilloporids are stress sensitive and often described as ecological 'losers' [61]. In contrast, coral species with high symbiotic specificity (i.e. high fidelity with respect to Symbiodinium) such as massive environmentally resilient poritids are ecological 'winners' [62,63,69,70].
The identification of the ecological pattern of endosymbiotic flexibility and stress susceptibility identified in tropical corals in our study conflicts with the broadly accepted idea that endosymbiotic flexibility enhances resilience in tropical reef coral–Symbiodinium symbiosis [12]. For example, ecological surveys of the bleaching severity of the corals of Moorea following the 1994 mass-bleaching event revealed that the per cent bleaching was highest in Acropora (approx. 90–100%) followed by Pocillopora (approx. 70–90%), Montipora (approx. 20–50%) and Porites (approx. 13–43%; [69]). Furthermore, the 'winning' and 'losing' taxa are relatively similar biogeographically [61,62] with the generalists Acropora and Pocillopora showing a higher propensity to bleach, and higher mortality than the specifists such as Porites [63]. This pattern also holds for the stress of ocean acidification [64], where there is a decrease in the diversity and complexity of the reef through the loss of Acropora and Pocillopora at low pH, and an increase in the relative abundance and dominance of massive Porites. Indeed, the higher symbiotic flexibility of the generalist species does not appear to be reflected in a holobiont function that is ecologically beneficial under challenging environmental conditions. Support for the benefits of fidelity in symbiotic assemblages is also paralleled in temperate symbioses, where low diversity of vertically transmitted Symbiodinium appears to promote stability in cnidarian–dinoflagellate symbiosis [71]. For example, anemones exposed to varying irradiances host genetically similar Symbiodinium and are ecologically stable, a trait likely maintained via maternal inheritance of genetically similar symbionts [72].
A number of other interesting features emerge from our data. For example, the composition of the endosymbiotic communities in the generalists Acropora and Pocillopora varies widely among individuals within and among species in these genera at the single sampling interval represented in our study (figure 4). Furthermore, this pattern is manifest in the broader literature where spatial and temporal variability in symbiotic flexibility in generalists is commonly reported. For example, the Symbiodinium type hosted by the pocilloporid Seriatopora hystrix varies among colonies within a single reef (2–27 m at Yonge Reef) and between two reef sites on the GBR (Yonge and Day Reefs; [20]). Likewise, temporal flexibility is common in Acropora, where an ontogenetic shift occurs from clade A at 10 days old, to clades A, C and D in 83-day-old juveniles, a distribution also found in adult Acropora longicyathus [19]. Generalists corals such as acroporids and pocilloporids are some of the most ubiquitous and speciose reef-building corals in the Pacific [73]. They are characterized by branching morphologies that are plastic with respect to the environment. These genera exhibit high recruitment and rapid growth following disturbances [74], attributes that frame their description as weedy species. Generalist corals also show intraspecific and intergeneric variability in their feeding strategies (autotrophy and heterotrophy), and they acquire Symbiodinium using both vertical and horizontal transmission strategies [56]. The highly flexible acroporids and pocilloporids are generally dominated by clade C Symbiodinium but also host clades A and D. The within-clade ITS2 types found in generalist corals are often found in symbiosis with a broad range of hosts and are, for the most part, basal members of their respective clades (A1, C1, C3 and D1). These are the Symbiodinium types often described as generalist [13,27] and opportunistic [16,75]. The broad biological flexibility and capacity of generalist corals to interact with a wide range of Symbiodinium types is clearly advantageous under stable conditions. However, under prolonged environmental stress, the promiscuity and symbiotic entrepreneurialism of flexible hosts may drive competitive interactions within the symbiosis that destabilize and impair the overall functional of the symbiotic interactions. Such a transition is likely to have dire fitness consequences for the coral host.
At the other end of the endosymbiotic flexibility spectrum are the specifist coral genera exemplified by Porites; corals that display high inter- and intra-specific fidelity in their symbiotic partnerships in our study (figure 4), as well as over space and time in other studies [32,76]. For example, massive Porites colonies associate with Symbiodinium C15 from depths of 1–17 m across the GBR, and in corals collected in Japan [77], Johnston Atoll [11] and American Samoa [32]. Massive Porites is exemplified by a greater stability and persistence over time and under stressful conditions [32,57,61,76], traits attributed to slow growth [57], thick tissues/high tissue biomass [61,63] and thermally tolerant Symbiodinium C15 [78,79]. Characteristics of specifist corals such as Porites include symbiosis with endosymbionts such as Symbiodinium C15, which is commonly transmitted vertically across generations in the host [56], a mechanism that promotes coevolution and integration with the host, and is one of the most highly derived members of the most derived clade C [9,80]. This high fidelity symbiosis results in a tight integration between a specifist coral and a symbiont, that leads to success under both stable and stressful environments, with fidelity promoting spatial and temporal stability of the holobiont and positive fitness benefits.
The profound differences in the symbiotic flexibility in corals on a reef, and the associations between symbiotic flexibility and coral environmental sensitivity raise fundamental questions that pertain to differences in interaction states in the coral–Symbiodinium endosymbiosis (i.e. mutualism versus parasitism). As reviewed by Thompson [81], high partner fidelity driven by few symbiotic options, and tight vertical transmission favours the evolution of reduced antagonism, or mutualism in symbioses, thereby increasing persistence of intimate and long-term mutualisms. This is clearly exemplified in the potentially obligate mutualism of massive Porites–Symbiodinium C15. Symbiodinium C15 is highly derived [80], unlikely to be free-living [28] and has not, to date, been successfully cultured [82], all evidence in support of an obligate mutualism. In stark contrast is the flexibility of generalist coral–Symbiodinium associations, and the opportunistic nature of these symbioses. Acropora, for example, is characterized by high symbiont flexibility [18,83] and horizontal symbiont transmission/acquisition [56,83], and therefore is exposed to a variety of free-living Symbiodinium options [28], many of which are currently in culture. Despite the short-term benefits to thermotolerance due to acquisition or shuffling [84–86], ecologically these holobionts are linked with reduced fitness and higher mortality under stressful conditions. Together, this suggests that the flexible symbioses lean towards relationships on the less beneficial end of the spectrum [16,75].
Here, we have highlighted two extremes, a specifist (massive Porites sp.) and a generalist (Acropora sp.). These represent some of the most important and abundant taxa on modern reefs and exhibit differences in their environmental sensitivity, making them major drivers of reef community composition. The ability to detect the range and flexibility of Symbiodinium sequences present within a host sample and quantify the flexibility of the host through studies such as ours, allows for the discovery of such links between endosymbiotic flexibility and environmental sensitivity. This results in the ability to generate new testable hypotheses for predictions of the stability of a range of Symbiodinium–holobiont combinations under various environmental stressors, particularly in a changing global climate.
The examination of coral–Symbiodinium interactions has expanded our understanding of diversity in Symbiodinium from one [87] to many [9,88,89], and from functional equality [87] to inequality [6,16,90]. Here, we link symbiotic stability to holobiont resistance, and symbiotic flexibility to holobiont sensitivity (table 2). The identification of symbiotic flexibility has promoted the hypothesis that there is potential for 'adaptation' of corals to environmental stress via the acquisition of new or shuffling of existing populations of Symbiodinium (sensu the ABH [12,13]). The rate at which natural selection can occur to produce a more tolerant holobiont, however, may be outpaced by the rapid rate of change in environmental conditions related to anthropogenic global warming and ocean acidification, and the increasing frequency of these stress events. Additionally, the outcome of this flexibility in acquisition, or ability to shuffle, may not always be beneficial to the holobiont (e.g. declines in growth and energy acquisition; [7,16,91]). Notably, the long-term benefits of this flexibility in symbiosis are unknown, as research to date has focused on short-term tolerance (less than or equal to one to two bleaching events), and implications across longer temporal scales have not been explored. It is likely that the reefs of tomorrow will be shaped by the resilient and resistant coral–Symbiodinium assemblages of today, which are dominated by those specifist coral genera associated with fidelity in their symbiotic unions (e.g. massive Porites sp.). A further examination of generalist and specifist Symbiodinium–coral unions is necessary to determine the range of benefits or costs to the holobiont associated with characteristics such as immune response, photosynthate production and release, symbiont population control and calcification. This information will improve our ability to determine the consequences of flexibility or specificity in the coral–Symbiodinium symbiosis in a future of environmental uncertainty and dire predictions for the maintenance of reef-building coral ecosystems.
R.D.G., M.S. and H.M.P. designed research; M.S. performed field research; H.M.P., X.P. and M.S. performed lab research; R.D.G. and H.M.P. contributed new reagents/analytic tools; H.M.P. analysed data; and H.M.P. and R.D.G. wrote the paper. We are thankful for assistance from the staff of the Richard B. Gump South Pacific Research Lab, as well as logistical support of Moorea Coral Reef Long-Term Ecological Research site (MCR-LTER) researchers and staff. We also thank Nicholas Fabina regarding discussions of quantifying flexibility, and two anonymous reviewers for their comments. This research was supported by funding from the National Science Foundation to the MCR-LTER (OCE 04-17412), the National Marine Sanctuary Program and Hawaii Institute of Marine Biology Reserve Partnership (memorandum of agreement 2005-008/66882), a postdoctoral fellowship to M.S. from the UWA-AIMS-CSIRO collaborative agreement, a grant to R.D.G. (OCE-0752604), as well as funds from the Gordon and Betty Moore Foundation. H.M.P. was supported by a fellowship from the US EPA (FP917199). The authors declare that they have no conflict of interest. This is a contribution of the MCR-LTER, and is contribution no. 1512 of the Hawaii Institute of Marine Biology and no. 8727 of SOEST.
- Received June 25, 2012.
- Accepted August 6, 2012.
- This Journal is © 2012 The Royal Society


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From The Files Of November 5, 1992
Fulton County voters offered no help to Bill Clinton's presidential victory, preferring incumbent President George Bush by a margin of about 51 percent to about 32 percent, with independent H. Ross Perot garnering about 17 percent of county votes. County voters did help incumbent Republican Senator Arlen Specter to re-election as he outpolled challenger Lynn Yeakle here by a margin of 2,786 to 1,888.
Nancy Keebaugh Traxler was named the Hustontown Area Fire Co.'s "Fireman of the Year" at the company's 25th annual banquet October 30 at the firehall in Hustontown. Traxler has been a member of the company for nine years, an emergency medical technician for seven years and a member of the auxiliary for 25 years. She is married to fellow firefighter Larry Traxler. The President's Award went to Morris Crouse who is vice president of the company.
Work on the new Southern Fulton Elementary School is slightly behind schedule, but the project should be completed around August 1, 1993. Ground was broken for the $7 million plus program November 18, 1991, after the school board had awarded contracts.
A screaming good time was had by all who visited the Jaycees Haunted House over Halloween which many considered the best yet with all its added spine-chilling innovations.
Commander William G. Hine of Harrisonville will be the featured speaker for the Fulton County Veteran's Day program November 11 at the courthouse.
Hustontown Area Fire Co. has dedicated a memorial and flag display in memory of their fellow firefighters who lost their lives while on work detail May 1, 1990. The company has also constructed a display case inside the firehall. The display shows photos and other memorabilia of the deceased firemen, James F. Chesnut Jr., Richard L. Hershey and Thomas L. Lane.
Ronald Younker and Tina Hendershott were united in marriage March 7 at Pleasant Grove Christian Church. Parents of the couple are Mr. and Mrs. Ray Younker Sr., Hancock, and Mr. and Mrs. John Hendershott of Big Cove Tannery.
Births at the Medical Center: a son to Mr. and Mrs. Gale Rouzer of Harrisonville on October 21; a son to Mr. and Mrs. Shawn Knepper of Hustontown on October 22; a son to Mr. and Mrs. Russell Seville of Mc- Connellsburg on October 23; a son to Mr. and Mrs. George Moats of Warfordsburg on October 29.
Deaths: Arlie Strait, 67, Needmore; A. Preston Gerhart, 100, Mercersburg; Seibert Garlick Jr., 60, Needmore; Thelma Bergstresser, 80, Roaring Spring; Thelma Gardner, 63, Winchester, Va.; M. Helen Kesselring, 70, Chambersburg; Frank Carnell, 88, Warfordsburg.
Marriage licenses: Walter Barmont III, McConnellsburg, and Beth Fleck, Three Springs; Gregory Mellott, McConnellsburg, and Wendy Richards, McConnellsburg.
Kelli Wooldridge and Ray Collins were united in marriage at Corpus Christi Catholic Church, Chambersburg, September 26. Parents are Mr. and Mrs. Robert Wooldridge of Warfordsburg and Mr. and Mrs. Dale Collins of Meadville and Mr. and Mrs. Harry Collins, Layton, Fla.
30 Years Ago
From The Files Of November 11, 1982
The Pennsylvania Turnpike toll plaza at Fort Littleton is moving.
A new booth and plaza are being built a couple hundred feet south of the current site, and the booth that now stands there is going even farther south, to Washington, D.C. The Fort Littleton plaza has been called "obsolete" and needs to be both replaced and expanded. To accommodate this expansion, the plaza has to be moved south along Route 522. The old toll booth is considered to be a relic, and the Smithsonian Institution in Washington, D.C., has claimed it. The booth will make its trip to Washington in the middle of next summer when the new plaza is expected to be ready for operation.
McConnellsburg businessman David "Andy" Washabaugh III recently opened a combination convenience store, gas station and carwash on Lincoln Way West. The new business, "Handy Andys," will be managed by Mark Clevenger.
Peggy D. Black, daughter of Mr. and Mrs. Warren Black of Waterfall, became the bride of David J. Baum, son of Mr. and Mrs. Robert Baum of York, on September 18, at Fairview United Methodist Church.
Specials at the IGA: tangelos, 19 for 98 cents; Del Grosso spaghetti sauce, 68 cents; Clorox, 59 cents; Kraft macaroni and cheese, two for 79 cents; grapes, 59 cents lb.; Heinz homestyle gravy, 69 cents.
Births at the Medical Center: a daughter to Mr. and Mrs. Paul Politis of Mc- Connellsburg on November 1; a son to Mr. and Mrs. James Canary of Big Cove Tannery on November 2.
Marriage licenses: Jeannette M. Cochran, Waterfall, and Eugene A. Dumer, Baltimore; Lisa Henry, McConnellsburg, and Mark Knepper, Hustontown; Vickie Seville, McConnellsburg, and Gerald Zeger, Harrisonville; Beatrice Tyler, Warfordsburg, and Ronald Keefer, McConnellsburg; Lori Neil, Willow Hill, and Thomas Elmer Jr., Spring Run.
Deaths: John L. Elbin, Warfordsburg; Wilhelmina Mellott, Hustontown; Sheldon Madden, Ormond Beach, Fla.
40 Years Ago
From The Files Of November 23, 1972
David Moses, administrator of Fulton County Medical Center; Fred
Mellott, board president; and Dr. G.T. Lorentz, chief of the medical staff, disclosed at a meeting of representatives of county businesses and civic organizations last Thursday night that there is a very real danger that the Medical Center may be forced to close unless more physicians are brought into this area. Mr. Moses said that the local hospital has been operating at only 45 percent of total capacity, and this situation cannot continue if the hospital is to remain financially healthy.
Pennsylvania's deer hunters, who have recorded remarkably large deer harvests for a number of years, should again have a really good season in 1972, according to the Game Commission. The regular two-week antlered deer season will open one-half hour before sunrise on November 27 and close at sunset on December 9. The regular statewide antlerless deer season is scheduled for Monday and Tuesday, December 11 and 12.
The home of "Buzz" Johnson, east of McConnellsburg, was considerably damaged by fire on Friday when a chimney from a wood and coal heating stove in the living room became overheated.
Coach Jack Frank has announced that Foster Earley has earned a slot on the 1972- 73 York College Spartan freshman basketball team. Foster, son of Mr. and Mrs. Eugene Earley of McConnellsburg, is a 6-foot forward on the squad.
Deaths: Earl B. Mellott, Warfordsburg; Mrs. Lulu Lodge, McConnellsburg; Irvin M. Hann, Everett; Roy E. Lowery, Altoona; Robert F. Rhodes, Mercersburg; Jesse R. Flood, Neelyton; Mrs. Larue Barthalow, Harrisonville; Richard R. Hull, Hancock; Tracey Lynn Brumbaugh, Saxton.
Birth at the Medical Center: a daughter to Mr. and Mrs. Monty R. Burrall of Mc- Connellsburg on November 12.
Specials at the IGA: turkeys, 10 thru 22 lbs., 39 cents lb.; pumpkin, 25 cents, 1 lb., 13- oz. can; sweet potatoes, 15 cents lb.; Duncan Hines cake mixes, 3 lb. boxes, $1; cranberries, 39 cents lb.
50 Years Ago
From The Files Of November 29, 1962
County, township and borough officials met with Fulton Industrial Development Association members on Monday night to discuss the economic future of Fulton County and, in particular, the best means of providing sewage disposal for the new Sagner building. Funds from the Area Redevelopment Program are available, but the plant, which is in Todd Township, would have to be annexed into the borough before application could be made. Another meeting is planned for December 10.
Sixty-six residents of Pennsylvania who reached their 100th birthday during 1962 have received testimonial plaques from the Pennsylvania Medical Society. Records show three out of four centenarians are females.
Engagement announced: Gayle Partchey, Harrisonville, to Raymond R. Mellott Jr., Needmore.
Marriage licenses issued: Kenneth N. Ray and Bonnie L. Cornelius, both of Mc- Connellsburg, and Clair J. Hollenshead and Joanne R. Koontz, both of Needmore.
Marriage announced: Esther L. Warthin, McConnellsburg, to Robert E. Lee, Hagerstown, on November 10 at Presbyterian Church in Hagerstown.
News of men in service: Pvt. Marlin T. Douglas, 23, of Needmore, has completed an aircraft maintenance course at the Aviation School at Fort Rucker, Ala. Army Pvt. Willis L. Kerlin, 22, of McConnellsburg, has completed advanced infantry training at Fort Gordon, Ga. A/C-1 William Warthin, McConnellsburg, left this week for an Air Force base in Germany, following a tour of duty in Newfoundland.
In response to a survey, over 500 people say they would be interested in work at the new Sagner factory.
Born to Needmore residents: a son, Dan Thomas, to Mr. and Mrs. Blaine Gordon; a son, Charles Webster, to Mr. and Mrs. Vernon Leese.
Postal rates as of January 7 will increase to 6 cents for first class, 8 cents air mail and 4 cents for postcards.
Kenneth Hall, New Enterprise, and Dorothy Brown, Fort Littleton, were married on Thanksgiving evening.
Beginning December 1, The Fulton County National Bank will pay 3 1/2 percent interest on time savings certificates.
Births at the Medical Center: a son to Mr. and Mrs. Delmar Eggleman, Burnt Cabins, on November 19; a son to Mr. and Mrs. Orville Mellott, Needmore, on November 19; a son to Mr. and Mrs. Carl Curfman, Three Springs, November 19; a daughter to Mr. and Mrs. John Spielman, Mercersburg, on November 22; a daughter to Mr. and Mrs. Ralph Hoover, Everett, on November 23; a son to Mr. and Mrs Donald Long, Hustontown, on November 23; a son to Mr. and Mrs. Lee Sheffield of Waterfall, on November 24.
Death: Charles W. Yaukey, 47, Chambersburg, brother of Humphrey Yaukey of Mc- Connellsburg.
60 Years Ago
From The Files Of November 27, 1952
Mr. and Mrs. Harry W. Abercrombie of Pittsburgh will spend Friday with their son-in-law and daughter, Mr. and Mrs. Daniel Grissinger of this place, before leaving on Saturday for a month's vacation in Florida.
William Brewer of upper Ayr Township is listed in critical condition in Chambersburg Hospital.
John and Jim Taylor and Donnie Black of Waterfall and George Taylor of New Grenada drove to Potter County last week for a day of bear hunting.
Rev. Ernest W. Brindle has been elected president of the McConnellsburg Ministerium.
Births: a daughter to Mr. and Mrs. Carl R. Pittman of McConnellsburg on November 19; a daughter to Mr. and Mrs. Paul N. Morton of Burnt Cabins on November 1; a son to Mr. and Mrs. H. Josef Stabler of Mc- Connellsburg on November 23; a daughter to Mr. and Mrs. Scott Gordon of Mercersburg on November 25.
Recognizing the sanctity of Christmas Day, the Game Commission decided on Saturday that all hunting in the state will be prohibited on December 25.
More births: twin daughters to Mr. and Mrs. Marvin Plessinger of Warfordsburg on November 13; a son to Mr. and Mrs. John Powell of Needmore on November 17.
A massive crowd jammed the local American Legion Home last Wednesday night to hear Shep Fields and his Rippling Rhythm Orchestra. A personal touch was added when the Wilson Wheelers of town renewed their acquaintance with the band's pianist, Andy Wright Jr. Both Wright and the Wheelers grew up in Beaver and were well acquainted before Wright struck out into the music world and Wheeler became a forester.
As of November 22, 318 persons in Fulton County were receiving public assistance.
The seniors of Southern Fulton will present "Hillbilly Wedding" on December 4-5. Taking part will be Benny Truax, Marietta Mellott, Shirley Hanks, Nancy Sheetz, Lois Barton, Gladys Gordon, Goldie Houdersheldt, Sarah Duvall, Jimmy Hollenshead, Merrill Gordon, Gary Dick, Mildred Hill, Clifford Wilt and Jack Ritz.
Death: Mrs. James K. Lynn of Harrisburg.
Mr. and Mrs. Avery Bohrer, former residents of Ridge, W.Va., have purchased the John Truxel farm at Warfordsburg.
Larry Shultz, son of Mr. and Mrs. George Shultz of Warfordsburg, is suffering from blood poisoning, the result of running a nail in his foot.
Mr. and Mrs. William Kirk Sr. of Warfordsburg are minus a barn roof as a result of the wind and rainstorm last Friday.
Deaths: Mrs. Fannie Elizabeth Shriever of Hancock; Mrs. Rebecca Margaret Baughman of Chambersburg.
70 Years Ago
From The Files Of November 26, 1942
Deaths: Mrs.
Priscilla Belle
(Keyser) Beatty of Cove Gap; Daniel Amos Fisher of Belfast Township; Andrew McGehee of Chambersburg.
Recent births: a daughter, Jeanine Marie, to Mr. and Mrs. H.E. Russell of Huntingdon on November 11; a son to Mr. and Mrs. Fred Martz of Hustontown, last Wednesday; a son to Dr. and Mrs. Lyle Sherwin of Mc- Connellsburg on November 8.
The following county men will enter the service on December 3: John V. Johnson, Norman F. Edwards, Forrest Garland, Harry E. Fix, Robert E. Doyle, Earl S. Alloway, Robert W. Colledge, Arthur L. Strait, Willard C. Mellott, John B. Stenger, Fred R. Sipes, Luther J. Strait, William C. Whetstone, Meade W. Stahle, Thomas W. Morton, Walter R. McElhaney, Clyde M. Craig, Maynard L. Mellott, Paul R. Michael, Emory N. Lanehart, Clyde R. Fletcher, William G. Buterbaugh, Wayne P. Strait, George C. Garland, Leo H. Gordon, Raymond E. Seville, Raymond Ward, Perry H. Seiders, Maurice E. Shaw, George W. Shaw, Robert W. Hann, James E. Keith and Albert Spriggs.
On Monday, Paul S. Shimer moved his barber shop from the Glen-Nel Hotel to the Morton building, just north of The Fulton County National Bank.
Mr. and Mrs. Daniel Grissinger of Hagerstown announce the birth of a daughter on Tuesday evening.
80 Years Ago
From The Files Of November 24, 1932
The Democratic victory parade in
McConnellsburg on Friday evening was a great success, although not as long as the Republican parade held before the election. "He laughs best who laughs last."
Death: Albert Allen Bollinger of Orbisonia.
Friends of Aunt Ray Sipes gathered at her home at Harrisonville last Wednesday and cut and stored five truckloads of wood for winter's use. The wood was donated by Homer L. Sipes, and the following helped cut and store it: Wilmer Sipes, Elmer Schooley, DeKalb Metzler, Arlie Deshong, Reuben Mellott, Rev. Chris Haumann, Ross Bard, Loy Hollenshead and Earl Williams.
Mr. and Mrs. McCauley Lynch of town were both injured in an automobile accident in Ohio last week. They were touring the state and ran into a fierce snowstorm. Several trucks were bogged down and one ran into their car as it was attempting to get free.
Bert Colledge of Todd Township was badly injured on Monday when hit by a car driven by Jack Cutchall. Mr. Colledge was admitted to Chambersburg Hospital with a broken leg, dislocated knee cap and a torn tendon.
Blaine Huston and Pauline O. Helman were wed October 31, 1942.
Clair H. Strait and Anita S. Knepper were married on October 20.
Charles Gelbert, star shortstop of the St. Louis Cardinals, received a leg injury while hunting with friends near Harrisonville last Wednesday. A gun was accidentally discharged, severing a nerve in his leg.
Mrs. Will Hamil of Knobsville received a badly sprained left ankle, bruises of the hands and an injured side when struck by a car driven by Raymond Wible, also of Knobsville, last Friday.
George Deshong of near Andover shot his little toe off last Tuesday morning. He had placed the muzzle of his gun on his boot when it accidentally discharged.
A thief entered the chicken coop of Elmer McCusker of Hancock last week and stole 20 chickens – the entire flock.
John Shives and Iva Flowers were married last Monday.
The stork visited Wells Valley last week and left new sons at the homes of Mr. and Mrs. James Sheeder and Mr. and Mrs. Herman Foster.
Work has begun on the new road from Waterfall to Hustontown, giving work to many of the unemployed in that section.
Thanksgiving specials at the A&P: lard, 2 lbs., 13 cents; butter, 2 lbs., 49 cents; sugar, 25 lbs., $1.15; fruit and nut cakes, 23 cents; coffee, 25 cents a pound.

subset: commoncrawl


text:
Odd choice of fonts. Only one way to improve on it. —JH
Odd choice of fonts. Only one way to improve on it. —JH
I pretty much agree with Anil Dash on the topic of wacky April Fools' jokes for websites, so instead I thought that today might be a good day to share a piece of genuine idiocy from the archives.
By the time Tobias and I began working together in 1999, we'd been friends for a decade, and had spent most of the previous years in close contact by phone. Our biographers will report this as a period of august correspondence in which we developed the philosophical framework that would inform our later collaboration, but the truth is that much of this time was spent goofing off, and naturally the arrival of the internet helped this project immensely.
Since we'd always been the types to tackle exhaustive projects, we both spent most of the nineties utterly exhausted. Many of our late night conversations were wits-end grievances about the impossibility of doing something or other, and these commonly degenerated into a discussion of Dumb Ideas for Typefaces. One of these, which I suggested in 1995, was that the OCR-A font — used on bank statements and designed for optical character recognition — really needed to be outfitted with a set of swashes. Using Adobe Illustrator, I ginned up the image above in about ten minutes, and sent it to Tobias. His response, which arrived within the hour, was a file named ¡¡¡Estupido-Espezial!!!.sit, which contained the following:


text:
Charleston County is pushing ahead with a plan to take more control and build the controversial extension of Interstate 526, even though the project's contract remains unsigned and such tasks usually are left to the state Department of Transportation.
County Council Chairman Teddie Pryor said the county is working along two tracks to move forward on the $558 million project, which has made little visible progress since council voted in favor of it in December.
County staffers are looking for a qualified company to manage the project, Pryor said. And they are working with the S.C. Transportation Infrastructure Bank and the DOT to complete the revised, three-way contract necessary to move the project forward.
Pryor said some people have questioned whether the county has the skills and resources to take on such a large construction project. But he's certain that it does. "We've never built an interstate," Pryor said, "but we do a good job with roads."
The completion of I-526 across Johns and James islands will be a parkway with speed limits between 35 and 45 mph, he said, and the road will not be designated an interstate because of those speed limits. "Now it's a regular road. It's a highway."
County Councilman Dickie Schweers, an opponent of the I-526 project, said he thinks the county taking it on is a bad idea. "I think that's the reason the SCDOT exists," he said, "to manage mega-projects."
He also said that he wasn't aware that the county had officially put out a call to companies interested in managing the project. The December vote to move forward with I-526 included a provision that allowed staffers to take actions on the project without first presenting them to County Council, he said.
Barrett Tolbert, the county's contracts and procurement director, said the county posted a Request for Qualifications for a management company on July 12. That's the first step in narrowing down the list of qualified, interested management companies, he said.
And he made it clear that the county is looking only for a project manager now, not a construction company to actually build the road. The county will invite some of the companies who respond to the request to submit formal proposals, he said. Interested companies must apply by Oct. 1.
The county is being careful to conduct the selection process appropriately, Tolbert said. "We're dotting all our i's and crossing all our t's."
Jake Libaire, a project manager for the Coastal Conservation League, a group opposed to the project, said he's confused by the county's change in stance on building the road. Last year the county tried to turn over the entire project to the DOT, he said. At the time, some council members said the county shouldn't be in the business of building interstates. The plan for the road was the same then as it is now, Libaire said. "So what changed between then and now?"
Pryor said he thinks negotiations on the three-way contract are in their final stages, and that the Infrastructure Bank and DOT ultimately will approve of the county's plan to take over construction of the road.
DOT spokesman Peter Poore said staffers from his agency are working on the contract, known as an Intergovernmental Agreement, or IGA, but he can't estimate how long it will take to complete. The DOT commission will hold a workshop Thursday morning, he said, and the group will discuss the matter there. Agendas for that workshop and other Thursday DOT meetings will be released this week, he said.
Pryor said the county already will have narrowed the list of possible qualified management companies by the time the IGA is complete. "It will get us ahead of the ball game," he said. "All we have to do is select a firm."
Reach Diane Knich at 937-5491 or on Twitter @dianeknich.


text:
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Revision of Kwaice's mage leveling guide from Tue, 2008-12-30 16:37
(This is the first version of the guide.
I'll soon add spell icons, wowhead links and finish all the sections.)
- Introduction
- Basic Spells
- Talents
- Cast rotations
- Frostfire Bolt
- Gear
- Useful addons
- Macros
- Glyphs (coming soon)
First of all, i'll ask you to be nice with my english, i come from Belgium and french is my primary language. This is also the first guide i write myself
This guide is here to help you leveling up your mage, not by telling you where to go and which quest to pick up (there are very nice leveling guides on wow-pro for that) but by giving you some piece of advice with the talents you should choose, the gear to aim for and some useful macros as well as addons.
It only aims to help you during the leveling process, it will not cover pvp, dungeons nor raids.
So let's get started.
The mage is a caster. 99,9% of the time, you will want to keep the ennemis as far as possible from you.
Until 75 and the poweful Frostfire bolt, the deep Frost spec is, in my opinion, the best spec to pick up while leveling.
It offers much more control over your ennemies than fire and arcane and can dish out very nice damage.
The spells you will use most of the time as a frost spec mage are Frostbolt, Frost Nova and Ice Lance (at 65). You'll also use Fireblast quite alot.
- Frostbolt : 3 sec cast time damaging spell that slows down your enemy's walking speed. This is your primary nuke spell, not as powerful as fireball (damage-wise) but it will let you solo more easily without worrying much about enemies getting close to you.
- Frost nova : Roots the enemies in place, allowing you to get some distance if they managed to get close to you. Useful against any melee mob and even caster type for the burst it offers with the Shatter talent (see Talents section).
- Ice Lance : Instant low-mana spell that does 3 times its initial damage if the enemy is frozen in place. Can deal HUGE damage and is instant cast. It only comes at 65 though. Also useful to tag a mob.
Also, the following armors will be available depending on your level :
- Ice armor : Slows your enemy when they hit you in melee. Not really the best armor for leveling. It can be favored over mage armor/molten armor if you play on a pvp server and are often attacked while questing.
- Mage Armor : Increases your resistance to magic and allows 30% of your mana to regen while casting. In my opinion, this armor is better than ice armor for leveling, since it will lower your overall downtime.
- Molten armor : Increases your critical chance by 3% and inflicts x damage when you're hit in melee. The critical chance is neat, it means you will kill faster. I personaly used this armor once i got it (level 62).
Let's review the frost tree and the talents that will help you the most. If bolded, it means that you should ABSOLUTELY take the talent.
- Frostbite : Extremely useful talent. It allows your chilling spells (frostbolt, cone of cold, ice armor, improved blizzard, frostfire bolt) to have a 15% chance to root your ennemies in place, like a mini frost nova. Sometimes, you'll be unlucky and it will never proc during the battle, but when it procs, it basically means your ennemy is going to die very soon (See Shatter). It lasts like 5 secs.
- Improved Frostbolt : Frostbolt will be your primary nuke spell until you get Frostfire bolt at 75. You'll cast it more than any other spell and you'll even dream about it too (ok maybe not). Improved frostbolt shortens the cast time of your primary spell by 0.5 sec when maxed. While this may not seem much, it will speed up the leveling process and improve the overall damage you deal to your ennemies. Beside, you'll be glad to have this talent maxed when you will need to Shatter your ennemy and won't have much time left on your Frost Nova or Frostbite proc timers.
- Ice Floes : There is "Survival" written on this talent. Maxed, it shortens the cooldown of your :
- Frost Nova by 5 secs.
- Cone of Cold by 2 secs
- Ice Block by 60 secs
- Icy Veins by 36 secs
- Ice Shards :
Core talent. Improves the damage of your critical spells by 100%.
In short, a spell that does 50 base damage for example will, without Ice Shards, crit for 75 (50% more damage).
With Ice Shards, the same spell will actually crit for 100, so double damage.
Improves your burst damage tremendously.
- Elemental Precision
3% hit when maxed means that you will have less "missed" spells when fighting mobs. This talent is really only useful if you're often taking on enemies that are higher level than you. You shouldn't really have much use of it while leveling but if you growl each time your spells miss, then i guess it wouldn't hurt to take this talent Otherwise it's not really mandatory. The mana reduction alone isn't really worth 3 talent points.
- Permafrost
Your chill effects last 3 sec longer and the walking speed of your enemies is decreased by another 10%.
Not that useful for leveling in my opinion. It allows you to kite the enemies longer but by the the time your enemies get to you, they should be already half dead anyway. It is pretty useful in pvp.
- Piercing Ice
6% more damage on your frost spells, i think it speaks for itself.
- Icy Veins
Activated, this allows you to cast your spells 20% faster without suffering pushback. It is nice when you need to burst down your enemy but i never really used it while leveling up. It is very useful in raids and dongeons though.
- Improved Blizzard
Depending on your playstyle, this could be a must get. Improved Blizzard adds a chill effect to the base spell allowing it to slow your enemies. Coupled with Permafrost (+3 secs chill and +10% slow), this means that the enemies taken in your blizzard are slowed by 75% for 4,5 secondes AND that Frostbite can proc, rooting them in place.
Also note that Blizzard can now crit since patch 3.0.2 (not too sure which patch).
If you like to take on many mobs at once and aoe them down, then this talent definitely is for you. It all depends on your playstyle.
- Arctic Reach
This talent can seem weak but it is very useful while leveling. Being able to cast your opening frostbolt on the enemy from x yards instead of x yards results in more frostbolts launched before it gets to you. It can save you a frost nova that could be useful when fighting the next mob. Also more range on Cone of Cold and Blizzard and a higher Frost Nova radius is very nice, allowing you to avoid hits more easily.
- Frost Chanelling
All your spells cost 10% less mana. This means less downtime while leveling. Also the -10% threat can be nice if you like to do dungeons from time to time.
- Shatter
THE core talent of the frost tree. If you don't put points in it, you'd be better just respecing fire or arcane.
This talent allows you to burst down your enemies everytime they are frozen in place. Your crit chance against them will raise by 50%.
Enemies frozen in place are the ones affected by the following spells : Frost Nova, Frostbite, Hungering Cold (Death Knight) and Frost Trap (Hunter).
Chill effects will NOT affect Shatter. Enemies slowed by a frostbolt won't make your crit chance go up by 50% but if frostbite procs, then it will.
- Cold Snap
Very nice talent that resets all your frost cooldowns. Can save your life if you need to frost nova an enemy chasing you when you're at 5% hp or so, or pop an ice block that was on cooldown to give you some fresh air. It is also a required talent point to unlock the very useful Ice Barrier.
- Improved Cone of Cold
Not really needed while leveling. Beside the "I-hope-frostbite-will-proc" Cone of Cold that you should use while leveling, you'll only use Cone of cold to slow a pack of enemies that are close to you or to kite one enemy while your frost nova is on cooldown.
Cone of Cold can be nice when it procs Frostbite but it's not reliable enough. We don't really use the spell for its damage, so 35% more damage for 3 talent points isn't really worth it imo.
- Cold as Ice
Maxed, it shortens the cooldown of your :
- Cold Snap by 1 min 36 secs.
- Ice Barrier by 6 secs
- Summon Water Elemental by 36 secs
- Winter's Chill
Each time your damaging spells hit, you have 2% more chances to crit. Stacks up to 5 times to make it 10%. Allows you to crit your enemies more often. Take this.
- Shattered Barrier
I personaly don't really like this talent for leveling. I didn't find much use for it and it can sometimes be annoying when the auto frost nova unsheeps an add that was too close without warning.
- Ice Barrier
Your personal shield, that prevents pushback on your spells while it's active. Very useful against those annoying ranged mobs and against a pack of enemies that are too close to you. It takes a % of your spell damage to absorb more. Core talent that will make leveling easier.
- Arctic winds
+5% damage on all your frost spells. It doesn't seem like much but it will make you kill mobs faster. Also the chance to be missed by melee and ranged attack can be very nice. You'll love seeing missed ranged hits
- Empowered Frostbolt
+4% crit and your Frostbolt gains an additional 10% of your spell power. Your primary nuke is now even stronger. Core talent, don't level without it.
- Fingers of Frost
Your chilling spells (Frostbolt, Cone of Cold, Improved Blizzard) have a 15% chance to make your 2 next spells act like your target was frozen in place, even if they are moving. Shatter, anyone ?
Must have. It's basically a 15% chance to improve your crit rate by 50% for your next 2 spells even if your target is not frozen. You will definitely like it.
- Brain Freeze
Burst damage. A free instant cast fireball, even untalented, hurts. Makes your killing even faster.
- Summon Water Elemental
The Water Elemental is a VERY strong guardian. Its bolt hurts alot and its Freeze is basically a frost nova that you can cast anywhere, even 35 yards away from you. It allows very nasty burst damage on demand.
I suggest using the water elemental macro from the guide for your water elemental though, as it can be annoying to click on your pet cast bar. This is how the macro works:
-The first click makes you summon your water elemental on defensive mode and makes it attack your current target.
-If you click on the macro again, 2 things happen : It makes your water elemental attack your other target if you changed targets. It also activates the Freeze ability of your Water Elemental, you still have to target it around your enemy to make it work though.
As you can see, you become very powerful when your elemental is active. You can run to an enemy with your elemental, start casting Frostbolt and just when Frostbolt is about to be launched, use the Freeze ability of your elemental to root your enemy in place. That will make your frostbolt crit almost all the time, chuncking a huge amount of hp off the mob. Even more deadly at 65 when you get Ice Lance.
The elemental also takes a percentage of your spell damage, intellect and stamina.
Too bad it only lasts 45 secs >_<
- Improved Water Elemental
Mainly useful while raiding. Recovers the mana of your raid and your elemental now lasts 1 min. Not very useful while leveling, your enemies will be dead very quickly anyway.
- Chilled to the Bone
Your Frostbolt, Frostfire Bolt and Ice Lance do an additional 5% damage. What else can i say, still more damage on your primary nuke spell as well as Ice Lance. Your enemies also take more time to get close to you <3
We won't use this talent with Frostfire Bolt, i will suggest another spec for it
- Deep Freeze
A 5 secs stun ability only available when your target is frozen in place (Frostbite, Frost Nova, you know the music). Pretty nice, this ability makes you untouchable and lets you solo elites without problems if they are not immune to stuns.
You can make solid combos that prevent your enemies from even moving, like Frostbolt > Frost Nova or Frostbite > Frostbolt + Ice Lance + Deep Freeze, allowing you to cast another full frostbolt while your enemy is stunned (if he isn't dead that's it).
Frost Nova is a spell you'll want available when one or more ennemies managed to get close to you. Same goes for Cone of Cold.
The shortened cooldown on Ice Block has saved my life many times, those seconds you gain with this talent can mean the difference between life and death when fighting mobs and especially elites.
Very useful, juste like Cold Snap and Ice Floes, it adds more survavibility and the shortened cooldown on Summon Water Elemental is very very nice to have while leveling (and anything else actually).
This is how i would put my talent points while leveling up. I'll also explain the cast rotation depending on which spell is available. I'll go up to 75 and then i invite you to read the Frostfire section of the guide.
10-14 : 5/5 Improved Frostbolt
15-17 : 3/3 Frostbite
18-20 : 3/3 Ice Shards
21-23 : 3/3 Icy Floes
24 : 1/3 Piercing Ice
25-27 : 3/3 Shatter
28-29 : 2/2 Arctic Reach
30 : Cold Snap
31-33 : 3/3Frost channeling
34 : 2/3 Piercing Ice
35-37 : 3/3 Winter's Chill
38-39 : 2/2 Cold as Ice
40 : Ice Barrier
41-44 : 4/5 Arctic Winds
45-46 : 2/2 Fingers of Frost
47-48 : 2/2 Empowered Frostbolt
49 : 5/5 Arctic Winds
50 : Summon Water Elemental
51-53 : 3/3 Brain Freeze
54 : 3/3 Piercing Ice
55-59 : 5/5 Chilled to the Bone
60 : Deep Freeze
Now we'll venture a bit in the fire tree until 75 and the Frostfire bolt respec. You can also put points in the frost tree wherever you feel like it.
61-63 : 3/3 Incineration (more crit with Fireblast and Cone of Cold)
64-65 : 2/2 Improved Fireblast (-2 secs on Fireblast CD)
(Improved Fireball is useless anyway, since we only cast it on Brain freeze procs)
66-70 : 5/5 Ignite (more damage on fire crits, even if it's not our primary nukes, it helpsl)
71-73 : 3/3 Impact (your spells have a 10% chance to stun your enemies for 2 secs, just to make you even more unreachable).
74 : Last point before the respec, put it wherever you like to. Burning Determination maybe.
75. Here we are, you can now learn Frostfire Bolt at your mage trainer. I strongly suggest respecing to take full advantage of this spell because it's simply monstruous imo.
I also invite you to head to the Frostfire section for more information on it.
Your cast rotation will vary depending on what level you are and what spells are available in your arsenal.
Note: You should always start at max range of your frostbolt.
Level 1-3 : Cast Fireball from max range and melee the mobs to finish them.
Level 4-5 : Just chain Frostbolt from max range and melee the mobs to finish them.
Level 6-11 : Same as before, but use Fireblast to finish the mobs. If they still live, just cast another Frostbolt / Fireblast or melee them down.
Level 12-25 : Start chaining Frostbolt until your enemy gets close to you. When it does, use Frost Nova (available at level 12) to froze him in place, take some distance and start chaining Frostbolt again.
Level 26-30 : You now have access to Cone of Cold. I like to use it whenever an enemy manages to get close to me because it can proc Frostbite. That will save me a Frost Nova, which has a longer cooldown, and allow me to cast another Frostbolt that will most likely crit, thanks to Shatter.
If it doesn't proc Frostbite, i will then use Frost Nova followed by a Frostbolt or i will use Fireblast to finish the mob, depending on its hp left.
The goal, with Cone of Cold, is to save Frost Nova as much as possible.
Level 30-39 : Same as before, excepted Cold Snap is available, meaning you can Frost Nova your enemies two times in a row to burst them even faster.
Level 40-44 : Ice Barrier is available. You'll have to worry less about being hit and will be able to cast your Frostbolt wihout pushbacks while the barrier is active.
Level 45-49 : You'll start criting even more. Fingers of Frost will speed up your killing tremendously.
Level 50 : Your Water Elemental is now available. I suggest saving it for tricky situations (elites, pack of mobs, etc..).
Level 51-59 : Now you have to monitor the Brain Freeze proc. When it does, launch the Fireball after your nukes. Don't interrupt your spells for a Fireball unless it can finish off your enemy.
Level 60-64 : Deep Freeze is available, meaning you can stun your enemies when they are frozen in place. I suggest stunning them when Frost Nova / Frostbite is about to break.
Level 65-70 : Killing machine : ON. Ice Lance is a crazy spell. It is instant cast, costs almost nothing and does HUGE damage if your enemy is frozen. As soon as you see the enemy frozen in place, be ready to launch an ice lance right after your frostbolt. Those 2 spells critting in a row will remove like 60% of your enemy's health.
Level 71-74 : Your spells can now stun, thanks to Impact. If an enemy manages to gets close to you, it means either that you were AFK or that he is immune to movement-imparing effects
Frostfire Bolt is a unique spell that takes the benefit from fire AND frost at the same time. It's like a fireball that can slow and that benefits from both fire and frost talent trees.
I will list the talents that benefit Frostfire Bolt.
- Ignite : +40% dot on crit
- Impact : 10% stun chance on hit
- Burning Soul : -70% pushback when casting and -10% threat
- Master of elements : 30% of the mana cost of the spell refund when you crit
- Playing with fire : +3% damage
- Critical Mass : +6% crit chance
- Fire Power : +10% damage
- Combustion : see the talent
- Pyromaniac : +3% crit chance
- Molten Fury : +12% more damage if your enemy is below 35% hp
- Empowered Fire : Your frostfire bolt gains an additional 15% of your spell power
- Hot Streak : When you score 2 cricital hits in a row, you get an INSTANT CAST pyroblast
- Burnout : +50% crit damage
- Frostbite : 15% chance to root the enemy in place
- Ice Shards : +100% crit damage
- Elemental Precision : +3% hit
- Piercing Ice : +6% damage
- Icy Veins : Only the faster casting time
- Frost channeling : -10% mana cost (note that FFB doesn't get -20% threat if you take this talent and Burning Soul, it only gets -10% threat max)
- Winter's Chill : +2% crit chance everytime it hits, stacks 5 times for a total of +10% crit chance.
- Arctic Winds : +5% damage
- Fingers of Frost : +15% chance to make your next 2 spells have a 50% more chance to crit (if you took Shatter, which you should)
- Chilled to the Bone : +5% damage and -10% walking speed
As you can see, the spell can benefit from many talents. I suggest going through the Fire path for Burnout, which is a huge dps boost and to take the essential Frost talents going with it.
This is the spec i used at 75 for leveling with Frostfire Bolt: 0/48/18
I then suggest putting your talent points like this :
76-77 : 5/5 Burnout
78-79 : 2/3 Frost Channeling
80 : Weeeeeeeee \o/ (3/3 Frost channeling if you plan to continue questing at 80)
Your frostfire bolt at 80 (0/50/21 pve) :
- has +9% chance to crit (critical mass + pyromaniac)
- does +19% more damage (31% more when the enemy is below 35% hp) (playing with fire + fire power + piercing ice + molten fury when under 35%hp)
- gains an additional 15% of your spell power (empowered fire)
- does +150% damage when it crits, refunding you 30% of its mana cost and also adding a 4 sec dot equal to 40% of the damage done by your crit (e.g. if frostfire bolt crits for 5000, it will also dot the enemy for 2000 in 4 secs. every 2 sec) (ignite + master of elements + ice shards + burnout)
- has +3% chance to hit (elemental precision)
- has a 10% chance to stun the enemy (impact)
- has a 15% chance to freeze the enemy in place allowing Shatter (+50% crit if the next spell lands) (frostbite)
- costs -13% mana (frost channeling + elemental precision)
Also, if you score two critical hits in a row with either Fireblast, Frostfire bolt or Scorch, your next pyroblast cast within 10 secs will be instant.
As you can see, the spell is very very sexy and benefits from soooooooo many talents at the same time that it is really worth a respec for the last 4 levels.
There are 3 useful stats that you will need while leveling and 2 less useful.
Let me start by saying that you should completely avoid STR and AGI unless you're trying to be some sort of battle mage that likes to hit stuff with your staff (yo).
If so, i ... don't think mage is the right class for you ... but if you like it, hey, why not.. you'll just level slower than 90% of the server... make it 99% actually.
Anyway, the main stats you should aim for are :
- Spell power : Increases the damage of all your spells. You should always try to get more as it will improve your overall damage per second.
You won't find many items with spell power in the lower levels but you'll start seeing more and more as you level, especially when you arrive in Outland. The same goes with crit.
- Critical rating : Improves the chances that your spells will critically hit. Crits is as important as spell power is. A crit will improves your killing speed tremendously. If, for example, you loot an item that lowers your spell power by 10 but increases your critical chances by 1%, well, equip it but don't equip an item that increases your crit chance by 0.2% and lowers your spell power by 35. It's all about balancing the both.
- INT : "Int" increases your mana pool as well as your crit chance. The amount of crit you gain from int depends on your level. (There is a nice Rating Buster, i suggest it in the Mage Armor. It's always nice to have some but you can level all the way to 80 without it. You shouldn't really focus much on it until raids.
- STA : Stamina improves your health pool. It's not really useful while leveling BUT if you play on a pvp server, then you might want to have more health to survive an ambush.
Overall, the important stats are spellpower / crit > int > spi > sta.
I recommand those addons while playing.
- Rating Buster : Rating buster converts stats into ratings and allows you to quickly compare two items to see which one is the best for you.
It can, for example, tell you how much critical rating an item with 57 int would actually give you and if that items will make you win or lose crit and spell power if you equip it. A must have in my opinion. It also works with any other class when properly configured.
- Range Display : Small addon that shows the distance between you and the mob. Very useful to know exactly when you are in max range for a frostbolt or any other spell. Easily configurable too.
- Food/Water macro :
#showtooltip nameofthefood
/cast [button:2] Conjure food; nameofthefood
Replace nameofthefood in the macro by the exact name of the food you can summon (shift-click the food to copy the text in the macro).
For example, at level 6, you can start using the macro by right-clicking on it to summon Conjured Muffins. Replace then nameofthefood by Conjured Muffins in the macro to make it work. The macro will be :
#showtooltip Conjured Muffins
/cast [button:2] Conjure food; Conjured Muffins
Right-click on the macro to summon food.
Left click on the macro to eat.
Sames goes for the water.
/cast [button:2] Conjure water; nameofthewater
- Auto-trinket macro.
#showtooltip Frostbolt
/use 13
/use 14
/cast Frostbolt
This macro will automatically try to use equiped trinkets when you cast frostbolt whenever they are available.
You can equip 2 trinkets at most.
Those trinkets go in slot number 13 and 14 in your character sheet.
Note: /use 13 and /use 14 can be placed individually in the macro or both at once.
If, for example, you have a spell power trinket in the first slot and your insigna of the horde/alliance in the second slot, then DON'T put /use 14 in the macro, as it will actually use your insigna every time you try to cast a spell, which you don't really want, do you ?
If you only want to use your first trinket, replace the macro by :
#showtooltip Frostbolt
/use 13
/cast Frostbolt
- Water Elemental macro
Ok, i can't live as a frost mage without this one. It makes life so much easier when you have your water elemental.
#showtootlip Summon Water Elemental
/cast Summon Water Elemental
/cast Freeze
As explained in the talent section :
The first click on the macro will summon your water elemental on defensive mode and make it attack your current target
Another click on the macro will :
- make your pet attack your other target if you swapped targets
- make your pet cast Freeze, which is a ranged Frost Nova that you have to target on the ground just like Blizzard
Coming Soon !
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Talents from lvl 75
Submitted by vtomi92 on Tue, 2018-07-24 10:37.
I've found out that the link with the talents from lvl 75 isn't working anymore. Maybe somebody has it, or somebody can help me with this.
RE: Talents from lvl 75.
Submitted by Emmaleah on Sat, 2018-08-11 19:43.
Things have changed considerably since this was written, which is why it has been moved to the archive section. It was well written at the time and some amount of the content is still useful. That being said, I think the link you refer to was a link to Wowheads talent calculater which doesnt exist anymore in that format, since talents don't exist in that format. If you want current talent / spec information I would suggest Icy Veins or Wowhead.
Emmaleah == bravely leading the way since ... um a long time ago...
Moved to Archive Section
Submitted by Jiyambi on Wed, 2010-10-20 23:22.
We are recently working on improving the quality of content here on WoW-Pro.com, both by limiting low ranked guides and by moving out of date guides into archives. For more information, see the full news post detailing this process.
Your guide has been moved to the archive section for the following reasons:
- The guide is significantly out of date.
If you improve your guide to address these issues, you are welcome to move the guide back to it's proper category. If someone else would like to adopt the guide, please leave a comment.
If you have any questions about this process, feel free to comment here.
Submitted by NYCTexan on Fri, 2010-09-10 03:35.
Maybe it's just me, but I'm highly disappointed at the lack of updated guides. Not just this class, but across the site for the most part. And I know that everyone's going to say, "Oh, but there's no point in updating - until we find out what Cataclysm does!" My answer? That's a load of CRAP... some of these guides are almost a year and a half to two years' old.
Perhaps it's a product of the game itself. I've witnessed this phenomenon a bit myself, within my own guild. The push to raid, the emphasis on gearscores, armor sets, etc., has robbed a lot of the fun. Some of us are NEVER going to see ICC... For whatever reason, desire, play style, time constraints, etc.
Some of us also like making and leveling alts... We could use current, updated information. Just because it's free, shouldn't make it worthless...
Hi there. Now, you are
Submitted by Jiyambi on Fri, 2010-09-10 04:21.
Hi there.
Now, you are entitled to believe whatever you want here, but I'm getting this sense that you think the contributors to this site are doing this full time and getting paid. You seem to think we aren't updating some of these guides because we are being lazy.
In that, you are quite mistaken - we are all gamers in our off time, just like you, and we all have real jobs or school, not to mention friends and family and actually playing the game, that take up time. So you see, we don't have endless hours to devote to the game.
We have to do a little something called prioritizing.
While I'd love to have an infinite amount of time in which to update every single guide on this site (I'm not being sarcastic, I LOVE making guides and that would be pretty awesome) - I just don't. I work and go to school, run a guild, and try to have some kind of social life as well. I just don't have time to update everything, and while there are other contributors to the site, they are often in the same situation as I am. We are all VOLUNTEERS and we volunteer as much as we can, but that's not enough to do everything and make everyone happy.
I am aware that you may have different priorities than I do, and that's fine. But since I'm the one volunteering my time and effort and not you, I get to decide what to update first. And I think for most of our users, we'd rather have some out of date guides now and have everything ready for launch date of cataclysm, than trying to play the catch up game now and have nothing ready at launch. Again, this is an opinion - you're entitled to yours.
Do please think about what you are doing next time, however, when you go on about how disappointed you are with people who are doing something for nothing that you are getting enjoyment out of. None of us are obligated to create this content for you to use - please keep that in mind. I'm not trying to say you shouldn't give suggestions or offer constructive criticism, but to be honest your comment rankled a bit and is a slap in the face for all the authors here on the site working their butts off so you can freely enjoy their creations.
Glyph of Slow Fall
Submitted by Sarok on Wed, 2010-08-04 20:35.
I found glyph of slow fall to be a bit redundant if you master this one skill. When falling, right before you might meet your doom, you blink. Blinking resets your falling position and you don't take as much damage or none at all! Of course you might as well take it since there aren't that many other good minor glyphs.
Glyph suggestion
Submitted by Lycander on Fri, 2010-04-02 01:34.
I've found that the Glyph of Eternal Water has been awesome for leveling. Sure, you lose the rooting ability of the elemental (and the associated frostbolt crit) but you've got a DPS pet helping you the entire way. It's helped my mage out a lot.
A warning on it, though. Anytime you take a taxi, or log out, your pet will be gone. On the plus side, the CD on the summon spell starts as soon as you summon, so if your elemental dies you'll likely be able to summon one right away.
Hope this helps.
Thank you Lycander and
Submitted by Shangra on Mon, 2010-07-19 12:23.
Thank you Lycander and Kwaice for your Glyph tips. I shall start moving my Mage now since I'm abit bored.
This guide is awesome, no
Submitted by runedragonxp on Tue, 2009-12-29 13:55.
This guide is awesome, no doubt on that.
But, the only small problem i found was the rotation,
after level 74, you don't tell us the new rotation for the fire spec,
can you maybe update that?
other than that, it's a great guide:)
Great guide, but I found gramatical error...
Submitted by Young Blood on Mon, 2009-10-05 01:41.
This guide is AMAZING!!! It really helped me out a lot!!!! I did find one small problem. In section 3, for the description of Empowered Icebolt, you wrote "You also gain 0,2 sec casting time" when it should say 0.2 Thanks again. Keep up the great work!! Please.
Actually ...
Submitted by Anaroth on Mon, 2009-10-05 08:50.
Actually we Europeans always spell it like that. Depends on which continent you are I think
Any chance you might be
Submitted by Lery on Tue, 2009-08-25 06:33.
Any chance you might be updating this guide? From what I am reading some people suggest switching to more of a Arcane/Fire build once at level 70 to 80. Then jusing it for raiding at 80.
Submitted by CEO160 on Sun, 2009-08-02 18:39.
I've been working on my macros lately and tried the conjuring food/water macro you have listed. It conjures perfectly but you say to left click to eat and it's not doing it. Any suggestions
Great guide
Submitted by Katharsis on Mon, 2009-06-15 22:40.
Wow, this is a great guide! My alt is a mage, and this is helping a LOT. Thanks!
Also, I'd recommend one, and only one, point in Improved Blizzard, for the reason you spoke about -- it slows AND gives you Frostbite -> Shatter chances. When I'm instancing, this annihilates mobs around the tank.
talent calculator
Submitted by miztajiggles on Tue, 2009-05-12 02:32.
Great guide. Just wanted to say that now whenever i go to the talent calculator link shown after the frostfire blot section, im told access denied and it says i need a username and password. anyone know why this is?
after patch 3.1
Submitted by nukestorm on Thu, 2009-04-16 19:42.
do you still recommend getting winters chill since it only gives 1% extra crit now after patch?
I disagree with the 6-11
Submitted by Nuclear898 on Thu, 2009-04-02 15:48.
I disagree with the 6-11 rotation, its alot better to do a fireball from max range (35yd), then go for frostbolt untill the mob has 1/4 HP then use fireblast and then melee or a last frostbolt, the DoT helps alot.
or maybe i just remembered
Submitted by nukestorm on Thu, 2009-04-02 10:44.
or maybe i just remembered it wrong
Submitted by nukestorm on Thu, 2009-04-02 10:37.
my frost bite talent only lasts about 2 seconds.
Frost AOE
Submitted by Holyterror on Mon, 2009-02-23 22:52.
If you are going to level frost then AOE grinding is the only way to go IMHO. Frostbite = death wish for AOE
It would be nice to have a Frost AOE guide. You can still follow Jame's guide but AOE when you can pull a few (most of the time)
DKs rule...others drool
DKs rule...others drool
Great guide. I do have a
Submitted by Nyon on Fri, 2009-02-20 00:09.
Great guide. I do have a question, however. What rotation do you suggest at 75, after you switch specs to get Frostfire bolt?
blast wave?
Submitted by Endd on Thu, 2009-02-26 02:01.
I'm guessing I shouldn't use pyroblast to start attacking, but at 75 should blast wave replace frost nova, and dragon breath replace cone of cold?
I love the guide
Submitted by Kranky on Mon, 2009-02-16 15:53.
I have been following the guide since level one and it has been a great help.
I just wanted to point out that you get Ice Lance at 66.
a.k.a. Sonnis on Anetheron
Sorry for the long delay;
Submitted by Arkaen on Thu, 2009-02-05 05:07.
Sorry for the long delay; I've been busy recently, I'll finish it soon!
Hi, I'm currently leveling
Submitted by Trollvink on Tue, 2009-02-03 14:26.
I'm currently leveling a mage, 'till level 29. Then I'm gonna twink him. When I looked for a guide for mages, this was the first one I found. I'm using this guide and I love it.
A few things I noticed:
- You can color your links by putting the colour between the text, before the URL code. like this:
[ url=YourLinkHere][ColourTypeHere]TextHere[/color][ /url]
This would be a nice addition for your spells. For instance, blue colour for Frost spells, and red for Fire.
At '6. Gear' you have 'Critical Rating', I think it should be 'Critical strike rating'.
At '6. Gear', again. You say INT, you might want to say the full word, Intellect. Sounds a bit friendlier and nicer for new players, in my opinion. Same with Stamina and Spirit, in that section.
You could replace all of the 'i' with capital 'I' if you mean 'yourself'.
There are alot of other things, but I wont nitpick any longer Outside of that, it's a wonderful guide. Covered everything I would ever need for leveling.
~ Trollvink
Hello, Kwaice. I just wanted
Submitted by Arkaen on Wed, 2009-01-21 05:32.
Hello, Kwaice. I just wanted to offer to proofread the guide for you, if you wanted. I saw a few errors earlier, and I'm a grammar freak sometimes, haha.
Talent build
Submitted by honj93 on Sat, 2009-01-10 14:25.
The other thing I wanted to say (sorry, had no time yesterday) is about your suggestions on talents and especially ice shards.
I agree that it is a core talent for a frost mage, but as at the beginning your crit chance with spells is about 5% and maybe a bit more depending on your gear your damage output will not be that raised by that talent. I suggest waiting utnil you get shatter to optimize the combinations of these two.
Also I would say to take Icy Veins, it's only 1 point anyway and because some people don't like to use spells on cd and hold them for a more difficult situtation I can guarrantee that I'm one of them and even if I feel bad everytime I do it use your spells, the cd will be over faster than you think. In this case I would suggest a macro if you don't feel comfortable clicking each time on spell icon, because if I remember right Icy Veins doesn't trigger the GCD.
And last I would say to not take Cold as Ice immediatly as at this moment it only affects Cold Snap.
So, I would say something like that (although it's not that perfect just to get a general idea):
10-14 : 5/5 Improved Frostbolt
15-17 : 3/3 Frostbite
18-19 : 2/3 Icy Floes
20 : 1/1 Icy Veins
21 : 3/3 Icy Floes
22-24 : 3/3 Icy Shards
25-27 : 3/3 Shatter
28-29 : 2/2 Arctic Reach
30 : 1/1 Cold Snap
31-33 : 3/3 Frost Chanelling
34 : 1/3 Piercing Ice
35-37 : 3/3 Winter's Chill
38-39 : 3/3 Piercing Ice
40 : 1/1 Ice Barrier
41-42 : 2/2 Cold as Ice
From here on just continue as you would, maybe only give less points to Arctic Winds.
P.S. For AoE grinders I stronly recommend :
impr Blizzard ofc
Icy Veins
Shattered Barrier
improved Cone of Cold
frost chaneling as blizzard uses lots of mana
and permafrost is also nice
Nice guide
Submitted by honj93 on Fri, 2009-01-09 12:14.
Quite nice guide and it will probably help many people who want to roll a mage. There are some things I would like to mention though:
1. Buy water. I know this may seem stupid, but when your conjured water is worse than the one you can buy you should buy it it will diminue greatly your downtime (personal experience ). After a certain lvl ( 10 I think) this is the case for lvls 15-20, 25-30, 35-40 and so on. If you have a main you shouldn't have money problems also not if you've picked 2 gathering proffesions (my priest alt has probably made about 20g and is only lvl 13 )
I have something other to say to but I don't have time now, will add it in the afternoon
Nice guide =) Recently
Submitted by Anti on Fri, 2009-01-09 11:02.
Nice guide =)
Recently rolled a mage alt as a side project while waiting for wrath, but had so much fun with him, he's now my main!
Looks like your guide cover's everything well, and the frostfire section was very helpful as i haven't really looked into that spec yet as levelled fully (currently 78) deep frost.
I specked more towards being an AOE machine and levelled from the early 20's to late 40's doing only about 10 quests, the rest AOE grinding. I just thought I'd mention it because i don't really think any mage guide is complete without a mention of the basic AOE cast cycle e.g
Ice barrier / mana shield -> round mobs up -> frost nova -> blink / run -> blizzard-> blizzard -> cone of cold (should be dead, unless they are a few level higher then you, if so start the rotation again)
great video here and the inspiration for me rolling a mage
and a horde version here (but watch the above one first)
if you want me to do a bit of an AOE wright up let me know and ill see what i can do when i get some time,
mana shield??
Submitted by Dicon on Sun, 2009-05-24 15:19.
I would suggest you never ...ever....use mana shield when AoE grinding for the following reason, if your low level AoE grinding your mana pool isn't going to be too big and will allow you to get off 2 Blizz, 2 nova a blink and CoC with very little mana left the reason you need your 2 nova is if you miss a mob or have an add
and need to regroup them, if you where to use mana shield each hit will take mana off you (and you will get hit) leaving you with an OOM situation and 10 almost dead mobs coming at you
If you don't have Ice barrier yet go in with no shield round up nova Blizz Blizz CoC
all mobs dead, you will have little mana left and your HP will be well down but its better that OOM, full HP 10 mobs running at you
Once you get higher level your mana pool will support you using mana shield but then you have Ice barrier so why would you
oh one more thing if mana is an issue forget blink and sidestep away, it might give you that little bit extra to finish them off
Submitted by Harper1990 on Fri, 2009-01-30 10:18.
But don't forget to change the Talents if you do take on an AoE section - after all, can you imagine AoE grinding with Frostbite???!
"Wise is the man who knows he does not know"...
...Or who knows why my mother keeps calling me.
"Wise is the man who knows he does not know"...
...Or who knows why my mother keeps calling me.
I tried this, I like it!
Submitted by Jhaessyn Jardeath on Wed, 2009-01-07 01:48.
I have a mage alt that I've let sit on the wayside. I read this, and redid his talents, and sought out items as you suggested...he rocks! Thanks so much for the helpul tips, and info.
Interested in doing a priest guide now?
To do is to be - Socrates
To be is to do - Voltaire
Do be do be do - Sinatra
To do is to be - Socrates
To be is to do - Voltaire
Do be do be do - Sinatra
Very Nice
Submitted by iRaid on Sat, 2009-01-03 20:24.
Very Nice Guide!!
I've never played a Mage before and after reading this it kind of makes me want to go and make a Mage alt.
After reading through it in
Submitted by Jiyambi on Fri, 2009-01-02 07:43.
After reading through it in more detail, I can say yet again that this looks quite impressive with a lot of really useful tips.
This looks great, and I see
Submitted by Jiyambi on Tue, 2008-12-30 21:49.
This looks great, and I see it has Jame's seal of approval! I have so many alts so these class-specific guides that are coming out are a big help. Thanks for making it!


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Copyright 2018 Mid-Hudson Multiple Listing Service, Inc. All rights reserved. The data relating to real estate
for sale or lease on this web site comes in part from MHMLS. Real estate listings held by brokerage firms other that Weichert
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This data has been updated as of 11/21/2018