Species diversity after prescribed burns at different intensities and seasons in a high altitude Pinus hartwegii forest/Diversidad de especies despues de quemas prescritas en diferentes epocas e intensidades en un bosque de Pinus hartwegii de gran altitud/Diversidade de epecies depois de queimadas prescritas em diferentes epocas e intensidades em um bosque de Pinus hartwegii de grande altitude.
In the Ajusco volcano, in Central Mexico, prescribed burnings of low and high intensity were applied in March and May 2002, along with one unburned control for March and another for May, considering conditions of open stands and closed stands, with the objective of evaluating the vegetation response of the Pinus hartwegii understory. Samplings were conducted every three months during one year. Number of herb and shrub species, density, covering, and frequency were recorded, and the Simpson diversity index was calculated. According to a principal component analysis, the March prescribed burnings of low and high intensity in open stands separate from the other treatments. The t test showed that the March low intensity burns in open stands yielded a higher Spring-diversity than the unburned control. For the shrub species diversity, the March and May high intensity burns in open stands formed a separated group. A [chi square] test revealed the species that appear preferably in burned areas. The results indicate that the use of prescribed burnings no later than March favor the richness and diversity of species of the understory.
KEYWORDS / Fire Ecology / Fire Effects / Forest Fires / Integrated Fire Management / Mexico /
En el volcan Ajusco, en Mexico central, se aplicaron quemas prescritas de baja y de alta intensidad en marzo y mayo de 2002, dejandose un control no quemado para marzo y otro para mayo. Tambien se consideraron condiciones de arbolado abierto y de arbolado cerrado. El objetivo fue evaluar la respuesta del sotobosque de Pinus hartwegii. Se hicieron muestreos cada tres meses durante un ano, registrando el numero de especies herbaceas y arbustivas, su densidad, cobertura y frecuencia, y se calculo el indice de diversidad de Simpson. De acuerdo con un analisis de componentes principales, las quemas prescritas en marzo, tanto a baja como alta intensidad, con arbolado abierto, formaron el grupo con mayores valores. La prueba t indico que la diversidad en primavera fue mayor en el tratamiento de quemas prescritas en marzo a haja intensidad en masas abiertas, con respecto a su control no quemado. En el caso de arbustos el grupo con mayores valores correspondio a las quemas de alta intensidad en marzo y mayo en masas abiertas. La prueba de [chi square] revelo las especies que aparecen preferentemente en localidades quemadas. Los resultados indican que el uso de quemas prescritas no despues de marzo favorece la riqueza y la diversidad de especies del sotobosque.
No vulcao Ajusco, no Mexico central, se aplicaram queimadas prescritas de baixa e de alta intensidade em marco e maio de 2002, deixando-se um controle na queimada para marco e outro para maio. Tambem se consideraram condicoes de arvoredo aberto e de arvoredo fechado. O objetivo foi avaliar a resposta do sorobosque de Pinus hartwegii. Realizaram-se amostras cada tres meses durante um ano, registrando o numero de especies herbaceas e arbustivas, sua densidade, cobertura e frequencia, e se calculou o indice de diversidade de Simpson. De acordo com uma analise de componentes principais, as queimadas prescritas em marco, tanto a baixa como alta intensidade, com arvoredo aberto, formaram o grupo com maiores valores. A prova t indicou que a diversidade na primavera foi maior no tratamento de queimadas prescritas em marco a baixa intensidade em massas abertas, em relacao a seu controle na queimada. No caso de arbustos o grupo com maiores valores correspondeu as queimadas de alta intensidade em marco e maio em massas abertas. A prova de 2 revelou as especies que aparecem preferentemente em localidades queimadas. Os resultados indicam que o uso de queimadas prescritas nao depois de marco favorece a riqueza e a diversidade de especies do sotobosque.
The pine forests of tropical areas host a significant biodiversity. For instance, for Mexico, 47 species and 20 infraspecific taxa are reported (Farjon and Styles, 1997), which is almost half of the ~110 species reported in all the world. Also, some pines are among the tree species reaching the highest altitudes worldwide, and the displacement of the tree line at both high latitude and high altitude forests is an important indicator of global warming. Pinus hartwegii Lindl. is probably the pine reaching the highest altitude in America and one of the highest altitudes in the world. According to Perry (1991) and Farjon et al. (1997) it is found at 2200-4300masl in Mexico, while P. flexilis James reaches altitudes of 3800m in North America (Steele, 1990) and P. wallichiana A. B. Jacks may be found at 4000m at the Himalayas (Sargent et al., 1985).
In pine forests, fire is necessary for the promotion and conservation of richness of species and diversity (Van Lear et al., 1995; Brown, 2000), among other ecological functions. However, the fire responses of tropical pines and their associated species in most of the world are poorly known, including high altitude tropical pines. According to Rodriguez-Trejo and Fule (2003) these are fire-dependant ecosystems, where the alteration of fire regimes by man through its exclusion (implying high intensity wildfires eventually) or excessive fire frequency coupled to rural socio-economic problems; both lead to deforestation, contributing to the reduction of biodiversity.
One alternative to counteract such alterations of fire regimes, is an integrated fire management, involving three key components: prevention and fighting of forest fires, community based-fire management (oriented to address the rural socio-economic issues), and ecological fire management (Rodriguez-Trejo, 2000; Myers, 2006). In order to accomplish integrated fire management, it is necessary to increase the use of prescribed burns coupled with research and monitoring of fire effects, particularly in countries like Mexico, where such information is scarce.
It is necessary to understand the positive effects of fire and make the best use of those effects to improve fire management and its subsequent benefits for the ecosystem and society. Based on the aforesaid, the objectives of the present study are to answer the following questions: a) what are the effects of prescribed fire and wildfires occurring in two different seasons and at two intensities on the richness and diversity of species of the understory of Pinus hartwegii ecosystems?; b) can different fire-treatments be classified in groups, based on principal components, with respect to their effects on the understory?; c) what are the indicative species of burned areas in the understory of P. hartwegii?; and d) what is the feasibility of fire use to promote diversity in these types of forests?
[FIGURE 1 OMITTED]
The study area is located south of Mexico City, Distrito Federal, central Mexico, on the Ajusco volcano. The climate, according to the classification of Koppen modified by Garcia (1981) is type C(w2)(w)(b')ig, semi cold, sub-humid, with rainfalls in the summer (Jun-Nov), mean annual temperature of 5-12[degrees]C, and mean annual precipitation of 1138.62mm. Soils are andosols. The fire season spans winter (Jan) through spring (May). Research was conducted on a slope of the volcano, that has a North-West aspect, mean slope of 55% at 3553-3626masl. The area is characterized by a young forest stand (<8m high) of Pinus hartwegii, and belongs to the Comunidad de San Miguel y Santo Tomas Ajusco. The young forest stand has been there for 14-18 years. The adult trees that originated this cohort were killed during a windstorm. These pine forests are likely maintained by fire (Rzedowski, 1978). Representative genera in the understory include: Festuca, Muhlenbergia, Alchemilla, Geranium, Potentilla, Rumex, Lupinus, and Penstemon (Rzedowski, 1981). However, the effects of fire on plant responses at the community level are not fully understood.
Establishment of the experiment
In March 21st and May 29th 2002, corresponding to mid-forest fire season and its final and severest part, prescribed burns were conducted, leaving non-burned controls for each date. Each plot was 0.6-0.75ha, making a total experimental area of 4.05ha. The coordinates of the Northwest vertex were 19[degrees]12'58.8"N and 99[degrees]16'11.7"W. Each plot had a 2m wide firebreak. No data was taken over a 5m wide perimeter belt on each plot, making an effective separation of 12m between pllots (Figure 1).
The low intensity prescribed burns were initiated by 8:00am as backing fires (against wind and slope). The high intensity prescribed burns were initiated by l:00pm as head fires (in favor of the wind and slope). The rate of spread was measured in 24 separate and randomly placed 10m long experimental units. Flame length was estimated visually with graduated 3m wooden rulers. The atmospheric data were obtained with pocket wind meters (wind speed), sling psychrometers (temperature, relative humidity), and compasses (wind direction). Fine fuels (grasses, needle litter) moisture content was estimated with Sackett tables modified by Burgan and Cohen (unpubl., referred by Rothermel, 1983) and using information on relative humidity, dry bulb temperature, month, fuels exposed or not exposed, time of the day, aspect and slope.
In each fire treatment area, approximately the lower half of the surface was covered with closed stands of juvenile pine trees (900-2500 trees/ha), while the upper half had open stands (300-700 trees/ ha) with large gaps. Crown cover was 0-30% in the open stands, and 70-100% in the dense stands. Mean loading of surface fuels prior to fire were 13.2Mg x [ha.sup.-1] in the open stands, and 11.35Mg x [ha.sup.-1] in the dense stands. The experimental area was not fenced to exclude cattle grazing, because an objective was to study the vegetation response to fire under conditions generally occurring in Mexico. Figure 2 shows the different treatments utilized in the experiment.
[FIGURE 2 OMITTED]
Fire intensity was estimated from flame length using the models of Byram (1959) and Alexander (1982):
L = 0.0775 x [I.sup.0.46]
I = exp((ln L - ln 0.0775)/0.46)
where L: flame length (m) and I: fire intensity (kW x [m.sup.-1]).
Post-fire tree condition (tree survival, mean live crown area and percentage of vertical live crown) was determined in another study in the same plot (Rodriguez-Trejo et al., 2207). In general, the values tended to be lower following May, high-intensity treatments and closed stands. In the unburned plots, first-year survival was equal or higher than 91.3%, while in the March low intensity (both densities) and March high or low intensity in both densities the survival was similar (larger or equal to 91.7%). The lowest survival corresponded to the May high intensity prescribed burn, with 20.9% (closed stands) and 75% (open stands). The mean live crown area by tree was 4.7-5.3m2 for the unburned controls, and 4.4-6.5[m.sup.2] for the March low intensity open stand burn and the March low intensity closed stand burn, respectively. The lowest mean live crown area again was for the May high intensity burn in closed stands (1.6m2), with a low value for the May high intensity burn in open stands. A similar trend was found for the vertical live crown (Table I).
Three months after application of the fire treatments, quadrats fitted and coincident in a vertex, were randomly delimited at each point. The quadrats were placed using a lm coordinate system (X, Y) on each plot and random numbers for determining randomly such coordinates. The dimensions of these quadrats varied according to the life form of the understory species. For forbs and grasses, quadrats of 1m per side (1[m.sup.2]) were used, and for shrubs, quadrats of 4m per side (16[m.sup.2]). The information obtained was species, number of individuals and two perpendicular crown diameters (or covering in percentage in case of forbs and grasses). To determine the total number of species, extensive collecting was carried out in each treatment. Three quadrats of each dimension (two dimensions) per combination of treatment levels (12) were surveyed, which produced 72 quadrats. As these sites were sampled on four occasions (every three months), the total of surveyed quadrats was 288. Sampling intensity in herbaceous vegetation was 0.09%, while for shrubs it was 1.42%. With the field data, the Simpson diversity index (D) was calculated (Krebs, 1978) for forbs and grasses and for shrubs, separately, as
D = 1 - [summation][(pi).sup.2]
pi = n/N
where pi: proportion of individuals of each species, n: number of individuals of each species, and N: number of total individuals of all the species.
The identification of species was performed at the Herbario de Preparatoria Agricola, Universidad Autonoma Chapingo, Mexico. Density, dominance (cover), frequency, as well as relative values of density, dominance, and frequency, were calculated to obtain importance values (Krebs, 1978). The fire treatments were applied in different months; therefore, the quarterly measurements did not correspond to the same time for the treatments of March when compared to those of May. The measurements of the March treatments correspond to Jun, Sep, Dec 2002, and March 2003, whereas those of May were conducted in Aug, Nov 2002, Feb and May 2003.
A principal components analysis was performed using as variables the average herb species diversities (summer, fall, winter and spring) and the shrub species diversities (corresponding to the same four seasons), in separate analysis, because of the different size of the experimental units. The data of the four samplings was used for the analysis. Additionally, a t test was carried out to compare the diversity of pairs of treatments. A non-parametric [chi square] test was employed to determine typical species of burned areas, comparing each treatment with the respective unburned control, and also burned vs non burned plots. The principal components analysis was performed with the procedure proc princomp, the t test with the procedure proc t test, and the non-parametric test with the procedure proc freq, all of SAS (v. 8.0 for microcomputers).
There was a high variability in the fire behavior for each treatment, as natural. High-intensity heading fires had a higher rate of propagation, and higher flame length and fire intensity than low-intensity backing fires. For example, the maximum rate of propagation in the March high-intensity heading fires in open stands was 5 times faster than in the March low-intensity backing fires in open stands. Also, the maximum flame length was 6 times longer and the fire intensity 49 times higher. May treatments also tended to show higher values in these parameters. Comparing the May high-intensity heading fire (that emulated a forest fire) with the March high-intensity heading fire, both in open stands, the former had a 16 times faster propagation rate, 33% larger maximum flame length and 87% higher maximum fire intensity. The fire behavior from each treatment is shown in Table II.
Table III lists the 52 species that were found. After one year following March low-intensity backing fires and high-intensity heading fires, 20 understory species were present in each treatment. In the March control area, there were a total of 13 species (65% of those in burned areas). After one year following the May, high-intensity heading fire, a total of 19 species were encountered, while a total of 20 species were present in areas burned using a May backing fire. By contrast, the May control plot had a total of 14 species (70% with respect to burned areas), showing less richness of species, especially of forbs, without fire (Table IV). Generally, low species richness is typical of high altitude sites like the present study area.
The families with the largest range of representation were as teraceae (21 species) and gramineae (5). The genus with the largest number of species (t) was Senecio.
In the principal components analysis for herbs, eigenvalues indicate that three components (mostly the first) account for approximately 97% of the variance (with 47.1, 74.6 and 97.0%, respectively). The first eigenvector has larger loadings on the third and fourth sampling seasons (0.7051 and 0.6534). The second eigenvector has high loadings on the first and second sampling seasons. This suggests that the first principal component is primarily a measure of the two last sampling seasons (winter and spring) for diversity, and that the second principal component is mainly a measure of the first two sampling seasons (summer and fall). Tables V to VII show the mean and standard deviation of the variables used, the correlation matrix and its eigenvalues, respectively.
The species diversity for all the samplings, as determined by the first principal open condition, are part of the highest values correspond to March prescribed burns at low and high intensities, open condition, and to May low-intensity, open condition, (Table VIII).
Figure 3A presents the component plot, showing the relationship between the first two components. Each observation corresponds to a prescribed fire treatment. The low-intensity prescribed burns in March, open condition, is the group with the highest values, while the group with the lowest values appears to be associated to the May unburned control in open stands. Other treatments that also have very high values in relation to the first component are the March high-intensity burns in open stands, and the May low-intensity burns in closed stands. However, in relation to the second principal component, the two last treatments have very low values, and the highest value is also associated to the March low-intensity prescribed burn in open condition, while the lowest value corresponds to the May high-intensity prescribed burning in closed stands.
The spring diversity (first sampling) was used in the t test because of its higher values. In this case, the only pair of treatments that exhibited statistically signiticam differences were the March low-intensity backing fire, when compared to their unburned control (p=0.0072).
[FIGURE 3 OMITTED]
Regarding the shrub species diversity, the first two components account for 93% of the variance (73.5 and 92.7%) and the first eigenvector has equally large loadings from the second to the fourth sampling seasons (fall, winter and spring, with 0.5174, 0.5462 and 0.5415, respectively). The first sampling (summer) had also a relatively high load (0.3753). For the second component, only the first sampling was relevant (0.8508).
The plot of the first two principal components (Figure 3) shows that the group with the highest values correspond to the May and March high-intensity burns in open and closed stands, respectively, followed by the group May low-intensity in closed stand, March low-intensity in closed stand and the May high-intensity burn in closed stand. Other groups are formed by the rest of treatments (see also Table VIII for the diversity values of each particular sampling). The values of treatments scatter in relation to the second principal component; however, the May high-intensity burns in open stands reach the highest value, and the lowest values correspond to the March burns (high and low intensity) in closed stands. According to the t test in this case the May high-intensity heading fire in open stands had higher spring-diversity than the May unburned controls in open stands.
The species which appear preferably (p[less than or equal to]0.05) in burned areas in different seasons (Table IX), for which they may be considered as indicators are A. procumbens, A. lanuginosa, E. schaffneri, O. jacquiniana, P. gentianoides, P. postrata, and S. callosus. Additionally, the non-parametric test was conducted without taking into account season nor intensity of fire; in other words, simply if it had been burned or not. The species which preferably (p[less than or equal to]0.05) appear in bumed areas (Table IX) are A. procumbens, A. lanuginosa, L. aschenbornii, P. gentianoides, P. postrata, D. jorullensis, S. callosus and S. cinerarioides. S. angulifolius is the species that is observed predominantly in unburned areas (considering burned and unburned areas), with the third and fourth samplings results only being significant for it (p=0.0396 in both cases).
Percentage of importance value
In the March high-intensity prescribed fire, the highest percentage of importance value (PIV) for the herbs corresponded to A. procumbens (values of 22.9-58.1) and F. tolucensis (14.9-45.2) in open as well as in closed condition, throughout the four samplings. In shrubs, S. callosus was the most important one (86.4-90.7 in open, and 24.9-100 in closed condition).
In the low-intensity prescribed burn of March, again A. procumbens and F. tolucensis prevailed among forbs and grasses, and S. callosus among shrub species (Table X). In the March unburned control, the herbaceous species cited previously dominated, and those which had greatest PIV among shrubs, were P. gentianoides and S. callosus (Table XI).
As for the May high-intensity burning, for herbaceous species F. tolucensis had values of 6.0-40.2, and A. procumbens of 27.0-64.2. With respect to shrub species, S. callosus (10.5-100) and P. gentianoides (27.5-64.8) were those with the highest values throughout the samplings.
With respect to low-intensity prescribed burning of May, F. tolucensis, A. procumbens, and E. schaffneri presented the highest PIV (19.1-43.9, 16.8-54.3 and 314.6, respectively), among herbaceous species. In shrubs, S. callosus prevailed with values of 85.4-100 in open condition, and 30-89 in closed condition.
In the May unburned control, again the herbaceous species F. tolucensis and A. procumbens were dominant, with PIV of 14.5-77.2 and 10.9-27.3, respectively. The shrubs P. gentianoides (41.9-100) and S. callosus (58.1-100) dominated in open condition, and Ribes affine (17.8-46.4) and S. angulifolius (50.5-58.5) in closed condition.
In the sites treated with fire, most of the species with greater PIV are typical of burned areas, as shown by the results of the non-parametric test. One of these species is A. procumbens, which, is present in the area of prescribed burns as well as in the control, both of March. In spite of the latter having fewer species, it reaches higher importance values in the first area. Similar behavior (PIV data not shown) is observed in other species, such as P. postrata and L. aschenbornii (March treatments) and A. procumbens, A. lanuginosa, E. schaffneri, L. aschenbornii, S. callosus and S. cinerarioides (May treatments).
The increase in species richness and diversity documented in the burned areas is similar to that observed in North American pine forests of Pinus palustris Mill., Pinus ponderosa or Pinus elliottii var. elliotti (Wright and Bailey, 1982; Wade et al., 2000; Arno, 2000). Fire was shown to play an essential role in Pinus echinata Mili. ecosystems by creating and maintaining open canopy conditions that allow for relatively high diversity understory communities (Sparks, 1998). Periodic fire is considered a key factor for maintaining overall diversity and rare plant species in Pinus elliotii var. densa rockland ecosystems from South Florida (Snyder et al., 1992) and in the Ajusco study area (Table III) two species are rare (G. spathacea and S. paradoxa) and one is endemic (P. ranunculoides; Silva et al., 1999; NOM, 2001). Garcia-Romero (2005) found that after forest fires in tropical high mountain grasslands on the Iztaccihuatl volcano, Mexico, Penstemon gentianoides dominates during the first two years, followed by Lupinus montanus 1.5 to 3 years after the fire, and eventually grasses replace forbs.
Increased richness in burned areas is attributed to species that not only re-sprout from root systems but also germinate from seeds. Furthermore, many of these species require direct solar radiation for germination and growth. Very likely, most of the species are present in seed banks prior to the fire (Whelan, 1997). Baskin and Baskin (1998) reported that fire is an environmental factor that alleviates physical dormancy of numerous species, including several species of Lupinus, and according to our study L. aschenbornii is an abundant species in burned sites. Martin-Martinez (2007) referred for Lupinus bilineatus Benth., also present in Pinus hartwegii forests in the region, 17.5% of germination for untreated seeds and 38.8% when treated with fire.
Biomass accumulation during the fire-free interval creates adverse space, nutrients, water, and light conditions for many associated species, leading to a reduction in the number of species (Grime, 1979). The burned areas in the present study had not been affected by fire for at least five years, which contributed to an increase in number of species after the application of the treatments. The prescribed fires likely reduced aerial competition for light by decreasing mostly the aboveground biomass of grasses, but also of herbs and shrubs, as well as the foliar area of tree crowns. These plants are also likely to depend on fire-induced inputs of nutrients. According to DeBano et al. (1998), ash is rich in cations. This is reminiscent of results after fire in Pinus ponderosa ecosystems in Oregon and Argentinian savannas (Kerns et al., 2006; Kunst et al., 2003), where increased herbs richness and diversity, likely related to post-fire increased resources availability after, respectively, fall burns and mid-fire season burns, in comparison to other seasons of burn and unburned controls.
Diversity appears to be increased by the fire, and withdrawing fire in many ecosystems reduces it (Keane et al., 2002). Likewise, the variability of the fire with respect to season, intensity, pattern, frequency and space creates the greatest diversity in components of the ecosystem (Collins and Gibson, 1990; Swanson et al., 1990; Brown, 2000), which helps explain the different responses found in this study. Moreover, according to Hiers et al. (2000) a variable fire season and other components of the fire regime help maintaining a greater variability of native species and, likely, are a key for conserving biodiversity in the Pinus palustris Mill. ecosystems in particular and in fire-dependent ecosystems in general.
In the same study area at the Ajusco volcano, Rodriguez-Trejo et al. (2007) pointed out that first year tree survival was 96% in the unburned control as well as after the March low-intensity prescribed burn, but it was only 48% after the May high-intensity fire.
The higher diversities associated to open conditions and March prescribed burns are related to the availability of direct solar radiation, nutrients and water because of the reduced competition, in comparison to the closed stands (Whelan, 1997). Also, the March burns corresponded to the beginning of spring, while the May burns corresponded to mid-spring, so that this last treatment implied a delay for the recuperation of the understory species (Brown, 2000).
The results of the present study show the value of using prescribed fire in March at low intensity in order to increase herbs diversity. Nonetheless, the results also point out the importance of varying fire intensity (and fire season) to yield higher shrub species richness. Under conditions common in Mexico, that include a very high incidence of human-caused forest fires, limited resources to suppress fires and with the fire season reaching the highest number of forest fires in April and May in the study area (Gobierno de la Ciudad de Mexico, 2005), it seems most convenient to conduct prescribed fires no later than March. The results suggest that the objectives of increasing species diversity while limiting and reducing hazardous forest fuels can best met with prescribed fires before the end of March.
The areas prescribed-burned in March had more understory species and herbs diversity compared with the unburned control, due to the released growth space, the re-sprouting of many species, and the existing seed bank. The other burn treatments also resulted in an increased number of species, but there was a greater risk of fire escape (because of the season of the intensity of the fire) and tree mortality. The total number of species found (52) was relatively low, due to the area altitude.
Species indicative of burned sites, as indicated by the non parametric test, also presented a high percent importance value in the burned areas, although as the species in these areas are more numerous, lower values would be expected.
According to the principal component analysis, the herb species principal component values tended to be greater in the March-burned sites in open stands. In the case of shrubs, the highest values were obtained following the May and March high-intensity burns, in closed and open stands, respectively.
According to the results, and considering that the vast majority of forest fires in the region are human-caused and the relatively limited resources to fight wildfires, it may be a good strategy to conduct the prescribed burns no later than March. This may go in hand with other objectives like the reduction of hazardous fuels, that would limit the negative environmental impacts (tree mortality, erosion, pollution) of wildfires. Prescribed burning may also be compatible with moderate grazing.
The authors thank the communities of San Miguel and Santo Tomas Ajusco for the permission to conduct this work on their lands, to CONACYT installation project No I35626-B and to the University of Chapingo (Ajusco Project) for financial support, the herbarium of the Preparatoria Agricola, UACH, for the invaluable support identifying the species, and CONAFOR and the Mexico City government for the authorizations and help for conducting this experiment. Finally, thanks to the reviewers. Finally, thanks to the reviewers. Their keen observations improved this work significantly.
Received: 03/01/2007. Modified: 04/03/2008. Accepted: 04/10/2008.
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TABLE I POST-FIRE TREE CONDITION Treatment Survival Mean live Vertical (%) crown area live crown ([m.sup.2]) (%) COs 100.0 4.7 96.8 CCS 91.3 5.3 53.3 MRLOS 91.7 4.4 56.8 MRLCS 100.0 6.5 84.3 MRHOS 75.3 2.9 46.6 MRHCS 91.7 3.1 58.5 MYLOS 82.5 2.6 68.5 MYLCS 94.4 3.7 63.3 MYHOS 75.0 3.3 57.8 MYHCS 20.9 1.6 36.9 C: Unburned control, MR: March burns, MY: May burns, L: low intensity, H: high intensity, OS: open stands, CS: closed stands. TABLE II CHARACTERISTICS OF THE PRESCRIBED BURNING TREATMENTS Treatment Temp Relative ([degrees]C) humidity (%) MRLOS 7-18 49-70 MRLCS MRHOS 16-18 30-49 MRHCS MYLOS 12-18 20-25 MYLCS MYHOS 16-19 12-18 MYHCS Treatment Wind Wind direct. speed (km x [h.sup.-1]) MRLOS N, E [less than or equal to] 12.8 MRLCS MRHOS N [less than or equal to] 20 MRHCS MYLOS N [less than or equal to] 4 MYLCS MYHOS N [less than or equal to] 20 MYHCS Treatment Light fuel Rate of moisture propagation (%) (m x [min.sup.-1]) MRLOS 10-14 0.10-1.00 MRLCS 12-15 0.04-0.08 MRHOS 8-10 0.20-5.00 MRHCS 9-I1 [less than or equal to] 0.19 MYLOS 6-7 0.20-1.00 MYLCS 8-9 [less than or equal to] 0.17 MYHOS 3-4 0.30-80.00 MYHCS 5-6 [less than or equal to] 0.34 Treatment Flame Fire length intensity (m) (kW[m.sup.-1]) MRLOS 0.2-1.0 8-260 MRLCS 0.1-0.4 2-35 * MRHOS 0.5-6.0 58-12770 MRHCS 0.1-0.4 2-35 * MYLOS 0.2-1.0 8-260 MYLCS 0.2-0.4 8-35 * MYHOS 0.5-8.0 58-23867 MYHCS 0.25-0.5 13-58 * MR: March burns, MY: May burns, L: low intensity, H: high intensity, OS: open stands, CS: closed stands. * Estimates of fire intensity were based in flame lengths. Do not include below ground heat pulse for high density stands, where pine needles are the most abundant fuel and is produced below-ground smoldering combustion. TABLE III SPECIES LIST FOR BURNED AND UNBURNED PLOTS IN THE SOUTH OF MEXICO CITY Species Family Arenaria lanuginosa (Michx.) Rohrb Caryophillaceae Cerastium nutans Raf . Caryophillaceae Commelina coelestis * Commelinaceae Baccharis conferta H.B.K. Asteraceae Bidens anthemoides (DC.) Sherff * Asteraceae Cirsium ehrenbergii Sch. Bip. Asteraceae Erigeron galeottii (A. Gray) Greene Asteraceae Eupatorium adenophorum Spreng Asteraceae E. schaffneri Sch. Bip. Asteraceae Gnaphallium americanum Mill. Asteraceae G. inornatum DC. Asteraceae G. spahcilathum H.B.K. Asteraceae Helenium integrifolium (H.B.K.) Benth & Hook Asteraceae Hieracium fendleri Sch. Bip. Asteraceae Selloa plantaginea H.B.K. Asteraceae Senecio angulifolius DC. Asteraceae S. callosus Sch. Bip. Asteraceae S. cinerarioides H. B. K. Asteraceae S. reticulatus DC. Asteraceae S. sanguisorbae DC. * Asteraceae S. toluccanus DC. Asteraceae S. vulgaris L. Asteraceae Sonchus oleraceus L. Asteraceae Stevia hirsuta DC. Asteraceae Draba jorullensis H.B.K. Cruciferae Pernettya postrata (Cav.) CD. Ericaceae Gentiana spathacea H.B.K. * Gentianaceae Geranium potentillaefolium DC. Geraniaceae Agrostis tolucensis H.B.K. * Graminae Brachypodium latifolium Fourn. * Gramineae Calamagrostis tolucensis (H.B.K.) Trin. Gramineae Cinna poaeformis (H.B.K.) Scribn. & Merr. * Gramineae Festuca tolucensis H.B.K. Gramineae Ribes afine H.B.K. Grossulariaceae Phacelia platycarpa (Cav.) Spreng * Hydrophyllaceae Luzula lenticulata Liebm. Juncaceae L. racemosa Desv. Juncaceae Lamium purpureum L. * Labiatae Salvia prunelloides H.B.K. * Labiatae Stenantium frigidum Kunth. * Liliaceae Astragalus micranthus Desv. * Fabaceae Lupinus aschenbornii Scher Fabaceae Oxalis jacquiniana H.B.K. Oxalidaceae Peperomia campylotropa Hill. Piperaceae Plantago linearis Plantaginaceae Pinus hartwegii Lindl. Pinaceae Alchemilla procumbens Rose. Rosaceae Potentilla ranunculoides Humb. & Bompl. Rosaceae Salix paradoxa H.B.K. * Salicaceae Castilleja arvensis Cham. & Schlecht. * Scrophulariaceae Penstemon gentianoides (H.B.K.) Poir Scrophulariaceae Eryngium monocephalum Cav Umbelliferae * Collected in the study area or plots, but not encountered in sampling units. TABLE IV NUMBER OF SPECIES BY TREATMENT Treatment Number of species recorded in quadrats Forbs Shrubs Total and grasses March low-intensity backing fires 16 4 20 March high-intensity heading fires 16 4 20 March unburned control 7 6 13 (65% *) May low-intensity backing fires 14 6 20 May high-intensity heading fires 14 5 19 May unburned control 9 5 14 (70% *) * Percentage with respect to low intensity backing fires. TABLE V SIMPLE STATISTICS FOR DIVERSITY ALONG ALL THE SAMPLINGS First Second Third Fourth Herbs Mean 0.543 0.437 0.414 0.433 Standard deviation 0.163 0.116 0.145 0.106 Shrubs Mean 0.041 0.142 0.167 0.198 Standard deviation 0.104 0.152 0.193 0.193 TABLE VI CORRELATION MATRIX FOR HERBS AND FOR SHRUBS First Second Third Fourth Herbs First 1.0000 0.1096 0.1673 0.0184 Second 1.0000 0.2662 -0.0262 Third 1.0000 0.8241 Fourth 1.0000 Shrubs First 1.0000 0.3445 0.3970 0.6139 Second 1.0000 0.8941 0.7116 Third 1.0000 0.8261 Fourth 1.0000 TABLE VII EIGENVALUES OF THE CORRELATION MATRIX FOR HERBS AND SHRUBS Eigenvalue Difference Proportion Cumulative Herbs 1 1.8856 0.7855 0.4714 0.4714 2 1.001 0.2044 0.2750 0.7464 3 0.8957 0.7771 0.2239 0.9703 4 0.1186 0.0297 1.0000 Shrubs 1 2.9386 2.1666 0.7347 0.7347 2 0.7720 0.5586 0.1930 0.9277 3 0.2134 0.1375 0.0534 0.9810 4 0.0759 0.0190 1.0000 TABLE VIII HERBS AND SHRUBS SPECIES DIVERSITY BY TREATMENT AND SEASON, LISTED IN ORDER OF OVERALL DIVERSITY, AS DETERMINED BY THE FIRST PRINCIPAL COMPONENT Treatment Sampling 1st 2nd 3rd 4th Herbs MYLCS 0.68 0.29 0.16 0.31 MRCCS 0.39 0.32 0.24 0.42 MRCCS 0.45 0.57 0.35 0.27 MYCCS 0.54 0.35 0.33 0.35 MRCCS 0.60 0.58 0.29 0.33 MRCCS 0.41 0.41 0.36 0.40 MYHCS 0.28 0.44 0.46 0.51 MYLCS 0.73 0.50 0.51 0.41 MYHCS 0.57 0.34 0.56 0.51 MYLCS 0.37 0.49 0.57 0.57 MRHOS 0.71 0.32 0.58 0.57 MRLOS 0.78 0.63 0.56 0.55 Treatment Sampling 1st 2nd 3rd 4th Shrubs MRCCS 0.00 0.00 0.00 0.00 MRCCS 0.00 0.00 0.00 0.00 MYLCS 0.00 0.00 0.00 0.12 MYCCS 0.00 0.00 0.00 0.13 MYLCS 0.00 0.00 0.00 0.15 MRLOS 0.00 0.16 0.00 0.00 MRHOS 0.00 0.06 0.14 0.06 MYLCS 0.00 0.28 0.24 0.25 MRCCS 0.00 0.31 0.44 0.20 MYHCS 0.15 0.18 0.38 0.53 MRCCS 0.00 0.41 0.47 0.49 MYHCS 0.34 0.30 0.33 0.45 MR: March burns, MY: May burns, C: unburned control, L: low intensity, H: high intensity, OS: open stands, CS: closed stands. TABLE IX P-VALUES OF THE [chi square] TEST. CHARACTERISTIC SPECIES FROM BURNED AREAS Species Sampling 1st 2nd 3rd 4th Considering season of application and intensity of fire: Alchemilla procumbens ns 0.0300 0.0419 0.0308 Arenaria lanuginosa ns 0.0528 0.0399 ns Eupatorium schaffneri ns 0.0005 0.0300 0.0308 Oxalis jacquiniana ns 0.0059 ns ns Penstemon gentianoides ns 0.0348 0.0348 0.0187 Pemettya postrata ns ns 0.0031 ns Senecio callosus 0.0133 0.0016 0.0002 0.0201 Considering burned and unburned areas: Alchemilla procumbens 0.0171 0.0027 0.0011 0.0007 Arenaria lanuginosa ns 0.0233 0.0049 0.0244 Draba jorullensis ns 0.0578 ns ns Lupinus aschenbornii ns 0.0578 0.0085 0.0407 Penstemon gentianoides ns ns ns 0.0531 Pernettya postrata ns 0.0244 ns ns Senecio callosus ns 0.0038 0.0001 0.0041 Senecio cinerarioides ns ns ns 0.0233 Ns: non significant. TABLE X IMPORTANCE VALUE (%) FOLLOWING BACKING FIRES IN MARCH, FOR THE FOUR SAMPLINGS Species Stand/Sampling Open 1 2 3 4 Forbs and grasses Alchemilla procumberrs 16.5 35.7 30.1 33.3 Arenaria lanuginosa 5.2 2.7 Cerastium nutans Cirsium ehrenbergi 3.1 Draba jorullensis 2.6 Eryngvum monocephallum 5.3 4.0 6.8 3.9 Eupatorium schaffneri 8.4 10.4 Festuca tolucensis 36.2 28.7 44.0 36.6 Gnaphallium inornathum 6.5 5.4 Oxalis jacquiniana 11.1 Pernettva postrata 11.2 10.9 9.1 Potentilla ranunculoides 2.4 4.4 4.4 3.2 Selloea plantaginea 3.3 Senecio toluccanus 3.0 2.6 Senecio rulgaris 2.0 Stenanthium frigidum 3.5 3.2 Total 100 100 100 100 Shrubs Lupinus aschenbornii Penstemon gentianoides 10.6 Senecio callosus 100 89.4 100 100 Senecio cinerarioides Total 100 100 100 100 Species Stand/Sampling Closed 1 2 3 4 Forbs and grasses Alchemilla procumberrs 39.1 53.0 58.5 54.3 Arenaria lanuginosa Cerastium nutans 5.1 Cirsium ehrenbergi Draba jorullensis Eryngvum monocephallum Eupatorium schaffneri 4.2 10.8 17.1 Festuca tolucensis 37.5 16.1 24.4 27.0 Gnaphallium inornathum Oxalis jacquiniana 2.7 Pernettva postrata 14.1 10.2 18.7 Potentilla ranunculoides Selloea plantaginea Senecio toluccanus Senecio rulgaris Stenanthium frigidum 7.2 Total 100 100 100 100 Shrubs Lupinus aschenbornii 34.0 46.8 74.5 Penstemon gentianoides Senecio callosus 66.0 42.3 9.2 Senecio cinerarioides 10.9 16.3 Total 100 100 100 TABLE XI PERCENTAGE OF IMPORTANCE VALUE, MARCH UNBURNED CONTROL, FOR THE FOUR SAMPLINGS Species Stand/Sampling Open 1 2 3 4 Forbs and grasses Alchemilla procumbens 27.5 27.6 12.2 7.6 Eryngium monocephalum 6.6 10.6 11.8 Eupatorium schaffneri 15.4 Festuca tolucensis 53.3 44.9 77.2 73.0 Pernettya postrata 19.3 5.4 7.5 Senecio reticulatus Stenanthium frigidum Total 100 100 100 100 Shrubs Lupinus aschenbornii 48.0 Penstemon gentianoides 100 52.0 Ribes affine Senecio angulifolius Senecio callosus 100 100 Senecio cinerarioides Total 100 100 100 100 Species Stand/Sampling Closed 1 2 3 4 Forbs and grasses Alchemilla procumbens 7.3 21.7 24.7 24.0 Eryngium monocephalum 4.5 6.9 Eupatorium schaffneri 6.0 16.6 Festuca tolucensis 64.4 52.1 68.4 76.0 Pernettya postrata 14.8 Senecio reticulatus 5.0 Stenanthium frigidum 7.5 Total 100 100 100 100 Shrubs Lupinus aschenbornii Penstemon gentianoides Ribes affine 12.8 20.3 66.6 Senecio angulifolius 58.5 33.4 Senecio callosus 87.2 Senecio cinerarioides 21.1 Total 0 100 100 100