Growth and Recovery of Oak-Saw Palmetto Scrub through Ten Years After Fire Paul A. SChmalzer
RESEARCH ARTICLE ABSTRACT: Oak-saw palmetto scrub, a shrub community of acid, sandy, well-drained soils in Florida, is maintained by periodic, intense fIre. Understanding the direction and rates of changes in scrub composition and structure after fire is important to management decisions. We followed changes in vegeta.tion along 15Cm linecintercepttransects that were established in 1983. Two stands (8 transects) burned in a prescribed fire in December 1986; the stands had previously burned 11 y (N=4) and 7 y (N=4) before. We sampled transects at 6, 12, 18, and 24 rna and then annually through 10 y after the • 1986 fire. We measured cover by species in two height classes, > 0.5 m and < 0.5 m, and measured height at four points (0, 5, 10, and 15 m) along each transect. Saw palmetto cover equaled preburn values by· one year postburn and changed little after that. Cover of oaks > 0.5 m (Quercus myrtifolia, Q. Growth and geminata, Q. chapmanii) equaled preburn values by 5 y postburn and changed little by 10 Y postburn. Height growth continued, increasing from a mean of 84.0 cm at 5 y to 125.9 cm at 10 y postburn. Bare ground declined to <2% by 3 Y postburn. Plant species richness increased slightly after fire and then Recovery of Oak gradually declined. These vegetation changes alter habitat conditions for threatened and endangered Saw Pal metto Scrub animals and plants. Crecimiento y Recuperaci6n del 'Oak-Saw Palmetto Scrub' Durante Diez Anos through Ten Years Despues del Fuego
RESUMEN: EI 'Oak-saw palmetto scrub' es una comunidad de arbustos de suelos :kidos, arenosos y After Fire bien drenados en Florida, mantenidos por fuegos peri6dicos e intensos. Conocer la direcci6n y las tasa de cambio en la composici6n y la estructura del matorral despues del fuego es. irnportante para las decisiones de manejo. Seguimos los cambios en la vegetaci6n a 10 largo de transectas de intersecci6n linear de 15 m que fueron establecidas en 1983. Dos lotes (8 transectas) quemadas en un fuego Paul A. Schmalzerl programado en Diciembre 1986; los lotes fueron quemados previamente hacia 11 afios (N=4) y 7 afios Dynamac Corporation (N=4). Muestreamos las transectas a los 6, 12, 18 Y 24 meses y luego anualmente durante 10 afios Mail Code DYN-2 despues del fuego de 1986. Medimos la cobertura de las especies en dos c1ases de altura, >0,5m y <0,5 m, y medimos la altura en cuatro puntos (0, 5, 10 y 15 m) a 10 largo de cada transecta. La cobertura post Kennedy Space Center, FL 32899 USA fuego de 'saw palmetto' se igua16 a la cobertura pre quema al afio y cambi6 poco despues de eso. La cobertura de robles >0,5 m (Quercus myrtifolia, Q. geminata, Q. chapmanii) igual6 los valores previos a las 5 afios y cambi6 muy poco hasta los 10 afios posteriores a la quema. EI crecimiento en altura • continuR aumentando, de una media de 84 cm a los 5 afios a 125,9 cm a los 10 afios posteriores a la quema. El suelo descubierto disminuy6 a <2% durante 3 afios posteriores a la quema. La riqueza de especies de plantas aument6 levemente despues del fuego y luego gradualmente disminuy6. Estos cambios en la vegetaci6n alteraron las condiciones para los animales y plantas amenazados 0 en peligro.
Index terms: Florida scrub, prescribed burning, vegetation recovery
INTRODUCTION ment of presuppression conditions in all landscapes (Abrahamson and Abraham Florida scrub is a rare and declining eco son 1996a, 1996b; Duncan et al. 1999). system (Myers 1990, Menges 1999) and is Long-unbmned scrub may become fIre habitat for a number of threatened and resistant and require combinations of me endangered plant and animal species chanical treatment and bmning for resto (Christman and Judd 1990, Stout and ration (Schmalzer and Boyle 1998, Marion 1993, Stout 2001). Management Schmalzer and Adrian 2001). of remaining scrub is critical to the surviv al of these species. Scrub is. a fIre-main- Florida scrub is characterized by evergreen, . tainedsystem (Myers-1990, Menges 1999), xeromorphic shrubs including oaks (QueT" . but landscape fragmentation and fIre sup cus L. spp.), repent palms (Serenoa repens pression have reduced fIre size and fre [W. Bartram] Small, Sabal etonia Swingle quency (Myers 1990, Duncan and ex Nash), and ericaceous shrubs (Lyonia 1 Author e-mail: Schmalzer 2001). Prescribed burning is spp., Vaccinium spp.). Scrub ecosystems [email protected] the primary management technique for vary regionally (Myers 1990, Schmalzer scrub reserves (Menges 1999); however, et al. 1999) and along local environmental Natilral Areas Journal 23:5-13 prescribed burning after decades of fIre gradients (Abrahamson et al. 1984, suppression does not result in reestablish- Schmalzer and Hinkle 1992b). Specific
Volume 23 (1),2003 Natural Areas Journal 5 types of scrub include sand pine scrub, anN '<1"1.0«: r<)t-r<)0\ 00 oak scrub, rosemary scrub, and scrubby t- Ic.ic
r<)~ O\anO\an "! OONO Recovery of oak scrub and scrubby flat Ic An understanding of the direction and rates an r<) r<)~' METHODS A December 1986 prescribed fIre burned 10 transects in two scrub stands. We had 6 Natural Areas Journal Volume 23 (1),2003 established 15-m line-intercept 8N~8 r-"" transects in these stands origi ION 0 I 0 0 nally in January 1983 and resa mpled them in January 1985. The two stands were 11 y (N=4) and 00 1.0 0\ 7 y (N=6) from the previous fire. I~O I 10 We sampled cover of vascular plants by species on the burned transects at 6, 12, 18, and 24 mo ~c-r: tr) 0 ...... 1100 IN I postfire and then annually through ten years after the 1986 fire. Cover by species was sam pled in 0-0.5 m and 0.5 m ...... > 001 lvi height layers (Muelller-Dombois and Ellenberg 1974); cover was estimated to the nearest 5 cm. o <,)""""",,8 V) "" Bare ground was recorded when e 000 0000 I~O there was no plant cover in ei ~ ther height strata. Nonvascular plants (mosses, lichens) were "" ...... 8r-~N ...... ~o ...... ~ uncommon at this site. We re 001 10000 NO I ...... N000 I corded vegetation height at four points (0, 5, 10, and 15 m) along each transect. Previous analysis ~N ...... r-'N 1.0 Nerioocio I leri (Schmalzer and Hinkle 1992a, 1992b) indicated that two transects were wetter and had ~ little or no cover of scrub oaks; Nr-Or-O 1.0 1.0 V) V) 000~0""'00 I 0 I leri 100\ those transects are not consid V) ered here. One transect burned in November 1994; the sample 800\ ...... 1.08 o 00 V) size for the last three time-since 000~~ ..... 00 I 0"" I leri I~oci fire periods was reduced to sev en. The last sampling included here is from January 1997. All 8~~N 0\""1.0 0\r-1oO ...... 0N~V) transects burned again in June 10 ...... Noo~0 100000eriOo\o...... V) ...... 1997. ~ We analyzed data using SPSS r-1oO~1000 0\ 800.00.~ ...... ool~~ 0010 10 NO 10.0 (SPSS, Inc. 1999). Normal ...... ity of distributions was tested using Kolmogorov-Smirnov o ""1.0 8<,)r-08v)""N .,-{C'f),.--(OO tests; most were not normal ONO~O e 1 ..... 0 I 0 o\erioo~o\ V) ...... (p<0.05). For selected parame ~ ters, a nonparametric repeated measures analysis of variance r 00 "" (ANOVA) (Friedman test) was oo I N ""01 NO conducted' usirrgSigmaStat 2.0 (Fox et al. 1995); if the ANOVA was significant, postburn peri ods were compared to preburn using Dunnett's test. RESULTS Before the 1986 fire, saw pal- Volume 23 (1),2003 Natural Areas Journal 7 metto, myrtle oak, and fetterbush (Lyonia lucida [Lam.] K. Koch) were the species A. with the greatest cover in the > 0.5 m stratum in these stands (Table 1). Sand live and Chapman oaks were relatively com 60.------__~ mon, with lesser amounts of other erica • Serenoa repens ceous shrubs and few herbs (Table 1, Ta ble 2). Mean height was about 108 cm 50 (Table 1). Nearly all the species present before the 40 fire were present by 6 mo postburn (Table "#.--- 1, Table 2); the only exception was Hy '-"' .... 30 pericum reductum (Svenson) W.P. Adams, which had been present in minor quanti ()~ ties (Table 2). A few herbs and subshrubs 20 appeared after burning that were not re corded previously: Carphephorus spp., Galactia elliottii Nutt., Panicum spp., and 10 Pteridium aquilinum (L.) Kuhn (Table 2). * The subshrub dwarf huckleberry, Gaylus- accia dumosa (Andrews) Torr. & A. Gray, O~----._----_r----_.------._----,_----_.----~r_--~ also occurred after burning (Table 2). Most o 20 40 60 80 100 120 140 woody species (e.g., Quercus myrtifolia) showed increased cover in the < O.S m Time Since Burn (months) stratum (Table 2) followed by increased cover in the> O.S m stratum for the larger shrubs (Table 1). Cover of shiny blueberry B. (Vaccinium myrsinites Lam.) increased, and the increase persisted for most of the peri- od (Table 2). 30 ,--______--, Dominant species showed different growth • Lyonia lucida rates after fire. Cover of saw palmetto >O.S 25 m tall was reduced 6 mo postburn but was similar to preburn by 12 mo postburn (Fig 20 ure lA). Saw palmetto cover varied within anarrow range between 1 and 10 y post bum (Table 1, Figure lA). The repeated "#.--- 15 '-"'.... measures ANOVA was significant m > (p<0.001), but only the 6-mo postburn o 10 sample differed from preburn (p 8 Natural Areas Journal Volume 23 (1),2003 ------~------ measures ANOVA was significant A. (p<0.001), and the 6-, 12-, 18-, and 24-mo 60rr======J------~ p()s!burn samples _differed _from preburn • Quercusmyrlifolia (p<0.05). Cover (>0.5 m tall) of sand live 501~------~ oak (Figure 2B) was reduced significantly 6 and 12 mo postburn (ANOVA, p DISCUSSION ~~---r----._--_,----,---_,----._---.--~ Oak-saw palmetto scrub displays remark o 20. 40 60 80 100 120 140 able resilience at the fire return intervals Time Since Burn (months) experienced during this study. All the spe cies that occuu-ed before burning exhibit Figure 2. Changes in percent cover of myrtle oak (A), sand live oak (B), and Chapman oak (C) from preburn (time = 0) through 10 y (120 mol postburn. Sample size is n = 8 transects from preburn through resprouting life histories, although some 7 y postburn and n = 7 for the last 3 y. Shown are means and 95% confidence intervals. Repeated are also capable of seeding and clonal measures ANOVA indicated significant (p<0.001) differences among time periods. Asterisks indicate spread (Menges and Kohfeldt 1995). Re- time periods different from preburn (Dunnett's test, p<0.05). Volume 23 (1),2003 Natural Areas Journal 9 sprouting and the high proportion of be lowground biomass (Johnson et al. 1986, A. Guerin 1993) allow rapid reestablishment 160 of cover of the dominant shrubs. However, Cover >0.5 m the growth rates of individual species dif 140 • fer such that there are short-term shifts in dominance; in particular, saw palmetto 120 reestablishes cover more rapidly than the I woody shrubs. By 4 to 5 y after fire, cover ..-.. I of major species is nearly identical to that ?F. 100 '-" I I before burning. Few changes occur through .... I Q) ! > 10 y postburn except for continued height 0 80 I () growth. These observations support and extend earlier work showing rapid recov ~ 60 ery of this vegetation (Schmalzer and Hin II kle 1992b) and are similar to trends in 40 postfrre recovery of scrubby flatwoods on ! the Lake Wales Ridge (Abrahamson 20 * 1984a). These observations also fit the ! conceptual model of vegetation dynamics * of xeric uplands in peninsular Florida de 0 veloped by Menges and Hawkes (1998) as 0 20 40 60 80 100 120 140 the type scrubby flatwoods, which are per Time Since Burn (months) sistent under moderately frequent fires (5- 20 y). Extended periods without fire allow struc B. tural changes toward xeric hammock 80 (Menges et al. 1993, Schmalzeret al. 1994, Menges and Hawkes 1998). Height growth Cover <0.5 m continues, scrub oaks become frre-resis 70 • tant, and fires may not readily reverse these conditions, particularly under typical pre 60 scribed conditions (Abrahamson and Abra hamson 1996a, 1996b; Schmalzer and ~ 50 Boyle 1998; Schmalzer and Adrian 2001). '-".... Q) Growth rates vary regionally (Schmalzer > 0 40 and Adrian 2001), and the modal fire fre () I quency may be less than 20 years in some ro.... * * 0 30 scrub and scrubby flatwoods of the Atlan I- tic Coastal Ridge, Merritt Island, and Cape Canaveral due to more rapid increases in 20 I cover and height. * t I 10 ! ! ! ! Sprouting from underground buds (e.g., oaks, ericaceous shrubs) or buds protected 0 by plant structures (e.g., saw palmetto) 0 20 40 60 80 100 120 140 occurs in many fire-prone ecosystems Time Since Burn (months) (Whelan 1995, Bond and van Wilgen 1996). In many fire-adapted shrublands there is a range of regeneration patterns Figure 3. Changes in total percent cover >0.5 m tall (A) and total percent cover <0.5 m tall (B) from from obligate seeders to obligate sprouters preburn (time =0) through 10 y (120 mo) postburn. Sample size is n =8 transects from preburn (e.g., Keeley and Zedler 1978, Christens through 7 y postburn and n = 7 for the last 3 y. Shown are means and 95% confidence intervals. Total en 1985, van Wilgen and Forsyth 1992). cover is the sum of cover values of individual species. Repeated measures ANOV A indicated significant (p 10 Natural Areas Journal Volume 23 (1),2003 ------._------ 1981) and Quercus coccifera garrigue 35 * (Malanson and Trabaud 1987, 1988) is similar to that of oak-saw palmetto scrub. 30 Bellingham and Sparrow (2000) suggest 25 ed a general model in which resprouting (aboveground) is the predominant response ...-.. ~ 20 * to the least severe disturbance regimes; but resprouting is also a common response "0c:: ::J in disturbance regimes of high severity e 15 C!) that destroy most or all aboveground bio mass and which occur at medium to high ~ 10 ctl L. frequency. Very frequent severe disturbanc CO es would tend to eliminate woody shrubs 5 by preventing sufficient carbohydrate stor If * * age between disturbances. Sites with low 0 • • • I productivity (e.g., xeric, infertile) also fa vor resprouting. The disturbance response -5 of oak-saw palmetto scrub is generally o 20 40 60 80 100 120 140 consistent with this modeL Fire in scrub is severe in that it kills most aboveground Time Since Burn (months) biomass. It is of intermediate frequency; Figure 4. Change in percent bare ground from preburn (time = 0) through 10 y (120 mo) postburn. very frequent fires on xeric sites in Florida Sample size is n = 8 transects from preburn through 7 y postburn and n = 7 for the last 3 y. Shown are favor sandhill vegetation dominated by means and 95% confidence intervals. Repeated measures ANOVA indicated significant (p<0.001) grasses (and the fIre resistant Pinus palus differences among time periods. Asterisks indicate time periods different from preburn (Dunnett's test, p<0.05). tris Mill.) rather than shrubs (Myers 1985, Menges and Hawkes 1998). One consequence of the rapid reestablish ment of shrub cover is the rapid decrease in amount of bare ground (open sand or 160 gaps). Young and Menges (1999) showed that gaps and open sand declined rapidly 140 after fIre in scrubby flatwoods on the Lake Wales Ridge. Openings are important for many scrub herbs and small shrubs, in 120 cluding endemic species (Hawkes and Menges 1996, Menges and Hawkes 1998, 100 I Boyle et aL 2001). Although scrub on E I I Merritt Island lacks the endemic flora of .s- I ...... c: 80 I the Lake Wales Ridge (Schmalzer and ·mOJ f ! Hinkle 1992a), there are species that re :c 60 I * * quire openings such as Selaginella areni * cola Underw., Lechea divaricata Shuttlew. 40 IH ex Britton, and Lechea cernua SmalL These species did not occur on this study site but ! * are present in other scrub sites on Men-itt 20 * Island. -- --- 0 The temporary openings created by burn 40 60 80 100 120 140 0 20 ing did allow some expansion of herbs and Time Since Burn (months) small shrubs in this scrub. Modest increas es in cover by .4ristida stricta, Carphep Figure 5. Vegetation height from preburn (time = 0) through 10 y (120 mo) postburn. Sample size is n horus spp., Galactia elliottii, and Panicum = 8 transects from preburn through 7 y postburn and n = 7 for the last 3 y. Shown are means and 95% confidence intervals. Repeated measures ANOVA indicated significant (p<0.001) differences among spp. may have occurred by sprouting time periods. Asterisks indicate time periods different from preburn (Dunnett's test, p<0.05). (Menges and Kohfeldt 1995) followed by Volume 23 (1),2003 Natural Areas Journal 11 increased growth in the more open condi (Woolfenden and Fitzpatrick 1984, Brein example of the southern Lake Wales Ridge. tions that persisted for a few years. Gay inger et al. 1998), and they close quickly Florida Scientist 47:209-250. lusaccia dumosa and Vaccinium myrsin in this scrub. Prescribed burning regimes Bellingham, PJ., and A.D. Sparrow. 2000. ites sprout and spread clonally (Menges that will maintain an open landscape but Resprouting as a life history strategy in and Kohfeldt 1995). Abrahamson (1984b) leave sufficient scrub patches at optimum woody plant communities. Oikos 89:409- 416. found that they increased rapidly and then height is one of the management challeng declined after fIre. Here the increase was es in this system. Bond, WJ., and B.W. van Wilgen. 1996. Fire and Plants. Chapman & Hall, New York. less dramatic but persisted longer, partic 263 pp. ularly for the Vaccinium. ACKNOWLEDGMENTS Boyle, O.D., S.K. Maliakal, M. Petru, and B.S. Menges. 2001. Gapology in the Florida Analysis utilizing historic photography This study was conducted under NASA scrub: incorporating spatial and demograph (Duncan et al. 1999) showed that open contracts NAS 10-10285, NAS 10-11624, ic approaches. Pp. 35-37 in D. Zattau, ed., ings in scrub on Merritt Island were much and NASlO-12180. I thank Dr. W. Knott, Proceedings of the Florida Scrub Sympo more common in the 1943 landscape. NASAIKSC Biological Sciences Branch, sium 2001. U.S. Fish and Wildlife Service, Openings declined during two decades of and B. SummerfIeld, NASAIKSC Safety, Jacksonville, Fla. 63 pp. fIre suppression. Prescribed burning since Health and Environmental Division, for Breininger, D.R., and D.M. Oddy. 2001. Fire 1981 has reestablished openings in a scrub their continued support and L. Maull, J. and Florida Scrub-Jay source-sink dynam by flatwoods landscape (with pines) but Mailander, and S. Turek for fIeld assis ics in mesic flatwoods. Pp. 3-7 in in D. Zattau, ed., Proceedings of the Florida Scrub not in a scrub landscape (lacking pines). tance. Trade names and commercial enter Symposium 2001. U.S. Fish and Wildlife More effective fIre suppression in the scrub prises or products are mentioned for infor Service, Jacksonville, Fla. 63 pp. landscape may have contributed to the mation purposes only; no endorsement by Breininger, D.R, and P.A. Schmalzer. 1990. greater loss of openings. In the scrubby NASA is implied by their use. Dr. E. Meng Effects of fire and disturbance on plants and flatwoods landscape the presence of dead es and an anonymous reviewer provided animals in a Florida oak/palmetto scrub. pine wood on the ground and pine stumps useful comments on an earlier draft of the American Midland Naturalist 123:64-74. may have caused local hot spots contribut manuscript. Breininger, D.R., V.L., Larson, B.W. Duncan, ing to more persistant openings (Myers and RB. Smith. 1998. Linking habitat suit 1990). ability to demographic success in Florida Paul A. Schmalzer is a Plant Ecologist scrub-jays. Wildlife Society Bulletin 26: 118- with the Dynamac Corporation at Kennedy The rapid recovery to previous conditions 128. Space Center, Florida. His research inter occurs in scrub only when the sprouting Breininger, D.R., M.A. Burgman, and B.M. ests include fire ecology, restoration of ability of the dominant shrubs remains Stith. 1999. Influence of habitat quality, scrub ecosystems, and barrier island veg catastrophes, and population size on extinc intact. Disturbances that remove roots and etation. tion risk of the Florida Scrub-Jay. Wildlife rhizomes of oaks, palmettos, and erica Society Bulletin 27:810-822. ceous shrubs cause long-lasting changes Christensen, N.L. 1985. Shrubland fire regimes in composition and structure (Breininger LITERATURE CITED and their evolutionary consequences. Pp. and Schmalzer 1990, Schmalzer et al. Abrahamson, W.G. 1984a. 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