Table 1.

Significant Spearman's rank correlations (p) of bee abundance and species richness with habitat variables following application of a Bonferroni correction (a level 0.05/14 = 0.004) for treatment units (thin only, burn only, thin + burn, and control) in a pinyonjuniper woodland, Arizona. Nonsignificant findings are not shown. Total bees Habitat variable May Flower cover (per m^^) Number of flowering species
August

needed to determine long-term effects of fuets-mitigation treatments on pollinator assemblages in pinyon-juniper woodlands.

Acknowledgments
I wish to thank Dave Hutinian for bis suppoti oi this project, field assistants Nora Sutherland and Windy Edgar for tbeir bard wotk and enthusiasm, Rachel Biggs for her meticulous data entry, and Dave Egan for his editorial review.

Bee species

0.58 0.51

< 0.0001 0.0002

0.54 < 0.0001 0.48 0.0005

References
Huffman, D.W., RZ. Fule, J.E. Ctouse and K.M. Pearson. 2009. A comparison of fite hazard mitigation alternatives in pinyonjuniper woodlands of Arizona. Forest Ecology and Management 257:628-635. Huffman, D.W., PZ. Eul, K.M. Pearson and J.E. Crouse. 2008. Fire history of pinyon-juniper woodlands at upper ecotones with ponderosa pine forests in Arizona and New Mexico. Canadian Journal of Forest Research 38:2097-2108. Kendall, D.M. 2003. Effects of fire on insect communities in pion-juniper woodlands in Mesa Verde country. Pages 279285 /'" M.L. Floyd (ed). Ancient Pinon-Juniper Woodlands: A Natural History ofMesa Verdi Country. Boulder. University Press of Colorado. Kleintjes, P.K., B.F. Jacobs and S.M. Fettig. 2004. Initial response of butterfiies to an overstory reduction and slash mulching treatment of a degraded pion-juniper woodland. Restoration Vo/ogy 12:231-238. Larson, B.M.H., P.G. Kevan and D.W. Inouye. 2001. Flies and flowers: Taxonomic diversity of anthopbiles and pollinators. Canadian Entomologist 133:439465. Michener, C D . 2000. The Bees ofthe World. Baltimore MD: The John Hopkins University Press.

Flower cover (per m^) % cover litter % canopy cover

0.44 -0.41 -0.44

0.0020 0.0034 0.0016

variables, was most strongly correlated witb bee abundance and species richness (Table 1). The bigb flower cover in tbe tbin plus burn treatments may be attributed to the more open canopy found in these units, since canopy cover in tbe thin only and thin plus burn treatments was significantly lower than in the control and burn only plots {p < 0.0001). Cover of soil and litter did not differ significantly among treatments, indicating that these nesting substrates are equally available in both treated and untreated areas. The amount of coarse woody debris and number of snags did differ significantly among treatments, but neither substrate was correlated with bee abundance or diversity. This result is unsurprising, since most ofthe solitary species observed are from ground-nesting genera. There were no significant treatment effects on fly abundance and richness, or correlations with measured habitat variables for either sampling period. For land managers seeking to reduce hazardous fiiel loads and simlutaneously support insect pollinators, overstory thinning followed by prescribed burning may be the best option. A study of fuels-mitigation alternatives at this site (Huffman et al. 2009), suggests that tbin only or thinning plus prescribed burning treatments are the most effective in terms of fire hazard reduction, and the highest abundance and diversity of bees were observed in tbin plus burn units during this study. In addition, "lethal patches," in which high-intensity fire kills all ofthe overstory trees in a limited area, may be botb typical of pinyon-juniper woodlands (Huffman et al. 2008) and of benefit to pollinating insects. Treatment units at the study site were imbedded within a partially treated matrix (approximately two-thirds thinned and burned, one-tbird either burned or untreated) including several unintentional "lethal patches." Soil substrates in these areas were powdery and probably unsuitable as a substrate for ground-nesting bees, but they did support large patches of floral resources (mostly composites) that attracted many insects, including honeybees and bumblebees that were rarely observed on plots. Further studies are

Germination and Survival of Tree Seeds in a Tropical Montane Forest Restoration Study (Costa Rica)
Gabriel C. Sady (Environmental Studies Dept, University of California, Santa Cruz, CA 95064, gahriel.sady@gmail .com), Karen D. Holl (Environmental Studies Dept, University of California, Santa Cruz, kholl@ucsc.edu), Rebecca J. Cole (Environmental Studies Dept, University of California, Santa Cruz, cole.rebeccaj@gmail.com) and Rakan A. Zahawi (Organization for Tropical Studies, Apdo 73-8257, San Vito de Coto Brus, Costa Rica, zak.zahawi@ots.ac.cr) lanting tree seedlings is a common strategy to facilitate tropical forest recovery, but it is resource intensive. Planting trees in small patches or "island" as a restoration strategy (Zahawi and Augspurger 2006, Cole et ai., forthcoming) imitates the nucleation process that occurs naturally over longer time scales from random dispersal events. Moreover, the island planting strategy is less expensive 28:2 121

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Table 1. Mean ( SD) tree canopy and ground cover, surface soil temperature (daily maximum and minimum), and light (photosynthetically active radiationPAR values were averaged from 5:30 a.m. to 5:30 p.m.) In four microhabitats (Agua Buena, Costa Rica). Values with the same letter are not significantiy different across microhabitats.
Variable Canopy cover (%) Grass cover (%) Forb cover (%) Bare ground (%) Max, soil temp (C) Min. soil temp ("C) PAR (fjmoi/mVs) Control 1.7 3.4 62.9 42.4' 31.5 29.6" 11.5 13.8 21.0 2.4 17.1 0.9" 235 118' Island Edge 84.4 13.2" 32.2 32.4" 49.5 34.4' 32.2 19.8'" 29.7 3.P 13.6 1.5 157 78 Island Interior 96.9 2.r 21.3 29.8" 40.0 25.0 49.7 18.7" 22.1 1.4 18.8 1.4^ 217 98 Plantation 95.7 3.5^ 6.4 4.0" 24.4 13.9" 76.7 15.7= 21.8 4 . 4 ' 17,8 1.2*^ 186 1 70

than plantations and therefore may be more practical for restoring large areas, but there has been little study of its long-term efl^ectiveness. Several factors limit tropical forest recovery, including lack of seed dispersal and germination, competition from pasture grasses and other ruderal vegetation, and microclimatic extremes (reviewed in Holl 2002b). A large-scale, long-term restoration experiment in southern Costa Rica is testing the strategy of planting tree islands to overcome these limitations and facilitate tropical forest recovery (Cole et al., forthcoming). As part ofthis re.search, we studied seed germination and survival of three small-seeded (< 1 cm dia) tree species that naturally disperse into abandoned pastures during early succession (Cole et al., forthcoming) to determine which planting strategy best facilitates seedling establishment. We conducted the study at four experimental restoration sites near Agua Buena, Coto Brus, Costa Rica (8''44'36" N, 8258'04" W) from July 2007 through June 2008. All sites had been used for more than 18 years for agriculture, until 2004 when we established in each site three 50 m x 50 m treatment plots: control (no seedlings planted), tree plantation (entire plot planted with 313 seedlings spaced 2.8 m apart), and tree islands. The island treatment plot included two large islands (12 m x 12 m) used in this study that were each planted with 25 trees (16 timber trees and 9 softwoods). Two softwoods, poro [Erythrina poeppigiana) and guaba {Inga edulis)^ are ni trogen-fixers that are widely used as shade trees in cofl^ee plantations. Mayo bianco ( Vochysiaguatemalensis) and amarillon ( Terminalia amazonia) are slower-growing timber species used extensively for reforestation in the region. These four native or naturalized species were used in both planting treatments. See Cole et al. (forthcoming) for more experimental design details. We collected seeds from fruiting trees in the area of three small-seeded species: Tabebuia rosea (wind-dispersed). Cassiafistulaand Psidium guajava (both animal-dispersed). We planted 2,592 seeds without pretreating them on 24-29 June 2007 (early rainy season) in four microhabitats: plantation (P); control (C); interior (I) and at the edge (E) of large tree islands. Seeds were planted more than 5 m from the edge of plantation and control plots and at least 3 m from the edge in island interior plots. In each 122 June 2010 ECOLOGICAL RESTORATION

microhabitat, we planted two separate 1 m^ quadrats per species, each with 81 seeds 10 cm apart in a 9 x 9 grid. We placed seeds on top of the leaf litter to simulate natural dispersal. Seeds were also planted in a shade house near one site to determine timing of germination. We recorded germination in the field one week after planting and monitored germination of new seeds and survival of seedlings every two weeks through August 2007. Thereafter, we monitored seed germination and seedling survival at about three-month intervals through June 2008, when we also measured seedling height. During 6-15 March 2008 for each seed quadrat, we estimated percent cover classes of grasses, forbs, and bare ground (0%, > l % - 5 % , > 5%-10%, > 10%-25%, > 25%-50%, > 5O%-75%, > 75%-95%, and > 9 5 % 100%) and used midpoints for analyses. We also quantified canopy cover with a densiometet. For three days at each site February-^March 2008, surface soil temperature (2 cm depth) and photosynthetically active radiation (50 cm height) were recorded every five seconds and averaged for each 30-minute period using sensors (LI-COR Biosciences, Lincoln NE) set out in each of the microhabitats. Survival was calculated as a percentage of already germinated seeds. We used a randomized complete block ANOVA and Tukey's multiple comparison procedure to test the effect of microhabitat on most variables, and multivariate repeated measures ANOVA to compare seedling survival over time. We used stepwise regression to test whether understory vegetation or canopy cover explained differences in germination, survival, and seedling height. Survival data were arcsine-square root transformed and seedling height data were log transformed. Overstory cover was highest in the plantation and island interior, slightly lower at the island edge, and extremely low in the control microhabitat (Table 1). Percentage of grass cover was highest in the control and lowest in the plantation, with intermediate values at island edges and interiors; bare ground showed the reverse trend. At the island edge, maximum soil temperature was higher and minimum temperature was lower than in other tnicrohabitats. Light (photosynthetically active radiation) levels were highly variable within microhabitats and did not differ significantly (Table 1).

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high grass cover [P- = 0.42, F= 9.3, df = 1,26,; = 0.009), whereas Tabebuia seedling survival was low in most plots. g 40 Cassia seedlings were significantly taller in the plantation ab after 1 year (Figure lC). Tabebuia seedling height showed a 0 30 similar trend but it was not significant due to high variance. "o c Result uFthis and previous sttidies clearly show that pas1 20 ture grass competition limits seedling survival and growth (D 0 10 (Chapman et al. 1999, Holl 2002a). Therefore, restoration efforts in abandoned agricultural land in the tropics should o aim to reduce grass cover. One of the most effective ways to achieve this goal is to establish a tree canopy that shades 50 out light-demanding grasses. Our results are consistent with past research (e.g., Hooper et al. 2002, Doust et al. 2006) showing that gerr 30 mination of small-seeded species in restored tropical forests is generally low beneath a range of canopy environments, and this, therefore, is one factor that will slow recovery. I 10 ^ Higher Cassia seed germination at island edges suggests that w intermediate canopy cover favors seedling establishment in 0 this species. The intermediate canopy cover of the island restoration strategy created broader daily temperature 0.8 changes than in other microhabitats, which may have triggered germination. r 0.6 I Other factors besides microclimate, such as high herbiv> ^ ory and seed prdation, may have reduced germination and $ 0.4 survival. Smaller seedlings may also have been smothered m by leaf litter (Vzquez-Yanes and Orozco-Segovia 1993). .^ 0.2 a Seed prdation and poor seed-soil contact (Doust et al. X 2006) may explain the complete lack of germination in ND 0.0 the field for Psidium, as seeds germinated in a shade house Control Edge Interior Plantation in trays with soil. Island The low seed germination and seedling survival suggest Figure 1. Mean ( SD) percent germination (A), survival (B), and height that small seeds must be dispersed or seeded in large quan(C) after one year of Cassia fistuloso and Tabebuia rosea seeds across tities for successful establishment in recovering tropical four microhabitats: control (abandoned pasture), tree island edge and forests (Holl 2002b, Hooper et al. 2002). Planting tree interior, and tree plantation (Agua Buena, Costa Rica). Values with the same letter are not significantly different; ND = no data. seedlings, using either the island or plantation strategy, serves to reduce grass cover and enhance survival and Overall field germination was 20.6% for Cassia and growth of naturally establishing seedlings. The higher 17.7% for Tahebuia. Psidium seeds germinated in the shade seedling growth in both plantations and island interiors house but not in the field. More Cassia seeds germinated was likely due to the planting of the two nitrbgen-fixing at island edges compared to other microhabitats, whereas tree species, which may function as nurse trees (see Holl Tabebuia germination did not differ across microhabitats 2002b). The island-planting strategy creates more heterogeneous abiotic conditions, which are likely to better mimic (Figure lA). . There was a trend toward higher Cassia survival in the tropical forests which are quite patchy naturally. island interior and plantation microhabitats after 1 year, but it was not significant given the high variance (Figure Acknowledgments IB). Tabebuia seedling survival was higher at island edges We are grateful for field research help from R. Gomez. M. Loik, after six months, but by the end of the study few seedlings J.L. Reid, and J.A. Rosales. Financial support for this project was survived in any treatment, and none survived in the control provided by a NSF grant (DEB 05-15577), the Richard A. Cooley (Figure 1B). 'Ihe number of seedlings remaining after one Memorial Endowment, and UCSC College Eight Project Funds. year did not differ by treatment for either species {Cassia: p = 0.474; Tabebuia: p - 0.349). Given low germination References and survival, only a small percentage of seeds resulted in Chapman, C.A., LJ. Chapman, L. Kaufman and A.E. Zanne. surviving seedlings {Cassia: 3.7% 5.1%; Tabebuia: 0.88% 1999. Potential causes of arrested succession in Kibale 1.7%). Cassia seedlings had lower survival in areas of 50
ft 1 I Cassia azzz2 Tabebuia

1 20

fl

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123

We used these exclosures to evaluate the short-term effects of cattle exclusion on the herbaceous understory in a ponderosa pine forest. We established ten paired plots consisting of a grazing treatment exclosure and a similarly sized grazed control. Of the ten pairs, five were located in a thinned and burned forest and five in unrestored areas. The plots were dispersed across the 1,200 ha Little Spring pasture of the Mt. Logan allotment, where elevations range Ponderosa Pine Understory Response to from 2,000 to 2,250 m, and the mean annual precipitation Short-Term Grazing Exclusion (Arizona) from 2002 to 2007 was 413 mm (WRGG 2007). Gattle Christopher D. Sorensen (School of forestry, Northern Ari- grazing occurs on the Mt. Logan allotment only from July zona University, Box 15018, Flagstaff, AZ 86011) and through October at a maximum stocking rate of 13.6 ha/ Christopher M. McGlone (Ecological Restoration Institute, AUM (Whit Bunting, BLM, pers. comm.). In 2006. howNorthern Arizona University, Box 15017, Flagstaff, AZ ever, grazing pressure was reduced by a break in a fence that 8601L 928/556-9004, chris.mcgbne@nau.edu) allowed cattle to stray into the adjacent Tuweap allotment. The unrestored sites within the project are predomiivestock grazing is a pervasive land use practice on public lands in rhe American Southwest, making it an nantly dense stands of ponderosa pine with thick litter important variable in land management practices. While and duff layers and a sparse understory. The restored ateas most grazing allotments are stocked each year, shorr rest were thinned and burned to emulate pre-1870 tree density periods (1-2 years) are used ro minimize potential negative and distribution. Restoration treatment areas have a more impacts of grazing and to sustain range resources (Curtin diverse and abundant understory community and substan2002). Moore and others (1999) specifically recommend tially fewer trees per hectare (399.2) than the prerestoration deferment of livestock grazing on ponderosa pine {Pinus conditions (784.6) and the untreated controls (873.5). All ponderosa) forests after thinning and burning as ecological restoration treatments on our study plots were completed restoration treatments. Few data are available, however, by 2001 (see Roccaforte et al. 2009 for details), and cattle on the influence of short-term grazing exclusion on the were reintroduced in the summer of 2002 (Whit Bunting, understory plant community of pine forest restoration BLM, pers. comm.). The dominant tree species is ponderosa pine, though projects. In this study, we attempt to quantify the impacts there is a considerable amount of Gambel oak {Quercus of cattle grazing on plant cover, species richness, and nonnative species on a ponderosa pine restoration project in gambelii) on site. The main native species are silver lupine {Lupinus argenteus) and the important perennial forage northern Arizona. grasses bottlebrush squirreltail {Elymus elymoides) and westIn die summer of 2005, the Arizona Game and Fish Department erected several 35 m x 35 m cattle exclosures at ern wheatgrass {Pascopyrum smithii). In 2003, cheatgrass the Mt. Trumbull, Arizona, ecological restoration research {Bromus tectorum) invaded the site. Gheatgrass is the most site on the Grand Ganyon-Parashant National Monument. abundant non-native species present, and this annual grass

National Park, Uganda: Growth and mortality of seedlings. African journal of Ecobgy 37:81-92. Cole, R.J., K.D. Holl and R.A. Zahawi. Forthcoming. Seed rain under cree islands planted to restore degraded lands in a tropical agricultural landscape. Ecological Applications. Doust, S.j., P.D. Erskine and D. Lamb. 2006. Direct seeding to restore rainforest species: Microsite effects on the early establishment and growth of rainforest tree seedlings on degraded land in the wet tropics of Australia. Forest Ecobgy and Management 234:333-343. Holl, K.D. 2002a. Eifect of shrubs on tree seedling establishment in abandoned tropical pasture. Journal ofEcology 90:179-187. . 2002b. Tropical moist forest. Pages 539-558 in M.R. Perrow and A.J. Davy (eds). Handbook of Ecological Restoration. Cambridge UK: Cambridge University Press. Hooper, E., R. Condit and P. Legendre. 2002. Responses of 20 native tree species to reforestation strategies for abandoned farmland in Panama. Ecological Applications 12:1626-1641. Vzquez-Yanes, C. and A. Orozco-Segovia. 1993. Patterns of seed longevity and germination in the tropical rainforest. Annual Review of Ecology and Systematics 24:69-87. Zahawi, R.A. and C.K. Augspurger. 2006. Tropical forest restoration: Tree islands as recruitment foci in degraded lands of Honduras. Ecological Applications 16:464-478.

Ungrazed ' Conlroi Ung razed ' Treated Grazed * Control Grazed * Treated

Figure 1. Median (25th, 7Sth percentiies) cheatgrass cover per plot, separated by both grazing and restoration treatments. Results of Kruskal-Wallis ranked signs analysis were significant for year x grazing x restoration (treated vs. control) interaction ( x ' = 32.01, p = 0.001 ).

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