The role of animals in the colonization of the Krakatau Islands by fig trees (Ficus species)

Abstract. Since the biologically extirpating eruption of Krakatau (Sunda Strait) in 1883, Rakata (Krakatau's remnant) and two closely adjacent islands, Sertung and Panjang, have been colonized by over 200 species of vascular plants. They now carry species-poor mixed tropical forest, including some twenty-three species of Ficus . Data on the sequence of colonization over the last century by twenty-four Ficus species, twenty-three species of volant frugivores, and by agaonid fig-wasps, presumably from the large islands of Java and Sumatra, each some 44 km distant, are summarized. The potential of the volant frugivores as dispersers of fig seeds is assessed, the pollination problems involved in the colonization of islands by figs are reviewed and patterns of colonization by fig species and by their bird and bat dispersers are identified and discussed.
In 1930 a new island, Anak Krakatau, emerged from Krakatau's submerged caldera. This active volcano suffered a self-sterilizing eruption in 1952/1953 and has been colonized, under considerable constraint from its own volcanic activity, probably largely from the (selected) species pool present on Rakata, Sertung and Panjang, 2–4 km away. Its vegetation is at an earlier successional stage (grassland and Casuarina woodland) than that of the three older islands, and in 1992 the Casuarina woodland was in an early stage of transition to mixed forest. The colonization of Anak Krakatau by Ficus species, agaonid wasps and volant frugivores over a critical decade (1982–92) is reviewed, including preliminary assessments of the effects of pollinator limitation on four pioneer fig species and indications of a possible effect of the presence of avian raptors, particularly the peregrine falcon, on fig colonization and forest diversification.     

Interesting times on Krakatau: stand dynamics in the 1990s

The Krakatau Islands, Indonesia, have provided an opportunity for ecologists to track primary succession from the 'clean slate' of 1883, through forest closure in the 1920s, to the contemporary period, in which successional changes take the form of alterations in composition and stature of forest stands rather than gross changes in ecosystem type. This paper reports on permanent forest plots established on the islands in 1989, and fully surveyed again in both 1992 and 1997. Since 1989, the plots have been subject to natural disturbance phenomena in the form of varying combinations of, for example, deposition of volcanic ejecta, landslides, lightning strikes, storm damage and drought. These effects have been concentrated between 1992 and 1997, during which the volcano Anak Krakatau has deposited ash on the islands of Sertung and Panjang, but not on Rakata. Data on stand responses are presented for growth rates (dbh (diameter at breast height, 1.3 m) increment), stem recruitment and mortality, biomass changes partitioned into mortality, ingrowth and growth of established trees), and compositional shifts. The discussion focuses on evaluation of questions and successional models framed earlier in the programme. One general finding is that the stand dominants as of 1989 have tended to decline in number within the plots, generally through low levels of recruitment failing to balance rates of mortality. The effects of disturbance to the plots appear to be evident in terms of mortality and recruitment, dbh increment, and changes in biomass. The patterns of change in the eight plots are quite varied, such that relatively few generalizations are possible. The difficulties of establishing meaningful baseline rates for tree growth and stand biomass are discussed.     

Tropical fruitflies (Diptera: Tephritidae) of the Krakatau Archipelago in 1990 and comments on faunistic changes since 1982

Three of the Krakatau islands, in Sunda Strait, some 40 km from both Java and Sumatra, are believed to have been totally devastated by the famous cataclysmic eruption of 1883. The fourth, Anak Krakatau, is an emergent volcanic island which rose from Krakatau's submerged 1883 caldera in 1930, suffered a self-devastating eruption in 1952/3 and several severely damaging ones since then. In 1990 the tephritid fauna was monitored on all the islands with Steiner traps. Nine species were found, five of which appeared to have been found by Yukawa in 1982 (Yukawa 1984) using similar methods and lures, when he noted a total of six species. As in 1982, the fauna in 1990 was characterized by the numerical dominance ofBactrocera albistrigata (de Meijere) although this appeared to be less marked than in 1982 andBactrocera papayae Drew & Hancock appeared to have increased in relative numbers. The distribution of three species over the islands of the archipelago had increased and four species not found in 1982 were present. Rakata, Krakatau's remnant and by far the largest and highest island, carried seven species (six in 1982), Sertung three (two in 1982), Panjang four (one in 1982) and Anak Krakatau four (one in 1982). Capture rates were lower than in 1982 but still higher than those obtained on the Javan mainland in 1982. There was a marked difference between the tephritid faunas of Anak Krakatau's two forelands, which were at different stages of biotic succession.     

Mammals,reptiles and crabs on the Krakatau Islands: Their roles in the ecosystem

In 1982 I studied the changes which had taken place in the fauna of the Krakatau Islands over the last fifty years, and elucidated the roles of terrestrial consumers and reducers in the developing ecosystems of these islands. Only two species of rat were found (Rattus rattus andR. tiomanicus) as observed by Dammerman (1948) in 1933, butR. tiomanicus had newly established its population on Sertung. No rats were found on Anak Krakatau. Although many skinks (Mabuya multifasciata) were observed on R. Besar, this species was not found on the other islands. The rat and skink are considered strong competitors to the sand crab (Ocypode kuhli), because while the crab was restricted to the sandy beach on the three islands where rat and skink live it had expanded inland on Anak Krakatau where there are no rats and skinks. The two banded monitor (Varanus sarvator) was found on every island and it mainly fed on crabs (Gecarcoidea natalis andMetasesarma aubryi). Thus it is suggested that the crab plays an important role in the process of faunal succession on the Krakatau Islands.     

Rapid recovery of genetic diversity of stomatopod populations on Krakatau: temporal and spatial scales of marine larval dispersal

Although the recovery of terrestrial communities shattered by the massive eruption of Krakatau in 1883 has been well chronicled, the fate of marine populations has been largely ignored. We examined patterns of genetic diversity in populations of two coral reef-dwelling mantis shrimp, Haptosquilla pulchella and Haptosquilla glyptocercus (Stomatopoda: Protosquillidae), on the islands of Anak Krakatau and Rakata. Genetic surveys of mitochondrial cytochrome oxidase c (subunit 1) in these populations revealed remarkably high levels of haplotypic and nucleotide diversity that were comparable with undisturbed populations throughout the Indo-Pacific. Recolonization and rapid recovery of genetic diversity in the Krakatau populations indicates that larval dispersal from multiple and diverse source populations contributes substantially to the demographics of local populations over intermediate temporal (tens to hundreds of years) and spatial scales (tens to hundreds of kilometres). Natural experiments such as Krakatau provide an excellent mechanism to investigate marine larval dispersal and connectivity. Results from stomatopods indicate that marine reserves should be spaced no more than 50-100 km apart to facilitate ecological connectivity via larval dispersal.     

Insect conservation in early succession on islands: lessons from Surtsey, Iceland, and the Krakatau Archipelago, Indonesia

Early successional communities on islands may include specialised aeolian-based pioneer arthropods scavenging on aerial fallout on volcanic lava or ash. Some such species appear to be restricted to such extreme habitats. Persistence of such species may be governed by volcanic activity and their future thus be beyond the control of people. Surveys to determine their presence elsewhere are needed to determine their possible status as refugees in later successional stages, and as a basis for any more informed conservation measures. Insect colonization patterns on Surtsey and Anak Krakatau are briefly reviewed and contrasted to indicate the background to insect community formation on remote islands.     

The recolonization of the Krakatau islands by termites (Isoptera), and their biogeographical origins

The termite assemblages of the Krakatau islands and five sites in Sumatra and Java were investigated, using a standardized transect sampling method. Five species were found on the Krakatau islands that have not been collected there before. The total number of termites recorded on the islands since the 1883 volcanic eruption is now twelve species, all of which are wood feeders. No termites nesting in or feeding on the soil were found on the Krakataus, suggesting that rafting in wood is the primary method by which the termites recolonized the islands.     

Effects of a predator control experiment on Grey-winged Francolin (Scleroptila africanus) populations

One of the key findings of the Grey-winged Francolin Research Project conducted on the Stormberg Plateau, Eastern Cape Province, South Africa during 1988–1992 was that predation, particularly by small mammals on incubating female Greywinged Francolin Scleroptila africanus and their nests, was a major cause of mortality. To assess the response to this predation, small mammal predators were trapped intensively during the francolin breeding season in two 500ha plots, one on the farm Buffels Fontein and one on the farm Boschoff's Kraal. Trapping was not simultaneous on the two plots, but rather during 1992–1994 on Buffels Fontein and during 1995 and 1996 on Boschoff's Kraal. The Yellow Mongoose Cynictis penicillata was the most commonly trapped predator species (83%) in both plots. Grey-winged Francolin numbers during trapping in the plot at Buffels Fontein declined less during a drought than those in the untrapped control plot at Boschoff's Kraal. This suggests that the control of predators may buffer the negative effects of drought on Grey-winged Francolin reproductive success. However, the increases of populations in both plots during a period of higher-than-average rainfall indicate that Grey-winged Francolin can withstand predation during normal rainfall years.     

Influence of Different Intensities of Pesticide Management on the Structure of Ground Beetle Communities (Coleoptera; Carabidae) During a Crop Rotation

Investigations were lead through in the years 1992 to 1995 at Halle (Saale) on a stand 24ha in size. Aim of the survey was to record the effects of graduate intensities of pest management on ground beetles. For this purpose 6 plots were installed on the field, 72 ‐ 200m each. Two plots served as control areas, without any application of pesticides. On two other plots intensive chemical pest management was applied. On the last two plots treatments were lead through considering the economic thresholds for weeds, fungal and insect pests according to the rules of integrated pest management. Ground beetles were sampled by using pitfall traps. In 1992 complementary methods to the pitfall trap catches, pick up of beetles after pesticide applications and semi-field-tests, on all plots were lead through. The graduate pest management was carried out 1992 and 1993 in winter wheat and 1994 in sugar beet stands. The last crop in 1995 was summer barley. In this year all plots were treated conventionally. In 1992 the differences between the total catches of ground beetles on the experimental sites were not high. In the following year most species were predominant on the control sites. In 1994 in sugar beet mechanical weed control was carried out on the control plots. The intensity of pesticide management between intensive and integrated plots differs only small. The amounts of ground beetles reached similar values on all experimental sites. In contrast to the expections the trap catches of ground beetles in the final investigations in summer barley were highest in the plots which had been treated integrated in the years before. The smallest amount was sampled in the former intensively managed sites. Renunciation of pesticides often is connected with economic losses which are not tolerable. If management is conducted considering the economic thresholds corresponding to the principles of integrated pest management negative effects on the economy of nature should not be expected for the longer term.     

Pollinating fig wasps: genetic consequences of island recolonization

The levels of genetic diversity and gene flow may influence the long-term persistence of populations. Using microsatellite markers, we investigated genetic diversity and genetic differentiation in island (Krakatau archipelago, Indonesia) and mainland (Java and Sumatra, Indonesia) populations of Liporrhopalum tentacularis and Ceratosolen bisulcatus, the fig wasp pollinators of two dioecious Ficus (fig tree) species. Genetic diversity in Krakatau archipelago populations was similar to that found on the mainland. Population differentiation between mainland coastal sites and the Krakatau islands was weak in both wasp species, indicating that the intervening 40 km across open sea may not be a barrier for wasp gene flow (dispersal) and colonization of the islands. Surprisingly, mainland populations of the fig waSPS may be more genetically isolated than the islands, as gene flow between populations on the Javan mainland differed between the two wasp species. Contrasting growth forms and relative 'immunity' to the effects of deforestation in their host fig trees may account for these differences.     

A casualty of climate change? Loss of freshwater forest islands on Florida's Gulf Coast

Sea level rise elicits short‐ and long‐term changes in coastal plant communities by altering the physical conditions that affect ecosystem processes and species distributions. While the effects of sea level rise on salt marshes and mangroves are well studied, we focus on its effects on coastal islands of freshwater forest in Florida's Big Bend region, extending a dataset initiated in 1992. In 2014–2015, we evaluated tree survival, regeneration, and understory composition in 13 previously established plots located along a tidal creek; 10 plots are on forest islands surrounded by salt marsh, and three are in continuous forest. Earlier studies found that salt stress from increased tidal flooding prevented tree regeneration in frequently flooded forest islands. Between 1992 and 2014, tidal flooding of forest islands increased by 22%–117%, corresponding with declines in tree species richness, regeneration, and survival of the dominant tree species, Sabal palmetto (cabbage palm) and Juniperus virginiana (southern red cedar). Rates of S. palmetto and J. virginiana mortality increased nonlinearly over time on the six most frequently flooded islands, while salt marsh herbs and shrubs replaced forest understory vegetation along a tidal flooding gradient. Frequencies of tidal flooding, rates of tree mortality, and understory composition in continuous forest stands remained relatively stable, but tree regeneration substantially declined. Long‐term trends identified in this study demonstrate the effect of sea level rise on spatial and temporal community reassembly trajectories that are dynamically re‐shaping the unique coastal landscape of the Big Bend.     

Alfalfa mosaic and cucumber mosaic virus infection in chickpea and lentil: incidence and seed transmission

Samples collected in 1994 and 1995 from commercial crops of chickpeas and lentils growing in the agricultural region of south-west Western Australia were tested for infection with alfalfa mosaic (AMV) and cucumber mosaic (CMV) viruses, and for members of the family Potyviridae using enzyme-linked immunosorbent assay (ELISA). In 1994 no virus was detected in the 21 chickpea crops tested but in 1995, out of 42 crops, AMV was found in two and CMV in seven. With lentils, AMV and/or CMV was found in three out of 14 crops in 1994 and 4 out of 13 in 1995, both viruses being detected in two crops in each year. Similar tests on samples from chickpea and lentil crops and plots growing at experimental sites, revealed more frequent infection with both viruses. No potyvirus infection was found in chickpeas or lentils in agricultural areas either in commercial crops or at experimental sites. However, bean yellow mosaic virus (BYMV) was detected along with AMV and CMV in irrigated plots of chickpeas and lentils at a site in Perth. When samples of seed from infected crops or plots of chickpeas and lentils were germinated and leaves or roots of seedlings tested for virus infection by ELISA, AMV and CMV were found to be seed-borne in both while BYMV was seed-borne in lentils. The rates of transmission found through seed of chickpea to seedlings were 0.1–1% with AMV and 0.1–2% with CMV. Seed transmission rates with lentil were 0.1–5% for AMV, 0.1–1% for CMV and 0.8% for BYMV. Individual seed samples of lentil and chickpea sometimes contained both AMV and CMV. With both species, infection with AMV and CMV was sometimes found in commercial seed stocks or seed stocks from multiplication crops of advanced selections nearing release as new cultivars. Seed-borne virus infection has important practical implications, as virus sources can be re-introduced every year to chickpea and lentil crops or plots through sowing infected seed stocks leading to spread of infection by aphid vectors, losses in grain yield and further contamination of seed stocks.     

How important were stepping stones in the colonization of Krakatau?

In theory, one factor determining the rate and nature of the assembly of island biotas is the presence or absence of stepping stone islands, yet no field studies have demonstrated stepping stone function in practice. Krakatau, in Sunda Strait, is about equidistant from Java and Sumatra. Sebesi lies about half way between Krakatau and Sumatra, but no island intervenes between Krakatau and the nearest coast of Java. We assess the evidence that Sebesi has acted as an important stepping stone for Krakatau's recolonization since the devastating 1883 volcanic eruption. About a quarter of Krakatau's resident land birds, two-fifths of its reptiles, bats and land molluscs, and about two-thirds of its termites, pteridophytes, butterflies and spermatophytes are unknown on Sebesi, evidently having colonized without stepping stone involvement. Identifiable Sumatran taxa do not outnumber identifiable Javan ones on Krakatau, nor do historical distribution records indicate movement from Sebesi to Krakatau in animal groups. Krakatau's biota is not a subset of Sebesi's in predominantly anemochorous or thallassochorous plant groups, butterflies, reptiles or bats, and is only marginally so in termites. It is a subset in predominantly zoochorous spermatophyte groups, except Ficus species, and in birds and land molluscs. Comparison with a weaker stepping stone candidate, Panaitan, provides no evidence for a stepping stone role for Sebesi in butterflies or termites. We discuss the dispersal and establishment constraints on colonization by the groups involved, and conclude that, overall, Sebesi had little impact as a stepping stone. Instead, it is more probable that divergence of the environments of the two islands has led to an increasingly independent recolonization of Krakatau.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 275–317.     

Reproductive dynamics of a tropical freshwater crocodilian: Morelet's crocodile in northern Belize

Morelet's crocodile Crocodylus moreletii has not been well-studied and many aspects of its life history are unknown. In particular there is a notable paucity of information on nesting and reproductive ecology. We studied the nesting ecology of Morelet's crocodile in northern Belize from 1992 through 1995. Nesting occurs at the onset of the wet season in mid-June and continues through mid-July (mean oviposition date =1 July±10 days). Eggs hatch from mid-August through mid- to late September. Nesting effort at our primary study site remained relatively constant during 1992, 1993 and 1995, but nearly doubled in 1994; this appeared to reflect a regional trend. Natural and man-made islands are heavily used as nesting sites. Nesting success in 1993 and 1994 was consistently higher on natural islands when compared with man-made islands or shoreline sites. Nest losses were primarily due to flooding and raccoon Procyon lotor predation. Losses from predation were greatest in 1994 when unseasonably low water levels facilitated predator access to nests. Females probably reach sexual maturity in 7–8 years after attaining a total length of 150 cm. Mean clutch size (25.0±7.6; range=9–42; n =73) did not differ among years. Mean clutch size, egg width (EW), egg length, egg mass (EM) and clutch mass were positively correlated with female snout–vent length (SVL). Mean EW was the best predictor of female SVL. A partial correlation analysis of egg and clutch attributes found that independent of female SVL, EM increases with increasing clutch size.     

Zonation of forest vegetation and soils of Mount Cameroon,West Africa

Mount Cameroon is an active volcano in a wet part of West Africa. The forest vegetation and associated soils on its southern slopes were studied in 1989, 1991 and 1995 in coupled 0.25 ha plots at altitudes of 180, 600, 1,100, 1,800 and 2,180 m. All lianas and trees >10 cm dbh were enumerated and their structural features quantified. The forests were of large stature throughout. The strangling Schefflera species made a substantial contribution to the very high basal areas at 1,800 m. The associated soils were dominated by andisols derived from volcanic ash that showed a distinct increase with altitude in soil organic matter and total N attributed to lower temperatures. Soil pH, exchangeable K⁺, Ca²⁺, Mg²⁺, effective cation-exchange capacity and percentage base saturation showed very marked increases explained by the influence of recent volcanic ashfalls. Available N and P showed less distinct trends with altitude. Although there is a large decrease in tree species richness with altitude, forest stand growth (as compared on a basal area basis) does not appear to be limited by soil fertility or temperature. The forest line (altitude treeline and extensive gaps below it) appear to be controlled by periodic volcanic activity, ashfalls and lava flows, which can destroy existing forest through soil burial and fire effects and inhibit regrowth on bare lava flows and deep deposits of volcanic ash.     

Hurricane Disturbance,Plant-Animal Interactions,and the Reproductive Success of a Tropical Shrub1

Hurricane disturbance may have strong effects on plant-animal interactions important in plant reproductive success. Components of reproductive success (flowering, pollination, seed predation) in the tropical shrub Ardisia escallonioides (Myrsinaceae) were examined from 1991–1994 in four southern Florida populations. Hurricane Andrew struck three of the four populations on 24 August 1992. Hurricane Andrew delayed flowering by two months in 1992. In 1993 and 1994, the three hurricane-damaged populations had increased flowering and inflorescence production compared to 1991 and 1992, while the undamaged population had no flowering. Hurricane disturbance had different effects on generalist versus specialist plant-animal interactions. Species composition and relative abundance of the generalist pollinator community that visits A. escallonioides was similar before and after the hurricane, indicating little effect of the disturbance on this interaction. In contrast, populations of a specialist flower galling moth (Periploca sp., Cosmopterigidae) declined in 1992 following Hurricane Andrew. Although moth populations increased at two of the three sites in 1993, the relative impact of moth predation on seed production was low due to extensive flower production. One moth population suffered local population extirpation for two years, reestablishing itself in November 1994. Hurricane disturbance resulted in a window of opportunity for massive seed production of Ardisia escallonioides in south Florida. Total seed production in 1993 increased twelve to seventy-three times the 1992 levels. Total seed production declined in 1994, but remained high compared to prehurricane levels.     

Sooty shearwater (

Rakiura Maori annually harvest sooty shearwater (Puffinus griseus) chicks from islands in Foveaux Strait and adjacent to Stewart Island, New Zealand. Chick availability and the number of chicks harvested were estimated during the 1994 and 1995 seasons on Poutama (Evening Island). Burrow entrance densities estimated using circular plots were significantly higher in 1994 (0.45 ± 0.03 per m) than in 1995 (0.41 ± 0.03 per m). A similar burrow entrance density (0.45 ± 0.04 per m) was obtained in 1995 using a transect sampling technique. The number of usable burrows estimated using circular plots in 1994 and 1995 was 387 508 and 337 732 respectively. Of these, chicks occupied 24% ± 6% in 1994 and 29% ± 4% in 1995. It was estimated the muttonbirders harvested 13-24% (15 722) of the chicks present on Poutama in 1994 and 17-28% (22 092) in 1995. Muttonbirders targeted areas of the island with higher chick density and less fallen stems. Excluding chicks rejected by the muttonbirder (less than or equal to 750g), harvested chicks were significantly heavier and had less down than randomly encountered chicks. If larger heavier chicks are more likely to return and breed, then the preference for these chicks by muttonbirders would affect predictions of harvest impacts.     

Effects of experimental temperature elevation on high-arctic soil microarthropod populations

 An experiment was conducted to measure the effects of summer warming on the total population densities of soil-dwelling microarthropods in the high Arctic and to compare these results with those from natural between-year and between-site variations. Small polythene tents were used to elevate summer temperatures over 3 years on polar semi-desert and tundra heath in West Spitsbergen, Svalbard, Norway. Soil cores were taken at regular intervals from tented and untented (control) plots and heat extracted for mites (Acarina: Oribatida) and springtails (Collembola). Species present were similar at both sites, but at the start of the experiment total springtail populations were greater at the polar semi-desert whilst oribatid mite densities were equal at both sites. No significant effect of temperature elevation on oribatid mite populations emerged, even after 3 years. By contrast, springtail numbers were significantly lower on tented versus control plots at the polar semi-desert at the end of year 3, but not so at the tundra heath. Collembola numbers declined at both sites during the warm dry midsummers of 1992/1993 and this was most marked at the better drained polar semi-desert site. Over the equivalent period total oribatid mite populations, while relatively more stable, increased significantly at the polar semi-desert as a result of an increase in the number of juveniles. Results are interpreted in the context of the ecophysiological adaptations of oribatid mites and springtails to soil temperature and moisture. The resulting survival characteristics are considered in relation to the temperature and moisture characteristics of the two sites. The experiment demonstrated that year to year variation in climate, interacting with physical differences between sites, produced an equal or greater effect on microarthropod numbers at any one site than the 8–10% increase in “heat availability” (day degrees above zero) resulting from the summer tent treatment. The limitations of the use of tents to elevate soil temperatures are discussed. Comparisons are made with microarthropod population data from other polar and alpine sites. Received: 13 May 1994/Accepted: 18 May 1995     

Two decades of compositional and structural change in deciduous old-growth forests of Indiana,USA

Aims Using a network of permanent plots, we determined how multiple old-growth forests changed over an 18–19-year period at a state-wide scale. This examination of change allowed us to assess how the compositional and structural stability of each forest varied with site characteristics (topography, physiography and productivity) and stochastic disturbance.Methods In 2011, we resampled 150 plots distributed across five old-growth hardwood forests in Indiana, USA that were originally sampled in 1992–1993. Within each plot, we relocated and remeasured the diameter at breast height (dbh) of all trees (≥10.0cm) present during the 1992–1993 sample, which allowed us to track their individual fates through time for growth and mortality calculations. Trees that grew to ≥10.0cm dbh since plot establishment were designated as ingrowth. The dbh and species of all saplings (stems ≥2.0cm but <10.0cm dbh) were also recorded. For each forest, we calculated density (stems ha^-1), basal area (BA; m 2 ha^-1) and importance value (relative density + relative BA)/2) of trees by species. For saplings, density per ha was calculated by species for each forest. We also calculated annual mortality rate (AMR) for three diameter classes (10–29.9, 30–59.9 and ≥60cm) and species richness (S), evenness (E) and Shannon–Weiner diversity (H?) for the tree and sapling layers. Differences between years were compared for each forest using paired t -tests and Wilcoxon signed rank tests.Important findings Although we observed commonality in changes across some sites, our results suggest that these forests differ in their rates and trajectories of change. Changes in total stand BA and density varied across sites and were influenced by past disturbance and mortality rates. We observed a general increase in the overstory dominance of Acer saccharum coupled with a general decrease in the dominance of Quercus section Lobatae (red oak group) species. Mortality of overstory trees present in 1992–1993 ranged from 27% to 49% over the study period (mean AMR 1.6–3.7%). Most sites experienced greater mortality of early and mid-successional species, but one site experienced heavy mortality of Fagus grandifolia, a shade-tolerant late-successional species. Shade tolerant species, A. saccharum in particular, dominated the sapling layer at most sites. However, recruitment of this species into larger size classes did not occur uniformly across all sites and the species was comparatively uncommon at one site. Overall, our results suggest that old-growth remnants, even within a single state, cannot be viewed as equivalent units with regard to research or management. Stochastic disturbance events and surrounding land use may have amplified effects on small scattered remnants. Therefore, continued monitoring of these rare, but biologically important forests is critical to their long-term management and protection.