5. Ecosystems Overview

5.1 Introduction

A central goal of the Long Island Pine Barrens Protection Act is to "protect, preserve and enhance the functional integrity of the Pine Barrens ecosystem resources, including plant and animal populations and communities thereof." (E.C.L. 57-0121(2)(a)). Hence it is necessary to survey the existing ecosystem so as to tailor the plan to fulfill its legislative charge.

5.2 Description of the Ecosystem

The Long Island Central Pine Barrens region is a complex mosaic of pitch pine woodlands, pine-oak forests, coastal plain ponds, swamps, marshes, bogs and streams. Characteristic of Pine Barrens natural communities is their evolution in the presence of frequent fires. The dominant tree species in the frequently burned areas is the pitch pine, (Pinus rigida) which is highly fire adapted and somewhat fire resistant (see below). Pitch pine woodlands are characterized by widely spaced pitch pine. This spacing allows abundant sunlight to penetrate the open tree canopy allowing dense growth of the shrubby scrub oak (Quercus ilicifolia), and smaller heath species such as black huckleberry (Gaylussacia baccata), blueberry (Vaccinium pallidum and V. angustifolium), sheep laurel (Kalmia latifolia) and wintergreen (Gaultheria procumbens). In less frequently burned areas various species of tree oaks codominate, and the tree canopy is more closed. Under these more shaded conditions, scrub oak and heath shrubs decline in importance. (Reiners 1965, 1967). The herbaceous layer tends to be fairly sparse but is more developed in sunlit conditions. Characteristic species of the herbaceous layer include bracken fern (Pteridium aquilinum), and Pennsylvania sedge (Carex pensylvanica). Located in freshwater wetlands are Red maple (Acer rubrum), tupelo (Nyssa sylvatica) and Atlantic White Cedar (Chamaecyparis thyoides).

Prior to European settlement, pine barrens species probably existed on at least the coarsest, fire-prone soil types, although the actual geographic extent of the Pine Barrens at that time is unknown. Post-settlement land use practices may have expanded the area occupied by Pine Barrens vegetation. (see Evolution and History of the Pine Barrens). As recently as 100-150 years ago the Pine Barrens may have covered as much as 250,000 acres, more than one quarter of the area of Long Island. (Cryan 1980). The largest contiguous, intact Pine Barrens is the majority of the 52,500 acres designated as the core preserve. Much of the remaining Pine Barrens of the Compatible Growth Area, especially in Brookhaven, is fragmented and dissected by residential development, agricultural fields, highways, factories, golf courses, sand and gravel mines and shopping centers.

As noted in the Geological Overview, several soil associations typify the Central Pine Barrens region. Soils of the pine barrens have developed on coarse sandy and gravelly unconsolidated sediments deposited by the last two advances of Pleistocene glacial ice. The advance approximately 60,000 years ago deposited the hilly Ronkonkoma Moraine, which runs from Nassau through the Central Pine Barrens and out to Montauk Point. The final glacial advance 23,000 years ago formed the Harbor Hill moraine along the North Shore and North Fork. (Figure 4-2). During glacial retreat, sand and gravel washed from the morainal deposits formed extensive outwash plains between the two moraines and south of the Ronkonkoma Moraine to the Atlantic Ocean. This glacially deposited material is heterogenous in composition and texture. The Pine Barrens have developed primarily in areas where the deposits are coarse, and have given rise to soils in the Plymouth-Carver association. (Figure 3-1). Plymouth-Carver soils are textured, well-drained to excessively well-drained, acidic, and nutrient poor. Pine Barrens also may be found on medium to moderately coarse-textured Haven and Riverhead soils, associated with Plymouth-Carver soils in the southern and northwestern portions of the Central Pine Barrens.

The combination of droughty, nutrient-poor soils and frequent fire have created a harsh environment to which relatively few species have been able to adapt. Consequently, present plant communities of the dry uplands generally are of low diversity and low productivity. (Whittaker and Woodwell 1969). Whether the effects of past disturbance on Pine Barrens species richness contributed to present diversity is unknown. Pine Barrens natural communities are characterized by plant and animal species that are uncommon in the moist, deciduous forests that surround the Central Pine Barrens. As a result, the biota of the Central Pine Barrens tends to be unusual, and includes many rare species especially adapted to Pine Barrens conditions.

Pine Barrens freshwater wetlands are characterized by a different kind of environmental rigor. Pristine Pine Barrens water typically is acidic and nutrient-poor; water levels may fluctuate widely both seasonally and from year-to-year, especially in coastal plain ponds. Many of the plants that have adapted to these unusual conditions cannot survive elsewhere, because of their inability to compete with the more cosmopolitan, "weedy" species found in nutrient-rich habitats. In fact, coastal plain ponds (in the Central and South Fork Pine Barrens) harbor one of the highest concentrations of rare plant species in New York.

Like the plant communities, wildlife of the Central Pine Barrens is generally low in diversity. The rigorous conditions preclude use by many species which are found in the richer mesic communities elsewhere on Long Island. On the other hand, there are numerous species which are particularly adapted to this harsh environment. This is especially true of the insects.

Some species of vertebrates are virtually ubiquitous throughout the Central Pine Barrens. Mourning dove (Zenaida macroura), American crow (Corvus brachyrhynchos), white-tailed deer (Odocoileus virginianus), raccoon (Procyon lotor), red fox (Vulpes fulva), masked shrew (Sorex cinerea), eastern mole (Scalopus aquaticus) and Fowler's toads (Bufo woodhousei) are both common and widespread. The distribution and abundance of most other species, however, is largely restricted by availability of plant communities. The most typical associates of these communities are discussed below.

The invertebrate fauna of the Central Pine Barrens is not well known. "Except for Lepidoptera [moths and butterflies], however, and a few groups such as tiger beetles . . . and cerambycids . . ., the insect fauna of most northeastern pine barrens has not been intensively studied . . ." (Wheeler 1991). Numerous insects take advantage of the looser soils in that they frequently deposit their eggs in the ground. In addition, the predominant oaks are heavily exploited by a wide array of insects. The moth and butterfly fauna is especially rich; twelve species of rare Lepidopterans are known from the Central Pine Barrens. (Southampton 1993).

5.3 Ecological Processes

5.3.1 Vegetation

Pine Barrens natural communities are distributed on the landscape in a complex mosaic determined by an interaction of environmental factors and history. The key environmental factors controlling vegetation types are: (1) soil saturation (depth to water table), (2) soil texture and nutrients, (3) fire regime, and (4) human disturbance (clearing, logging). Insect herbivory and frost damage are secondary factors that also may influence vegetation composition.

Natural community types are related to key environmental factors in the conceptual model shown in Figure 5-1, which is a modification of Whittaker's interpretation of the New Jersey Pine Barrens. (Whittaker 1979).

Figure 5-1: Ecological model of the Central Pine Barrens, Long Island, New York
(Please see the printed version of the Plan for this illustration.)

Ecological model of the Central Pine Barrens, Long Island, N.Y. Adapted from R. Whittaker's (1979) depiction of the New Jersey pine barrens. Oak-pine forests are considered to be a variant of pine-oak forests by the New York Heritage program. Scrub oak areas are variants of pitch pine-oak heath woodland. Although not generally thought of as a pine barrens vegetation type, oak forests are present in the Pine Barrens and represent one end of the upland vegetation continuum.

On the chart, the horizontal axis represents depth to the water table, which is critical in separating uplands from wetlands. There may be as little as a half meter difference in elevation between a swamp and a dry pitch pine woodland. (Zampella et al., 1992). The topographic interface between open water and uplands may be occupied by pond shores, shrub swamps, poor fens, cedar swamps and/or red maple swamps, depending upon site topography, hydrology, and history. It is difficult to accurately represent the complexity of interrelationships among wetland types in the conceptual model, partly due to the limitations of this generalized, simplified model, and in part due to the lack of detailed ecological information.

Upland forest vegetation types are arrayed along the vertical axis of the model (Figure 5-1), which represents the combined gradients of soil texture and ecological effects of fire. Fire history terminology can be confusing due to inconsistencies in usage. (Romme 1980). For this discussion, the term "severity of ecological effects" refers to a combination of attributes of individual fires (fire intensity, fire severity), and the overall fire regime of the site (fire frequency and mean fire return interval). Fire intensity has most commonly been used to refer to fire temperature (units of energy released/area). Fire severity refers to ecological effects such as consumption of forest floor organic matter and mortality of overstory trees. (Romme 1980). Fire frequency refers to the number of fires per unit time in a designated area. Mean fire return interval refers to the mean number of years between two successive fires in a designated area of specified size. (Romme 1980).

At the extreme upper end of the fire gradient in the conceptual model (Figure 5-1), the fire return interval is the shortest (i.e., fires are the most frequent), and the fires tend to be the most severe (e.g., stand-replacing crown fires that consume forest floor litter and duff). On the lower end of the gradient, the fire return interval becomes longer and the fires tend to be less severe. Fires tend to be the most frequent and the most severe on the coarsest soils, which tend to be vegetated by drought-tolerant, flammable vegetation. (see below).

Natural community types typical of the most drought-prone soils with the severest fire regime are the dwarf pine plains and pitch pine-oak-heath woodlands. These communities grade into the pine-oak forests, which are maintained by a less severe fire regime. (Figure 5-1). Tree oaks become more common, and pitch pine less common, as fire severity and frequency decrease. Pine-oak forests in which oaks predominate are referred to as oak-pine forests in Figure 5-1. Oak-pine forests are included within the pine-oak type classification by the New York Natural Heritage Program, but are separated in this discussion because of their decreased flammability and lower fire frequency. At the end-point of the fire frequency gradient, where fires are the least frequent and least severe, are oak forests. Although not a Pine Barrens vegetation type, oak forests are included in the model because they represent one end of the upland vegetation continuum. Small areas of oak forest with few or no pitch pines do occur within the Central Pine Barrens.

Pine Barrens vegetation types throughout the northeast are located in regions with humid climates and ample rainfall, and are surrounded by mesic hardwood forests. Despite these factors, the unusual Pine Barrens plants and animals manage to persist. How do the Pine Barrens resist invasion by deciduous forest species? It appears that as result of the reinforcing interactions of droughty, nutrient-poor soils with highly flammable, fire-adapted vegetation and frequent wildfire, an environment is created hostile to the mesic hardwood forest while remaining conducive to the Pine Barrens vegetation.

Pine Barrens soils typically are 80-96% sand and drain very rapidly. Only vegetation that can withstand these droughty soils, and the soil's low nutrient levels and acidity, can persist. Many Pine Barrens plants produce waxes, resins, or volatile oils which help leaves retain moisture, (Burg 1983) and which may also reduce insect herbivory. (Patterson, personal communication). The very existence of these waxy compounds which allows the vegetation to exist on the Pine Barrens soil also increases the potential for fire. The compounds are highly flammable. Furthermore they contain volatile oils which vaporize when heated. The vapors ignite at relatively low temperatures and greatly increase the likelihood that fires will reach the tree crowns. (Patterson, personal communication). The oils and resins also increase the amount of heat emitted during a fire. (Patterson, personal communication). Additional characteristics that favor fire include litter that decomposes slowly and thus tends to accumulate on the soil surface, litter of low water-absorbing capacity, litter of low mineral content, high plant surface-to-volume ratio, high dead-to-live plant tissue ratio, and "ladder fuels" that carry flames upward from the ground. (Rundel 1981, Latham and Johnson 1993).

Mutch hypothesized that "fire-dependent plant communities burn more readily than non-fire-dependent communities because natural selection has favored development of characteristics that make them more flammable." (Mutch 1970). Positive feedback between flammability and fire-dependence would favor the persistence of fire-dependent communities, and inhibit invasion by species of nearby alternative communities. Latham and Johnson have postulated that fire-facilitating species, including those of the Pine Barrens, are also tolerant of nutrient scarcity. (Latham 1993). Fire-facilitators thus may gain a competitive advantage from fire due to long-term decreases in nitrogen availability, as well as to outright elimination of fire-sensitive competitors. Fire-facilitating species produce biomass and litter that are highly flammable, thereby increasing the likelihood and severity of wildfire. Frequent severe fires decrease nutrient levels, further favoring the fire-facilitators. This Pine Barrens-stabilizing feedback loop may be destabilized by suppression of fire, addition of nutrients, or prolonged wet weather. Under these conditions fire frequency decreases, nutrient-demanding, fire-intolerant plant species increase, and Pine Barrens species decline. With prolonged fire suppression, vegetational succession leads to the replacement of Pine Barrens by oak forests. (Figure 5-1).

Many Pine Barrens plant species exhibit adaptations to fire, including pitch pine (Pinus rigida) and scrub oak (Quercus ilicifolia). McCune has classified North American pines into five groups, including a "fire-resistant" group and a "fire-resilient" group. (McCune 1988). Fire-resistant pines are tall, have thick bark, long needles, large seeds and are slow to initiate seed production. Fire-resilient pines have low-to-moderate fire tolerance as mature trees, but produce cones at a young age, produce abundant small, readily dispersed seeds, and have a high degree of cone serotiny. Serotinous cones are cones that open only after being heated to high temperatures, such as occur during a fire. Reproductive behavior of fire-resilient pines is typical of "r-selected" pioneer species that survive as seeds through infrequent catastrophic fires, and have high potential for explosive reproduction. (McCune 1988). McCune classifies pitch pine in the fire-resilient group, but it also possesses some attributes of the fire-resistant group, namely its relatively long needles and thick bark, which increase its fire-resistance. However, pitch pine also possesses the fire-resilient attributes of precocious cone production and production of abundant small seeds. Pitch pine is usually not serotinous, but in areas with an unusually high fire frequency (short fire return interval) the serotinous trait is favored. (Ledig and Little 1979, Givnish 1981). For example, pitch pine in both the New Jersey and Long Island dwarf pine plains have a high frequency of cone serotiny.

Pitch pine shares with three other species in McCune's fire-resilient group (P. leiophylla, serotina and virginiana) the capacity to resprout vegetatively from dormant epicormic buds, located both beneath the trunk's bark and on the root collar. (Stone and Stone 1954). In the frequently burned dwarf pine plains, vegetative pitch pine sprouts may bear cones as young as 3 years. (Ledig and Little 1979). The ability to resprout ". . . may provide a mechanism for improved survival of fire-susceptible species on sites subject to fire-free intervals shorter than required for plentiful seed reproduction." (McCune 1988). The ability to resprout declines with age in tall pitch pines. (Andresen 1959). However, the effect of age on resprouting of dwarf pines in the Long Island pine plains is not known. Windisch suggests that dwarf pines in New Jersey do lose their ability to resprout with age. (Windisch 1990). Although vegetative reproduction often predominates, production of seedlings is important for replacement of senescent trees. Pitch pine seedlings survive and grow best under the conditions of full sunlight, exposed mineral soil, and reduced competition, conditions that usually follow severe fires.

Scrub oak also is fire adapted, and rapidly recovers from even a hot crown fire. The plants have large root collars, just below the soil surface, which bear numerous dormant buds that resprout readily when above-ground branches are killed. (Unnasch 1990). New shoots grow rapidly, often setting fruit after three years, and reaching maximum size in 7-10 years. (Wolgast and Stout 1977). Acorn production reaches a maximum when the sprouts are 5-7 years of age, and slowly declines thereafter. (Wolgast 1973). Scrub oak seedlings can become established only during the first few years following fire, due to decreased predation by white-footed mice. (Unnasch 1990). In addition to rejuvenating acorn production, fire has been found to stimulate a 4 to 9 fold increase in scrub oak foliage and shoot production during at least the first four years following fire in a scrub oak habitat in Pennsylvania. (Hallisey and Wood 1976). Concentrations in the foliage of crude protein, phosporus (P), potassium (K), calcium (Ca) and magnesium (Mg) also increased following fire. (Hallisey and Wood 1976). Fire-stimulated increases in forage quantity and quality may be important for maintaining populations of lepidopteran species (primarily moths) that feed primarily on scrub oak.

Ericaceous shrubs including blueberries, huckleberry and wintergreen, and herbs such as bracken fern, also quickly resprout and regain former biomass and production levels following fire. (Hallisey and Wood 1976). Additionally, increased protein levels have been measured in post fire resprouts of the blueberry. (Hallisey and Wood 1976). Periodic fire is required to open the canopy and provide the light levels required by herbaceous species typical of grassy openings in the pine barrens.

Development of upland Pine Barrens community types is controlled by an interaction between the fire regime and the soil texture and fertility. Both fire and soil characteristics are used as the vertical axis in the conceptual model (Figure 5-1), based on the assumption that coarse, droughty soils should tend to be more fire-prone. Therefore, natural communities associated with the shortest average fire return interval also would tend to be correlated with the coarsest soils, as was found by Olsvig. (Olsvig 1979). She reported an increase in the percentage of coarse sand, a decrease in the percentage of silt and clay, and a decrease in nutrients (P, K, Ca, Mg) going from oak-pine forest, to pine-oak forest, to (transitional) pitch pine oak heath woodland, to dwarf pine plains and to successional heath areas. However, similar data gathered for Long Island by Seischab and Bernard in 1993 (unpublished), unlike the Olsvig research, failed to reveal a clear gradient in soil texture across the same range of community types. Thus it appears that although pine barrens vegetation in general occurs on coarse soils, the same soil type may support different pine barrens community types, both on Long Island and in New Jersey.

Although the presence of well-drained, droughty soils are a prerequisite for development of flammable, Pine Barrens community types, (Patterson, personal communication), it appears that differences in fire regime may to at least some extent override soil characteristics in driving genetic selection and shaping plant community types in specific locations. Differences in spatial scale, position on the landscape, and potential for human-set fires may also be involved. For example, vegetation on small patches of droughty soil surrounded by mesic soils and vegetation, small areas isolated by firebreaks (natural or human-created) from their surroundings, and areas distant from human ignition sources would be relatively unlikely to burn. In contrast, fire would be much more likely in large areas uninterrupted by firebreaks, and which therefore could be burned by fires originating in many different locations within the area. This is known as the "fireshed" concept. (Windisch 1993, unpublished). The presence of humans would increase the likelihood of ignitions, since fires today primarily originate from arson. Pre-settlement fire regimes would have been strongly influenced by locations of Native American populations.

The dwarf pine plains are thought to have historically had the shortest fire return interval, perhaps as frequently as every 10-30 years (Windisch 1992) or even as often as 6 years. (Cryan 1982). Pitch pine-oak-heath woodland communities may have burned an average of every 20-35 years (Windisch unpublished). The most extensive occurrences of this community are as transition zones between the dwarf pine plains and surrounding pine-oak forests. The mean fire return interval in pine-oak forests may have ranged from 40-60 years in pine-dominated areas to 50-100 years in oak-dominated areas. (Windisch 1992; Cryan 1980). Oak forests dominate natural areas with fire return intervals greater than 100 years. These estimates of fire return intervals are imprecise, and need further verification and correlation with vegetation types (see Status of Research in the Central Pine Barrens).

Fire severity may be just as important as the fire return interval in shaping Pine Barrens vegetation. Anecdotal evidence suggests that severe, stand-replacing fires that consume most or all of the soil organic matter may have contributed to the creation of the dwarf pine plains and pitch pine oak heath woodland communities. A model more detailed than that shown in Figure 5-1 should be developed, including all aspects of the fire regime. Such a model cannot be created yet due to insufficient information. Such a model could be prepared after completion of a fire history and historical land use study for the Pine Barrens.

Superimposed on the natural environmental gradients are human-caused disturbances such as land clearing, sand and gravel mining, draining of swamps, and logging. Pitch pine can readily reseed cleared and abandoned areas. Pine Barrens vegetation ultimately reclaims disturbed areas through the process of natural succession. (Figure 5-1). However, succession proceeds very slowly in the dwarf pine plains, where large areas of heath have persisted for many years after sites were cleared. (Olsvig 1979; Windisch 1990). Succession may require 50 or more years to reclaim cleared areas in the pine plains with tall pitch pines invading in some locations. (Windisch 1990). When these areas were cleared much or all of the topsoil probably was removed. Since most soil nutrients are located in the surface organic horizons, vegetation regrowth may be limited by a lack of nutrients. The addition of nutrient-rich organic matter in the form of compost has been effective in restoring disturbed pine plains in New Jersey. (Fimbel and Kuser 1993). A possible explanation for the slow revegetation could be due to a scarcity of pine seed. This scarcity is a result of the fact that many of the surrounding dwarf pines have serotinous cones that would not open until heated by fire.

Fire also may be a significant environmental factor for Pine Barrens wetlands which are likely to have been burned over during periods of drought in the past. Coastal Plain Atlantic white cedar swamps depend on fire, or other disturbance to create the sunlit conditions necessary for seed germination and seedling survival. (Laderman 1989). Without fire, white cedar may eventually be replaced by shade-tolerant red maple. The role of fire in the ecology of coastal plain ponds is unknown. Occasional fire may be necessary to eliminate shrub and tree species from coastal plain ponds, and to reduce organic matter. (Schneider and Zaremba 1991).

Scrub oak is often damaged by late spring frosts, and may be eliminated in frost-prone, low-lying areas such as the drainage swales in the southern and western portions of the pine plains, which are vegetated primarily with short heath shrubs. Insect herbivory may be important throughout the Pine Barrens. Pitch pines have been killed by periodic outbreaks of the pine looper, which may modify forest structure and composition.

5.3.2 Wildlife

The key environmental factors which control the distribution and abundance of wildlife are:

1. vegetative cover-type,
2. soil moisture and free water;
3. size and shape of habitat patches;
4. juxtaposition of habitat patches to other habitat types;
5. range expansions, introductions and extirpations;
6. levels of human disturbance; and
7. for many migratory organisms, the conditions which exist in other portions of their range. Vegetative Cover Type

Of these factors, the vegetative cover-type, or plant community, is the most important. Differences in plant parameters such as species composition, size, density, age, condition and structural height diversity can determine a community's suitability as habitat for a given species of wildlife. As a general rule, vertebrates tend not to depend upon single plant species while invertebrates are frequently dependant upon individual species for their survival. Soil Moisture Content

As mentioned above, the level of soil moisture bears an obvious relationship to the plant community: diversity and abundance generally increase as soil moisture increases. Invertebrate fauna shares this relationship with soil moisture as well. Vertebrate wildlife frequently depend upon invertebrates as a food source and, thus, their abundance and diversity tend to be greater in moister soils. When free water is available for an extended period of time, aquatic communities of both invertebrates and vertebrates can exist. Habitat Availability

The size, shape and juxtaposition of habitat patches are key determinants of wildlife distribution. When large blocks of habitat are divided, reduced in size, or separated from other blocks, habitat fragmentation occurs. This phenomenon frequently results from human alteration of the landscape and poses a significant threat to biodiversity, both regionally and globally.

The theory of island biogeography suggests that the size of a patch or "island" of habitat is directly related to the diversity of the wildlife it will support. In essence, bigger patches of habitat support a greater diversity of wildlife than smaller patches.

Size is also important because some species of wildlife will not use a patch of otherwise suitable habitat if it is too small. Such "area sensitive" species require at least a threshold-sized patch for their territories or home ranges. As an example, some species of wildlife are primarily adapted to finding their life needs in an interior niche of a forest. As consequence, they require large blocks of forest habitat in which an acceptable interior niche can be located and in which they can avoid edges.

The shape of habitat patches also affects the wildlife species composition. "Edge" occurs where two different habitat types meet. A given acreage of habitat type has the least edge if it is circular; linear habitat patches have the greatest amount of edge. This is a purely geometric relationship. Some species of wildlife, the generalists and opportunists, are adapted to edge environments while others, the specialists, suffer from the existence of edge, due to increased competition, predation or parasitism.

The physical relationship of a patch of habitat to other patches is important in determining wildlife distribution. Some species of wildlife require more than a single type of habitat. As an example, an animal may use terrestrial habitats during the non-breeding season but may require a wetland for a breeding site. Thus, the existence of an otherwise suitable habitats cannot support some species if a breeding site is not available nearby.

Biogeographic theory also suggests that the distance between an island and the nearest similar island is important. A patch nearer a patch of the same type will support more wildlife species than a same-sized patch which is further away. In other words, closer islands support a greater diversity of wildlife because opportunities for immigration are greater.

Travel amongst habitat patches is also facilitated by travel corridors. Corridors are simply linear (approximately) patches of habitat which provide adequate cover for travel but do not necessarily provide food or breeding sites. Conversely, barriers to wildlife movements must be considered in determining whether a suitable habitat is created. Natural or artificial barriers may preclude wildlife use of otherwise viable habitats.

An animal's range can shrink or expand due to natural or human causes. In many cases, the causes are only poorly understood, especially when they appear to be naturally induced. Changes in bird distribution are well recorded. Some human activities have a direct and virtually permanent effect on wildlife distribution. As an example, extirpations of nonmigratory species can preclude their use of otherwise suitable habitats. Humans also frequently introduce nonindigenous species of wildlife, domestic animals, disease vectors and plants. The effects of the introduced species can be beneficial to individual species but may, in some cases, be deleterious to indigenous communities. In such situations, the effects can result in increased competition, predation, disease and habitat degradation. On the other hand, populations can also be restored by humans.

Human disturbance itself can also restrict the use of available habitats. Some species are intolerant of the sounds, sights and even smells which are characteristic of developed landscapes. The species may purposefully abandon or avoid such areas. This phenomenon is generally observed by vertebrates. Invertebrates are especially affected by the use of pesticides and artificial lights.

For migratory species, such as many birds, most bats, and some fishes and insects, conditions in other parts of their ranges may restrict their total population sizes and/or migratory routes, and thus limit their abilities to exploit available habitats.

5.4 Description of Ecological Communities

Ecological communities are classified and defined in this plan according to the 1990 Reschke scale. (N.Y. Natural Heritage Program and N.Y.S. Department of Environmental Conservation). Reschke defines a community as ". . . a variable assemblage of interacting plant and animal populations that share a natural environment." A biological community, or complex of interrelated communities, together with its associated non-living environment constitutes an ecosystem. (Primack 1993).

Significant communities and rare species are referred to as "elements" by the New York Natural Heritage Program. These elements have been assigned ranks reflecting their rarity. The global rank reflects the rarity of the element throughout the world, and the state rank reflects the rarity within New York State. The definitions in Figure 5-2 are from the New York Natural Heritage Program.

Communities are dynamic entities that change with time, intergrade with each other spatially and temporally, to form a complex mosaic in the landscape. Individual community types may include a considerable range of variability. Ecological communities may occur naturally or be human-created. The community boundaries indicated in Figure 5-3 are necessarily artificial lines drawn across ecological gradients and represent a simplification of the complex, real landscape.

Figure 5-2: Global and State Ranking for Elements of Rare Communities and Species
Figure 5-3: Pine Barrens Ecological Communities5.5 Natural Pine Barrens communities
(Please see the printed version of the Plan for these illustrations.)

Natural communities present within the Central Pine Barrens that are currently recognized by the Natural Heritage Program are listed in Figure 5-4. Salt marsh communities are not Pine Barrens community types, but do occur within the Flanders area of the Central Pine Barrens. In addition to the communities listed in Figure 5-4, two additional communities have been recognized in this document: Pine Barrens vernal pools and wet Pine Barrens. Community definitions that follow are based in large part on those of Reschke. (Reschke 1990).
Figure 5-4: Natural Pine Barrens Communities in Descending Order of State Rarity
Dwarf pine plains G1G2 S1
Coastal plain Atlantic white cedar swamp G3G4 S1
Coastal plain stream G3G4 S1
Coastal plain poor fen G3? S1
Coastal plain pond G3G4 S2
Coastal plain pond shore G3G4 S2
Pitch pine-oak-heath woodland G3G4 S2S3
Salt panne G3G4 S3
Pine barrens shrub swamp G5 S3
High salt marsh G4 S3S4
Low salt marsh G4 S3S4
Chestnut oak forest G3G4 S4
Pitch pine-oak forest G4G5 S4
Salt shrub G5 S4
Red maple-hardwood swamp G5 S4S5

5.6 Upland Communities

5.6.1 Pitch Pine-oak forest G4G5 S4

The dominant trees are pitch pine mixed with one or more of the following oaks: scarlet oak (Quercus coccinea), white oak (Q. alba), black oak (Q. velutina) or red oak (Q. rubra). The relative proportions of pines and oaks are quite variable. The shrub layer consists of scattered clumps of scrub oak (Q. ilicifolia) and a nearly continuous cover of low heath shrubs such as huckleberry (Gaylussacia baccata) and blueberries (Vaccinium pallidum, V. angustifolium). Herbs such as: Bracken fern (Pteridium aquilinum), wintergreen (Gaultheria procumbens) and Pennsylvania sedge (Carex pensylvanica) are sparse. Scrub oak coverages tend to be highest in the pine-dominated stands and lowest in oak-dominated stands. (Reiners 1967). Heath shrub abundance decreases with increasing cover by scrub oaks and tree oaks. Forests dominated by oaks have been considered to be a separate forest type (oak-pine forest) by McCormick and Jones (1973), Olsvig et al. (1979), Whittaker (1979), and Windisch (1992). However, the New York Natural Heritage Program does not make this distinction, since there is no evidence that plant species composition differs in pine-dominated versus oak-dominated stands (Reschke, personal communication). Division of this community into pine-oak and oak-pine types makes sense for management purposes, however, since a high proportion of pitch pine indicates higher flammability and presumably a more recent fire incidence.

The wildlife community can vary dramatically as this dry forest type varies from pine-dominated to oak-dominated. Nevertheless, some animals are found throughout these communities. Common birds are rufous-sided towhee (Pipilo erythrophthalmus), bluejay (Cyanocitta cristata), red-tailed hawk (Buteo jamaicensis) and bobwhite (Colinus virginianus). Eastern cottontail rabbit (Sylvilagus floridanus) and eastern chipmunk (Tamias striatus) are common mammals. Three snakes, the hognose (Heterodon platyrhinos), black racer (Coluber constrictor) and garter snake (Thamnophis sirtalis), are typical reptiles. Except when surface water is available nearby, the Fowlers' toad (Bufo woodhousei) is the only common amphibian.

Numerous other birds use Central Pine Barrens uplands during migrations. Dozens of species of flycatchers, warblers, thrushes, vireos and other neotropical insectivores forage in woodlands during their northward and southward travels. The richer woodlands in stream corridors are especially valuable.

When pitch pine is the principal canopy species, pine warblers (Dendroica pinus) are found in the taller stands. In the more open stands, the prairie warbler (Dendroica discolor), common yellowthroat (Geothlypis trichas), field sparrow (Spizella pusilla), brown thrasher (Toxostoma rufum) and brown-headed cowbird (Molothrus ater) are prevalent. Blue-winged warblers (Vermivora pinus) and indigo buntings (Passerina cyanea) are found at habitat edges.

When oaks dominate the canopy, red-eyed vireos (Vireo olivaceus), northern orioles (Icterus galbula) and scarlet tanagers (Piranga olivacea) and are found in the tree-tops during the breeding season. They are joined by raptors, crows (Corvus spp.), grey (Sciurus carolinensis) and flying squirrels (Glaucomys volans) and grey tree frogs (Hyla versicolor). The vegetation in the forest forms distinct levels or stories. These create a host of different niches. The middle level hosts the hole-nesting birds and animals such as great crested flycatcher (Myiarchus crinitus) and eastern wood pewee (Contopus virens), hairy woodpecker (Picoides villosus) and raccoon. The lower shrub story accommodates the grey catbird (Dumetella carolinensis), ovenbird (Seiurus aurocapillus), wood thrush (Hylocichla mustellina) and others which nest on or near the ground. On or just below the ground surface are the ground dwellers: ruffed grouse (Bonasa umbellus), rufous-sided towhee, white-footed mouse (Peromyscus leusopus), box turtle (Terrapene caroliniana) and red-backed salamander (Plethodon cinereus). The larger blocks of forest support nesting great horned owls (Bubo virginianus) and whip-poor-wills (Caprimulgus vociferus). Both of these species forage in more open areas but nest in woods. The owl nests in trees while the goatsucker nests on the ground.

5.6.2 Chestnut oak forest G3G4 S4

The chestnut oak forest community description does not precisely describe the hardwood forests found in the Pine Barrens, but is the best fit among the forested uplands community types recognized by the Heritage Program. Chestnut oak forests are dominated by chestnut oak (Quercus montana) and red oak. Common associates are white oak, black oak and red maple. American chestnut (Castanea dentata) was common in these forests prior to the chestnut blight. Although chestnut and red oak do occur in the Pine Barrens, they are uncommon. The best description of the hardwood forests in the Pine Barrens is the description of the pitch pine-oak forest but with pitch pines absent, namely a forest dominated by a variable mix of scarlet oak, white oak and black oak. Chestnut may be common in localized areas, such as the steeply sloping morainal areas south of Route 111 in Manorville. Characteristic shrubs are black huckleberry, mountain laurel and blueberry. Common groundlayer plants are Pennsylvania sedge and wintergreen. Wildlife is similar to that found in the oak-dominated pine-oak forests.

5.6.3 Dwarf pine plains G1G2 S1

This low-diversity woodland community is dominated by dwarf individuals of pitch pine, and scrub oak. The canopy is generally from 1 to 3 meters tall, and may form a dense thicket. Black huckleberry and blueberry (V. pallidum) form a low shrub canopy under the pines and scrub oak. The only other common vascular species are hudsonia (H. ericoides), bearberry (Arctostaphylos uva-ursi) and wintergreen. The groundcover includes an especially diverse flora of foliose and fruticose lichens. Community variants occurring within the dwarf pine plains include areas of heath shrubs and scrub oak thickets in which pitch pine is rare or absent. As previously noted, disturbance, frost or an unusually short fire return interval may be involved in creating these variant communities.

Many hypotheses have been suggested to account for the diminutive stature of pitch pines in the dwarf pine plains, including location relative to prevailing winds and ignition sources, high fire frequency, low nutrient soils, aluminum toxicity, physical response to repeated burning, drought, and genetic selection. Aluminum toxicity has been ruled out by the studies of Andresen. (Andresen 1959). Therefore, the actual "cause" may be an interaction among the remaining factors. It seems reasonable that the primary factors shaping the Long Island pine plains would be similar to those that shaped the New Jersey pine plains; namely, a higher than average fire frequency with resultant genetic selection for rapidly resprouting dwarf trees that bear serotinous cones at a young age. (Givnish 1981). Garden experiments with dwarfed pitch pines in New Jersey have indicated a genetic basis for dwarf stature, shrubby growth form and precocious cone production. (Good and Good 1975; Givnish 1981). High fire frequency appears to be the selective agent favoring serotiny in the New Jersey plains. (Givnish 1981).

Similar garden experiments have not been performed with Long Island's dwarfed pitch pines, but it is likely that genetics are important in the Long Island plains as well. Analysis of the DNA of dwarf and tall pines is now being done, with results anticipated very soon. (Colosi, personal communication). Genetic selection is a process that may take hundreds of years (Colosi, personal communication), especially in the case of dwarf pines which reproduce following fire primarily by vegetative sprouting. (Windisch, personal communication). If indeed the Long Island dwarf pines are a genetic ecotype distinct from the surrounding taller pines, it is likely that the Long Island pine plains evolved in response to elevated fire frequencies prior to European settlement. The cause of that elevated fire frequency is unclear. The Long Island dwarf pine plains are located on a large, coarse glacial outwash fan that has given rise to some of the most rapidly draining, drought-prone soils in the Pine Barrens. There are relatively few wetlands within the dwarf pine plains area that could serve as natural firebreaks, so the pine plains may represent a large "fireshed" superimposed upon coarse, droughty soils and flammable vegetation. The combination could have given rise to a high incidence of fire both pre- and post-European settlement. Past patterns of Native American habitation also could have played a role in the development of the pine plains.

Fire has been infrequent in the pine plains for the last several decades. The most recent extensive fires were in the 1930's and 1940's, with large areas probably having been unburned in the prior decades. (Windisch 1994). The prolonged absence of fire may be resulting in the death of the oldest dwarf pine sprout clumps, and their gradual replacement by non-dwarf pines, particularly at the perimeter of the pine plains. (Windisch 1990, Cryan 1982). Windisch speculates that a stem-killing fire following a long fire-free period could result in the death of most or all of the senescent dwarf pine sprout clumps. (Windisch 1990). Dwarf pines cored and aged by Seischab and Bernard in 1993 (unpublished) were only 15-20 years old, but many of these trees appear to have been located on sites of former clearings, revegetated in recent decades. At this time we do not have enough information on dwarf pine ages and rates of senescence to assess the effects of fire suppression on the vegetation of the dwarf pine plains.

The wildlife community of the dwarf pine plains is dominated by a few songbirds: prairie warblers, field sparrows, brown thrashers and towhees. The northern harrier (Circus cyaneus), which is listed by the NYSDEC as threatened also nests here. Amphibians are virtually absent. An especially diverse assemblage of lepidopterans (moths and butterflies) is found in the dwarf pine plains, including several rare species. The dwarf pine plains support the largest, densest population of the coastal barrens buckmoth (Hemileuca maia), which thrives in the dense scrub oaks. These moths pupate beneath the soil's surface, safe from the typical autumn fires, and emerge in mid-October as adults. They have the capacity to pupate until the following fall if conditions for emergence are unfavorable. Buck moth larvae depend on the abundance of fresh, nutrient-rich young scrub oak sprouts that are produced after a fire.

5.6.4 Pitch pine-oak-heath woodland G3G4 S2S3

"A Pine Barrens community that occurs on well-drained, infertile sandy soils in eastern Long Island . . . The structure of this community is intermediate between a shrub-savanna and a woodland." (Reschke 1990). Widely spaced pitch pine and white oak form an open tree canopy with 30-60% cover. A few black and scarlet oaks also may occur. Typically there is a fairly dense shrub layer dominated by scrub oaks (Q. ilicifolia and Q. prinoides), with some heath shrubs such as huckleberry and blueberry.

This community resembles the pitch pine-scrub oak barrens ("Oak Bush Plains") vegetation type found in western Suffolk County. However, these community types may be distinguished based on the presence of tree oaks scattered throughout the pitch pine-oak-heath woodland. Tree oaks are rare or absent in the pitch pine-scrub oak barrens. (Reschke, personal communication). Also, openings in the pine and scrub oak canopy in the pitch pine-oak-heath woodland have heath shrubs such as huckleberry, blueberry and bearberry, whereas canopy openings in the pitch pine-scrub oak barrens are predominantly grassy or herbaceous, with high herb diversity. (Reschke personal communication, Reschke 1990). In a few localized areas heath shrubs predominate, scrub oaks are absent and pitch pines are few and scattered. In some locations these "heath variants" of the community type are a successional stage following clearing, or following destruction resulting from bombing practice during the 1940's. In low-lying areas frost damage due to cold air drainage may eliminate scrub oak and pitch pine, and result in a dominance of heath shrubs.

Pitch pine oak heath woodland occurs most abundantly in the transition zone between the dwarf pine plains and the surrounding pitch pine-oak forest, as well as in several other locations in the pine barrens. (Figure 5-3). The extent of this community was mapped in western Southampton by Eric Lamont, Martin Shea and others, and is referred to as "pitch pine-scrub oak barrens" in the Southampton Town Western GEIS. (Southampton 1993).

The fire regime needed to create and maintain this community is not rigorously documented. Reschke suggests that the historic fire interval probably was more than 15 years. (Reschke 1990). Windisch estimates a fire interval of 10-30 years. (Windisch 1992). In order for tree oaks sprouts to attain sufficient size to set acorns following a crown fire, a minimum fire interval of 15-20 years probably would be necessary. Some pitch pine-oak-heath woodland occurrences in the western Pine Barrens are in areas known to have had severe fires in recent decades. (H. Davis, personal communication).

Fire affects the fauna here in several ways. The prevalent dead, standing trees which result from frequent fires offer an abundance of cavities for hole-nesting birds. Black-capped chickadees (Parus atricapillus) and great crested flycatchers will use such trees in the more densely-wooded sections. Eastern bluebirds (Sialia sialis) and tree swallows (Tachycineta bicolor) will use scattered trees in more open sites. Fires which reduce the shrub layer provide suitable foraging conditions for the bluebird which frequently pursues its insect prey on the ground. Numerous small mammals, snakes, insects and other invertebrates avoid fire in underground burrows.

5.7 Wetland Communities

Over 4,300 acres of NYSDEC regulated freshwater wetlands are found in the Central Pine Barrens. The majority of this acreage occurs within the two principal river systems. The Peconic River encompasses about 2,000 acres and the Carmans River encompasses about 1,000 acres. One-hundred-sixty-two other wetlands comprise the balance of the acreage. There are 7 wetlands between 15 and 100 acres in size. The reminder are smaller than 15 acres each.

There are many different kinds of wetlands within the Central Pine Barrens. By far, the most common type is a hardwood swamp dominated by red maple (Acer rubrum). This type is found throughout the Central Pine Barrens and frequently serves as a border between uplands and other wetland types. It is found where soils are saturated or inundated for brief periods during the growing season. As soils become saturated more frequently or for longer periods, other wetland community types appear. The generally low levels of nutrients and the high acidity result in harsh environments in which only specially-adapted organisms can thrive. Coastal plain ponds and pond shores are found where water levels fluctuate greatly.

Numerous wildlife require the free water provided by the ponds, swamps, bogs, streamcourses and saltmarshes of the Central Pine Barrens. Mallards (Anas platyrhynchos), black ducks (A. rubripes) and wood ducks (Aix sponsa) are the most prevalent waterfowl. They breed and feed in Pine Barrens surface waters. Belted kingfishers (Ceryle alcyon), little green-backed (Butorides striatus) and great blue herons (Ardea herodias), ribbon (Thamnophis sauritus) and northern water snakes (Natrix sipedon) hunt here, as do snapping turtles (Chelone serpentina). Tree and barn swallows (Hirundo rustica) "hawk" flying insects on the wing.

The NYSDEC listed endangered tiger salamander (Ambystoma tigrinum) frequently breeds in Central Pine Barrens ponds when they are free of predatory fish. They are joined by grey treefrogs and Fowler's toads as well as spring peepers (Hyla crucifer) and wood (Rana palustris), green (R. clamitans) and bull frogs (R. catesbiana). In dry years, lesser yellowlegs (Tringa flavipes) and other sandpipers such as pectoral (Calidris melanotus), solitary (Tringa solitaria) and spotted (Actitis macularia) feed on exposed mudflats during their migrations. The spotted sandpiper also stays here to breed.

Many of the permanent surface water bodies of the Central Pine Barrens support fish. Fish diversity is generally low, especially in more acidic waters. Ponds support those species which favor warm, shallow weedy areas. Yellow perch (Perca flavescens), white perch (Morone americana), carp (Cyprinus carpio), goldfish (Carassius auratus) and sunfishes (Lepomis spp.) are common. Largemouth bass (Micropterus salmoides) and rock bass (Ambioplites rupestris) prefer the cleaner waters, as do golden shiner (Notemigonus crysoleucas) and chain pickerel (Esox niger).

As for moths, the Southampton WGEIS noted

. . . bogs and swamps support a number of highly specialized borer moths, including several rare species. These noctuid moths...have evolved strictly to live in conditions of extreme wetness, acidity, and nutrient poor soils. . . . Certain borer moths, while rare regionally, can be extremely abundant seasonally in localized areas of the Central Pine Barrens. . . . (Southampton 1993).

5.7.1 Coastal plain ponds G3G4 S2; Coastal plain pond shores G3G4 S2

Coastal plain ponds on Long Island are located on glacial moraines and outwash plains, with fine to coarse sandy substrates. The ponds are small (most < 2 ha) and shallow with gently sloping pond shores. (Zaremba and Lamont 1993). Coastal plain ponds are identified based primarily upon their plant species composition rather than geologic origin. The elevation of nearly all coastal plain ponds is essentially the same as the elevation of the regional groundwater table, indicating that the ponds are in direct contact with groundwater. Overton Road pond is 50-60 feet higher than the local groundwater table, indicating that it is "perched," presumably on localized layers of relatively impermeable substrate. (Zaremba unpublished). Currans Road South and Randall Road North ponds are about 10 feet higher than groundwater, and thus may be perched. (Zaremba unpublished).

Coastal plain ponds are commonly isolated, with no inlets or outlets, and depend on direct precipitation and groundwater inputs. However, many of the Central Pine Barrens ponds in the Peconic River headwaters are interconnected by surface water flow. Regardless of their water sources, pristine coastal plain ponds are characterized by fluctuating water levels and acidic, nutrient-poor, high quality water. These ponds harbor one of the largest concentrations of globally and statewide rare species on Long Island, including the tiger salamander, banded sunfish, six species of damselflies and dragonflies, and more than a dozen species of plants.

A distinctive community of plants has adapted to the pond shore conditions of fluctuating water levels, nutrient poor soil and water, and the acidic environment. Periods of both high and low water are essential in maintaining the structure, composition and diversity of the pond shore plant community. Periodic flooding is necessary to kill seedlings of woody shrubs and trees invading from surrounding upland, (Zaremba and Lamont 1993), and may also be important in transporting nutrients and seeds. Periods of low water are required to expose substrate for seed germination and growth. In low water years an enormous diversity of sedge, grass and flowering herb species appear, growing in concentric rings at elevations determined by the species' need for nutrients and tolerance of flooding. Recent studies have revealed that the natural heterogeneity of coastal plain pond shores appears to be a result of long-term fluctuations in water levels, spatial and temporal variability in nutrients, and reproductive strategies. (Schneider 1994). The ability of many seeds to remain dormant in high-water years plays an essential role in the capacity of these species to respond when pond margins are exposed in low-water years.

Five distinct vegetation zones (habitats) have been described for coastal plain pond shores by Zaremba and Lamont. (Zaremba 1993). Different species are characteristic of each of these different zones:

1. Upper wetland shrub thicket. This is often a Pine Barrens shrub swamp community.
2. Upper, low herbaceous fringe. A narrow band of vegetation on a peaty substrate that is only marginally above general high water levels, and may not occur at all ponds.
3. Sandy exposed pond bottom. The outermost pond bottom exposed during droughts is often very sandy and dominated by annual species. Soil moisture is related to groundwater depth. The highest concentration of rare species occurs in this zone.
4. Organic exposed pond bottom. The organic exposed pond bottom is more frequently flooded than the sandy zone, hence has a greater accumulation of organics. This zone has a low species diversity.
5. Permanently flooded zone with emergent and floating species. Pond bottom substrates are generally covered with loose layers of organic debris. Species diversity and overall cover are generally low.

5.7.2 Pine Barrens shrub swamps G5 S3

Pine Barrens shrub swamps are wetlands that often occur at the margins of coastal plain ponds, as a transition zone between the pond shore and the surrounding pine barrens forest. Shrub swamps also occur in wet depressions with little or no standing water. (Reschke 1990). Characteristic species include leatherleaf (Chamaedaphne calyculata), highbush blueberry (Vaccinium corymbosum), sweet pepper-bush (Clethra alnifolia), male-berry (Lyonia ligustrina), fetterbush (Leucothoe racemosa), buttonbush (Cephalanthus occidentalis) and winterberry (Ilex glabra). Sphagnum mosses are common in the groundlayer.

5.7.3 Coastal plain Atlantic white cedar swamp G3G4 S1

Coastal plain Atlantic white cedar swamps occur on "organic soils along streams and in poorly drained depressions . . . Atlantic white cedar (Chamaecyparis thyoides) makes up over 50% of the canopy cover." (Reschke 1990). Red maple (Acer rubrum) may be a codominant tree. Characteristic shrubs include sweet pepperbush (Clethra alnifolia), highbush blueberry (Vaccinium coryumbosum, and swamp azalea (Rhododendron viscosum). Several species of sphagnum moss dominate the groundlayer.

Atlantic white cedar regenerates only under conditions of full sunlight and moist, but not flooded, conditions. Mature stands of Atlantic white cedar are dense, with little light penetration of the canopy. Therefore white cedar regeneration occurs only following removal of mature trees by processes such as logging, fire, high wind, protracted flooding, or occasional salt water flooding resulting from severe storms. (Laderman 1989). Historically, fires or blowdowns were the most common canopy-opening events. The composition of stands following fire depends on the amount of organic soil consumed by the fire, the presence of hardwoods and shrubs in the killed stands, and available seed sources. (Little 1979). Fires during high water periods favor regeneration of white cedars. Intense fire when water levels are low results in peat burning, and a lowering of the forest floor. The resultant flooding prevents white cedar regeneration. A bog, shrub swamp or deciduous hardwood swamp may then form. (Laderman 1989). If substrate conditions are unsuitable, or sources of white cedar seed have been eliminated, white cedar may fail to become established and the area will instead become a red maple-hardwood swamp.

Flooding or draining due to human-caused alterations of the hydrologic regime also could cause the loss of white cedar. Degradation of water quality by direct storm water runoff, or due to groundwater contamination from fertilizers or septic tank leachate, has the potential to adversely affect white cedars and characteristic cedar-associated species. (Ehrenfeld 1983, Schneider and Ehrenfeld 1987).

Ironically, the complete protection of white cedar swamps will ensure their eventual senescence and loss. There are ". . . no simple, definitive guidelines for optimal management practices of cedar wetlands; there are too many biological unknowns for any simple formula." (Laderman 1989). Little proposed an approach to cedar management which involved clearcutting or strip cutting, the removal of slash, the control of competing hardwoods, and the control of deer browse. (Little 1950).

White cedar swamps were formerly extensive on Long Island, but have been reduced to a few scattered remnants by past logging and draining. (Laderman 1987). The largest white cedar swamp remaining on Long Island is in Cranberry Bog County Park in Southampton Town (between the Peconic River and Riverhead-Moriches Road).

The highly acidic waters of white cedar swamps limit use by wildlife. Nevertheless, swamp sparrows (Melospiza georgiana) can be found here and four-toed salamanders (Hemidactylium scutatum) inhabit the sphagnum zone. Most noteworthy is the rare Hessel's Hairstreak butterfly (Mitoura hesseli), which is dependant upon the white cedar as its host plant.

5.7.4 Red maple-hardwood swamps G5 S4S5

Red maple-hardwood swamps occur "in poorly drained depressions, usually on inorganic soils. This is a broadly defined community with many regional and edaphic variants." (Reschke 1990). On Long Island red maple and black gum (Nyssa sylvatica) are the dominant trees. The shrub layer may be quite dense. Characteristic shrubs are sweet pepperbush (Clethra alnifolia), highbush blueberry (Vaccinium corymbosum), arrowwood (Viburnum recognitum) and swamp azalea (Rhododendron viscosum). Herbs include cinnamon fern (Osmundea cinnamomea), Royal fern (O. regalis), and skunk cabbage (Symplocarpus foetidus).

Insectivorous birds use the red maple-dominated swamps at all times of year. A wide variety of migratory insectivorous birds uses the deciduous swamps during their spring and fall movements. The early flowering red maple attracts a rich insect life which is used by vireos, warblers, thrushes and many others, including dozens of species of birds which winter in the tropics.

During the colder months, a guild which includes kinglets (Sylviidae), nuthatches (Sittidae), woodpeckers (Picidae), titmice (Paridae) and brown creepers (Certhia familiaris) searches for insect larvae in the many dead trees in these swamps.

5.7.5 Coastal plain poor fen G3? S1

Coastal plain poor fens are sphagnum moss-dominated peatlands, with scattered sedges, shrubs, and stunted trees. (Reschke 1990). The largest such fen on Long Island, at Cranberry Bog County Park, is dominated by sphagnum and sedges. (Reschke, personal communication). This relatively rare (G3?S1) community appears to be a successional stage in transition to a shrub or hardwood swamp.

5.7.6 Coastal plain stream G3G4 S1

Two of the four major rivers of Suffolk County are found in the Pine Barrens: the Peconic River and the Carmans. These slow-moving, often darkly-stained streams may contain abundant submerged vegetation, including pondweeds (Potamogeton pusillus, P. epihydrus), naiads (Najas flexilis, N. guadalupensis), waterweeds (Elodea nuttallii, E. candadensis), stonewort (Nitella spp.), bladderwort (Utricularia vulgaris), duckweed (Lemna minor) and white water-crowfoot (Ranunculus trichophyllus).

Cool streams are home to the eastern mudminnow (Umbra pygmaea), banded killifish (Fundulus diaphanus), and, where stocked, brown trout (Salmo trutta). Swamp darter (Etheostoma fusiforme), which only exist in this region of the state, may occur where surface waters are unpolluted and free of turbidity and silt, in both coastal streams and swamps. The tessellated darter (Etheostoma olmstedi) is attracted to similar habitats, but also occurs in large open bodies of water. American eels (Anguilla rostrata) can be found in some streams as well as in the tidal waters. Three species of sticklebacks: the ninespine (Pungitius pungitius), the threespine (Gasterosteus aculeatus) and fourspine (Apeltes quadracus) are also common.

5.7.7 Low salt marsh G4 S3S4; High salt marsh G4 S3S4

Salt marsh occurs in the Hubbard Creek Marsh on Peconic Bay, in the Flanders area. Low salt marsh extends from mean high tide down to mean sea level and is regularly flooded by semidiurnal tides. Vegetation is a nearly monospecific stand of cordgrass (Spartina alterniflora). High salt marsh occurs from mean high tide up to the limit of spring tides and is periodically flooded by spring tides and flood tides. Vegetation consists of a mosaic of patches dominated by either dwarf cordgrass or salt-meadow grass (Spartina patens). Also common are spikegrass (Distichlis spicata) and black-grass (Juncus gerardi).

The very limited expanse of saltmarsh within the Central Pine Barrens supports the full range of wildlife typical of saltmarsh elsewhere on Long Island. Many waterbirds rely upon marine organisms for food. The NYSDEC listed endangered piping plover (Charadrius melodus) is known to breed here, as is the endangered least tern (Sterna antillarum) and the threatened common tern (Sterna hirundo). The diamondback terrapin (Malaclemys terrapin), our only saltmarsh reptile, lives here year round and nests in nearby uplands.

5.7.8 Salt panne G3G4 S3

A salt panne is a poorly drained, shallow depression in both low and high salt marshes, found in the Hubbard Creek Marsh on Peconic Bay. Soil water salinities fluctuate in response to tidal flooding and rainfall. Pannes in low salt marsh usually lack vegetation. Pannes in high salt marsh are irregularly flooded, and are vegetated by dwarf cordgrass, glassworts (Salicornia europaea and S. virginica), marsh fleabane (Pluchea odorata), salt marsh plaintain (Plantago maritima ssp. junkets), arrowgrass (Triglochin maritimum) and salt marsh sand spurry (Spergularia marina).

Pond holes within the pannes may contain mummichog (Fundulus heteroclitus) and sheepshead minnow (Cyprinodon variegatus).

5.7.9 Salt shrub G5 S4

Salt shrub occurs in the Hubbard Creek Marsh on Peconic Bay, in the Flanders area. Salt shrub is a shrubland community that forms an ecotone between salt marsh and upland vegetation. Salinity levels are generally lower than in the salt marsh and the elevation is higher. (Reschke 1990). Characteristic shrubs are the groundsel-tree (Baccharis halimifolia), saltmarsh elder (Iva frutescens). Typical herbs are the salt-meadow grass (Spartina patens) and switchgrass (Panicum virgatum).

Both pitch pine, red maple and black gum are found in this community, which is transitional between upland Pine Barrens and wetland communities such as red maple swamps and shrub swamps. The shrub and herb flora is very similar to that of red maple swamps.

5.7.10 Pine Barrens vernal pond (G3G4 S2 elsewhere in New York); Vernal pool (G4 S3S4 elsewhere in New York)

Pine Barrens vernal ponds, and vernal pools, are not yet documented by the Heritage Program for the Central Pine Barrens, but wetlands resembling these communities do occur. Alternatively, these wetlands may be considered variants of coastal plain ponds, red maple-hardwood swamps or pine barrens shrub swamps. These wetlands are seasonally fluctuating, groundwater-fed ponds dominated by grasses and herbs. At some sites these are mixed with low shrubs. These wetlands are often small, covered by a closed-over tree canopy, and carpeted with leaf litter. (Zaremba, personal communication). Plant species composition is similar to that of shrub swamps and red maple-hardwood swamps.

5.8 Human-created Communities

In this document a distinction between "natural" and "human-created" communities has been drawn. This dichotomy is artificial, and somewhat arbitrary, but nevertheless useful. The range of communities found within the Central Pine Barrens exists along a continuum where, at one extreme, communities exhibit relatively few effects from human activities. At the other extreme, the communities are the direct result of human manipulation of the environment (e.g., lawns and agricultural lands). Lands which have been previously burned, logged, cleared, farmed, or developed may show the effects of these activities for decades or centuries. Communities are treated as "natural" wherever the overwhelming forces which have shaped their present state have been natural. The strongest such force is succession, but fire and the natural dispersion of organisms also play major roles.

Historically, sun-loving herbaceous plants must have been the most common in grassy patches and open areas created by fire or other disturbance within the Pine Barrens. Widespread development and fire suppression has resulted in a loss of these openings. (Southampton 1993). Some of these herbaceous plants now are rare, and persist primarily in human-created habitats such as roadsides, successional old fields, and mowed areas.

5.8.1 Successional old field G4 S4; Successional Shrubland G4 S4

Successional old field is a "meadow dominated by forbs and grasses that occurs on sites that have been cleared and plowed (for farming or development), and then abandoned. This is a relatively short-lived community that succeeds to a shrubland, woodland or forest community." (Reschke 1990). Weedy non-native species typically are a major component of the flora. These include bluegrasses (Poa pratensis, P. compress), timothy (Phleum pratense), quackgrass (Agropyron repens), sweet vernal grass (Anthoxanthum odoratum), orchard grass (Dactylis glomerata), chickweed (Cerastium arvense), Queen-Anne's lace (Daucus carota) and dandelion (Taraxacum officinale). Native species include goldenrods (Solidago rugosa, S. juncea, S. nemoralis, Euthamia graminifolia), old-field cinquefoil (Potentilla simplex), asters, evening primrose (Oenothera biennis) and ragweed (Ambrosia artemisiifolia).

When woody cover increases to 50% or more, the community becomes a successional shrubland. (Reschke 1990). Characteristic woody species include red cedar (Juniperus virginiana), blackberries (Rubus spp.), hawthorne (Crataegus spp.), choke-cherry (Prunus virginiana), serviceberries (Amelanchier spp.), sumac (Rhus glabra, R. copallina), arrowwood (Viburnum recognitum) and multiflora rose (Rosa multiflora).

The mixed shrublands and hedgerows support a bird community typical of sunny but densely tangled vegetation. Grey catbirds, song sparrows, yellow warblers and bobwhite quail are common. Blue-winged warblers can be found near the taller vegetation. Cottontail rabbits flourish amidst the undergrowth. Woodcocks (Philohela minor) perform their courtship flights here in the spring. The numerous shrubs and saplings attract field sparrows and cedar waxwings (Bombycilla cedrorum). Fallen fruits feed meadow jumping mice (Zapus hudsonius), raccoons, box turtles and others.

Meadow voles (Microtus pennsylvanicus), black racers and garter snakes and kestrels (Falco sparverius) are most abundant in the grasslands. Grasshopper sparrows (Ammodramus savannarum) use grasslands if the taller herbs or woody plants provide them with suitable singing perches. Meadowlarks (Sturnella magna) and bobolinks (Dolichonyx oryzivorus) use areas where the grass is shorter, as do upland sandpipers (Bartramia longicauda) and rough-legged hawks (Buteo lagopus).

5.8.2 Cropland/row crops G5 S5

Areas of active agriculture in the Pine Barrens include sod farms, nurseries, and truck crops (vegetables and fruits).

Canada geese (Branta canadensis), white-tailed deer, mourning doves, raccoons, crows, blackbirds (Icteridae), starlings (Sturnus vulgaris) and English sparrows (Passer domesticus) all forage in cultivated farmlands and may be pests to farmers. Killdeer (Charadrius vociferus), horned lark (Eremophila alpestris), water pipits (Anthus spinoletta) and other birds use the bare earth of fallow fields.

5.8.3 Mowed lawn G5 S5; mowed lawn with trees G5 S5

"Residential, recreational, or commercial land in which the groundcover is dominated by clipped grasses and forbs. . . . Ornamental and/or native shrubs may be present." (Reschke 1990).

5.8.4 Mowed roadside/pathway G5 S5

Included in this category are infrequently mowed areas along roads, pathways mowed through meadows, old fields, woodlands, forests, or utility right-of-way corridors. The vegetation may resemble that of old fields and shrublands; either grasses, sedges and rushes, or forbs, vines and low shrubs may dominate. Mowed areas at the Naval Weapons Industrial Reserve Plant (NWIRP) at Calverton are grasslands dominated by little bluestem (Schizachyrium scoparius), spike grass (Danthonia spicata), switchgrass (Panicum virgatum), asters, goldenrods, false indigo (Baptisia tinctoria) and sweet fern (Comptonia peregrina). Diversity is lower in frequently mowed areas near the runway aprons, and higher at less disturbed sites. (NWIRP 1989).

Residential, industrial and institutional land uses generally involve both structures and landscaped areas. These developed areas frequently provide wildlife habitat. Northern orioles, chipping sparrows (Spizella passerina) and house finches (Carpodacus mexicanus) nest in shade trees. Blackbirds, starlings, robins (Turdus migratorius) and northern flickers (Colaptes auratus) feed on lawns. Deer and geese will graze on the larger expanses. Norway rats (Rattus norvegicus), barn swallows, grey squirrels and raccoons reside within many buildings.

5.9 Occurrences of Rare Pine Barrens Natural Communities and Species

5.9.1 Communities

The Natural Heritage Program records a total of 52 occurrences of state rare natural communities (S1-S3) in the Central Pine Barrens, as shown in Figure 5-5. Of these, almost all are within the Core Preservation Area. Only one occurrence of pitch pine-oak-heath woodland and six occurrences of coastal plain pond shores are in the Compatible Growth Area. It is important to note that there has not been a comprehensive documentation of all occurrences of every community type in the Central Pine Barrens. Therefore, the list in Figure 5-5 may be incomplete.
Figure 5-5: Occurrences of Rare (S1-S3) Natural Communities in the Central Pine Barrens
Natural Community Type
Number of Documented Occurrences
Coastal plain pond shore
Coastal plain pond
Coastal plain poor fen
Coastal plain Atlantic white cedar swamp
Pine barrens shrub swamp
Dwarf pine plain
Pitch pine-oak-heath woodland
Salt panne

It is noted that all of the ponds and pondshores in the Core Area are ranked B2 or B3 for overall biodiversity significance by the New York Natural Heritage Program. (Appendix 2-1).

Of the six pondshores in the compatible growth area, two are ranked B2 (Lake Panamoka, North Pond), three are ranked B3 (Artist Lake, Currans Road South, Overton Road Pond), and one is ranked B4 (Coreys Pond). Despite human impacts (physical disturbance of the pondshores, altered water chemistry), four of these six ponds remain of special biological value based upon their biodiversity rank, quality of their pondshores (EO rank, Appendix 2-1), and suite of rare species. (Zaremba, personal communication). These four ponds are Lake Panamoka, North Pond (near Lake Panamoka), Currans Road South Pond, and Overton Road Pond.

5.9.2 Plants

A total of 205 occurrences of 54 rare plant species (S1-S3) have been documented in the Core Preservation Area. In the Compatible Growth Area 35 occurrences of 18 rare plant species have been documented. (Appendix 2-2). Many additional rare species have been known to occur in the Pine Barrens in the past, but have not been documented in recent decades. There has not been a comprehensive documentation of all species of rare plants in the Central Pine Barrens. Further searches could add more species to those listed in Appendix 2-2.

By far, the greatest concentrations of known rare plant species occur in wetland habitats: coastal plain pondshores (19 species), coastal plain ponds (11), wet pine barrens (10) and red maple-hardwood swamps (5). (Figure 5-5). Mowed areas, roadsides, and other open sites support 8 rare plant species. Six (6) rare species are found in salt marsh communities. (Figure 5-5). Very few rare plant species are found in forest or woodland communities, and those few are associated with open, sandy areas including roadsides (Cyperus houghtonii, Minuartia caroliniana, Prunus pumila var depressa).

5.9.3 Animals

Rare vertebrate species are the banded sunfish, tiger salamander, eastern mud turtle, osprey, piping plover, common tern and least tern. All of the vertebrate species may also be found in habitats and communities outside of the Pine Barrens. Rare invertebrate species include moths, butterflies and damselflies. Most of these rare invertebrate species occur only in Pine Barrens habitats and are absent or uncommon elsewhere on Long Island. There has not been an attempt to comprehensively document all species of vertebrate and invertebrate animals in the Central Pine Barrens. Further searches could add more species (especially moths and butterflies) to those discussed below.

Banded sunfish: Enneacanthus obesus (G5 S2) is listed by NYSDEC as a Species of Special Concern. In New York, it is known from 4 sites within the Peconic River drainage. It prefers slow-moving, weedy areas.

Habitat needs: The banded sunfish's aquatic habitat should be protected from loss and degradation. The known sites are all on public land within the Core Preservation Area which effectively precludes direct disturbance and loss.

Tiger salamander: Ambystoma tigrinum (G5 S3) is listed by NYSDEC as endangered. This mole salamander is known from 61 sites within the Central Pine Barrens. Tiger salamanders use small ponds for breeding sites. Fish-free waters allow breeding salamander adults, eggs and larvae to avoid a major source of predation. During the non-breeding season, adults and post-larval young dwell in tunnel systems beneath adjacent woodlands. They frequently range as much as 1000 feet from breeding ponds. Little scientific information exists as to the extent of movements by individuals amongst breeding sites. Nevertheless, it is obvious that populations have dispersed over time. Tiger salamanders exist on Long Island from Nassau County eastward onto the South Fork.

Habitat needs: The majority of the tiger salamander's known populations (38 of 61) are within the Core Preservation Area. In the compatible growth area, numerous sites require protective efforts by state and local governments. Although there is no law which specifically protects the habitats of endangered or threatened species, existing laws are currently being used to protect a significant portion of the animal's habitats. The breeding ponds are fully protected by both state and local wetlands laws. Much of the surrounding upland habitat is also protected under the New York State Environmental Quality Review Act (SEQRA). Generally, at least half of the uplands within 1000 feet of each known breeding pond is preserved as large blocks of naturally-vegetated open space.

Protection for corridors which connect known sites should be considered, as without opportunities to travel amongst sites, populations could become genetically isolated and subject to local extinction.

Eastern mud turtle: Kinosternon subrubrum (G5S1) is listed by NYSDEC as threatened. It is known from only 4 locations in New York State; 3 of these are at least partially within the Central Pine Barrens. This species uses "shallow, slow- or non-flowing fresh or brackish water with soft bottom and abundant aquatic vegetation." It nests and hibernates in nearby uplands. On Long Island it is known to use areas as much as 400 feet from ponds.

Habitat needs: The majority (3 of 4) of the mud turtle's known populations extend beyond the Central Pine Barrens. One is on private lands within the Compatible Growth Area. As discussed above, there is no law which specifically protects the habitats of endangered or threatened species but existing laws are currently being used to protect a significant portion of the animal's habitats. The breeding ponds are fully protected by both state and local wetlands laws. The surrounding upland habitat is protected under SEQRA.

The following species are found within the Central Pine Barrens but are far more common in coastal areas elsewhere on Long Island. Each is listed as either threatened or endangered, and is currently protected through regulation and through management. These species will be discussed only briefly.

Osprey: Pandion haliaetus (G5 S4) is now known from over 250 nests on Long Island each year. Three (3) of these are found within the Central Pine Barrens.

Piping plover: Charadrius melodus (G3 S2) is known from over 70 actives nest sites on Long Island each year. One of these is within the Central Pine Barrens.

Common tern: Sterna hirundo (G5 S3) is known from over 60 actives nest sites on Long Island each year. One of these is within the Central Pine Barrens.

Least tern: Sterna antillarum (G4 S3) is known from over 60 actives nest sites on Long Island each year. One of these is within the Central Pine Barrens.

Invertebrate Wildlife: The Natural Heritage Program lists 42 recent occurrences of rare invertebrate wildlife in the Central Pine Barrens. Forty-one (41) are within the Core Preservation Area, the other occurrence is within the Compatible Growth Area.

5.9.4 Upland species

The species below are found within the Central Pine Barrens. Virtually all are restricted to Pine Barrens habitat types and are absent or uncommon elsewhere on Long Island. One species, the White M Hairstreak, is found within the Compatible Growth Area. All others are known from the publicly owned land within the Core Preservation Area. As a result, their habitats are protected.

Coastal barrens buck moth: Hemileuca maia maia is found from 10 locations within the Central Pine Barrens; it attains its highest densities, over 1000 adults per acre, in the dwarf pine plains. It requires scrub oak and dwarf chinquapin oak as a host plant.

Habitat needs: Scrub oak-dominated communities need to be maintained.

Frosted elfin: Incisalia irus is a butterfly which is known from a single swale. It appears to require wild indigo (Baptisia tinctoria) and blue lupine (Lupinus perennis) as host plants.

Habitat needs: Weedy areas with wild indigo and blue lupine are required. Although the single known site is within the Core Preservation Area, it is a managed railroad right-of-way. Natural succession could threaten the future of this habitat type. Mowing or prescribed burning may be useful in maintaining such areas.

White M hairstreak: Parrhasius M-album is a butterfly which is known from a single old field in the Compatible Growth Area. This species appears to use goldenrods (Solidago spp.) and milkweed (Asclepias spp.) as host plants.

Habitat needs: Natural succession could threaten the future of this habitat type. Mowing or prescribed burning may be useful in maintaining such areas.

5.9.5 Dwarf Pine Plains Species

A noctuid moth with no common name, Chaetaglaea cerata is known from a single occurrence.

A noctuid moth with no common name, Heterocampa varia is known from 2 occurrences and is believed to depend upon pitch pine and oaks.

Jair underwing, Catocala jair ssp. 2, is a moth which depends upon oak and blueberry leaves.

Herodias underwing, Catocala herodias gerhardi, is a moth which depends upon oak (especially scrub oak) and blueberry leaves. It is known from 2 locations.

Dusted skipper, Atrytonopsis hianna, is a butterfly known from a single location. It appears to rely upon big and little bluestem grasses as host plant.

Violet dart, Euxoa violaris, is a butterfly which is known from a single occurrence.

Zale species #1, Zale lunifera, is a moth which is known from a single occurrence.

Pink sallow, Psectraglaea carnosa, is known from 2 occurrences.

Habitat needs for dwarf pine plains species: These species all require that the dwarf pine plains' natural community be protected. The location of this area within the Core Preservation Area affords it some protection. It is likely that the dwarf pine plains will need to be actively managed, perhaps through burning, to maintain its existing habitat characteristics.

5.9.6 Wetland species

Hessel's hairstreak, Mitoura hesseli, is a butterfly which is known from 4 Atlantic white cedar swamps. It relies upon this cedar (Chamaecyparis thyoides) as its host.

Pitcher plant borer, Papaipema apassionata, is a moth which is known from a single bog. It relies upon the pitcher plant (Sarracenia purpurea) as its host.

Chain fern borer, Papaipema stenocelis, is a moth which is known from a single site. It feeds on the rhizomes of Virginia chain fern (Woodwardia virginiana).

Lateral bluet, Enallagma laterale, is a damselfly which is known from 2 ponds. Eggs are laid in emergent vegetation. Larvae are aquatic, and post larval stages (tenerals) inhabit nearby upland vegetation.

Barrens bluet, Enallagma recurvatum, is a damselfly which is known from 8 ponds. Eggs are laid in emergent vegetation. Larvae are aquatic, and post larval stages (tenerals) inhabit nearby upland vegetation.

Painted bluet, Enallagma pictum, is a damselfly which is known from 2 ponds. Eggs are laid in emergent vegetation. Larvae are aquatic, and post larval stages (tenerals) inhabit nearby upland vegetation.

Round-necked damselfly, Nehalennia integricollis, is known from a single pond.

Habitat needs for wetland species: The above species' aquatic habitats need to be protected from loss and degradation. The known sites are all on public land and within the Core Preservation Area which effectively precludes direct disturbance and loss.

5.10 Ecological Principles of Conservation Reserve Design

In order to preserve natural ecosystems and halt the extinction of species it is necessary to think on a large scale, at the level of ecosystems and landscapes, (Franklin 1993), and to establish large core reserves, or assemblages of linked reserves, surrounded by "semi-natural" buffer zones. (Franklin 1993, Noss 1992, Noss and Cooperrider 1994). Core reserves should be large enough and extensive enough to include the full range of communities, successional stages, physical habitats, environmental gradients, and ecosystem processes in a region. The buffer zone should be managed to minimize the ecological contrast between the buffer and the core. (Franklin 1993). The extensive literature on the optimal design of conservation reserves has been reviewed by Shafer. (Shafer 1990). Only the most important ecological considerations are discussed here. These considerations are general guidelines which were interpreted within the context of the Long Island Pine Barrens.

Ecological processes. In order to maintain functional, viable ecosystems in perpetuity, it is essential to maintain ecological and evolutionary processes, such as natural disturbance regimes (primarily fire in the pine barrens), hydrological processes (including surface and groundwater flow and fluctuating water levels in coastal plain ponds), nutrient cycles, genetic selection and biotic interactions (including predation). (Noss 1992, Noss and Cooperrider 1994). Large reserves are more likely to allow ecological processes to continue than are small, isolated reserves. In some cases active management may be required to maintain essential ecological processes that are no longer operable due to human alterations of the landscape.

Edge effects. Organisms located at or near the transition zone between a naturally vegetated, original habitat and an adjoining disturbed area (matrix) are subject to "edge effects," including increased sunlight and wind, altered humidity, altered temperatures, exposure to invasive exotic species, increased predation, increased exposure to human disturbance, etc. These edge effects may extend considerable distances into the natural habitat. Edge effects may favor some species while eliminating other species, or greatly modifying their growth, viability and genetic makeup.

The following material concerning wildlife reserves has been excerpted from McDougal's 1994 work.

Size. The general rule for optimizing the effectiveness of a conservation area is "bigger is better." Large reserves offer both more area per se and generally can be expected to exhibit a greater degree of habitat heterogeneity than small ones. They are also more likely to allow the continued operation of disturbance regimes at a range of scales. These processes act to maintain spatial and temporal heterogeneity in "whole" ecosystems but generally are not functional in small, isolated preserves. (Noss and Harris 1986; Pickett and Thompson 1978). Large reserves are more likely to allow for the persistence of viable populations of species with relatively large area requirements, those that depend upon more than one habitat type for their resource needs and ephemeral species which are dependent upon a dynamic mosaic of newly created habitat for continued presence in the region. Large reserves are also more likely to protect metapopulations (complexes of interdependent subpopulations which are affected by recurring extinctions, and linked by recolonization from large reservoir populations) than are small reserves. (Murphy et al., 1990).

Population size. Small populations are particularly subject to fluctuations and random events, and random extinctions due to demographic and environmental [variability] are more important in small populations. (Shaffer 1981). The first species to disappear from small reserves are frequently those at high trophic levels, or larger, more specialized members of feeding guilds. (Wilson and Willis 1975). Species with variable populations that are dependent on patchy or unpredictable resources seem particularly sensitive. (Noss 1983). Since nature reserves are essentially habitat islands (i.e., areas of natural landscape surrounded by a culturally modified matrix), the potential for recolonization of a species that becomes extinct within the reserve is often limited or non-existent. This makes it critical that the area of the reserve is large enough to maintain viable populations of its component species. When this is not the case, and as the surrounding matrix is further altered extinction may become an important process within the reserve. (Pickett and Thompson 1978). Large populations are more likely to maintain greater levels of genetic variability than small populations, and this can be an important factor in allowing adaptive response to changing environmental conditions.

Shape. Fragmentation of habitat into isolated remnant patches generates a number of problems including 1) reduction in the effective size of the reserve due to physical and biological edge effects, 2) increasing domination by species characteristic of human landscapes and 3) the loss of large, wide-ranging, or ecologically specialized species. (Noss and Harris 1986). Given two habitat patches of equal area but different shapes, these problems are often more pronounced in those with a significantly greater edge: interior ratio. These factors become particularly important in extreme cases where a reserve is so small or narrow that no interior habitat exists within it. In large reserves shape may be less important. In general it has been suggested that a design that maximizes interior habitat and minimizes edge effects is most desirable. When this is not possible a buffer zone adjacent to the core preservation area is important.

Buffer zones. Edge effects can be dramatic, and the higher the contrast between the 'natural' landscape of the reserve and the culturally modified matrix surrounding it the greater these effects will be. (Franklin 1993). The role of the buffer zone is that of a transition area which filters out the effects of [human] disturbances (Baker 1992), or minimizes their impact when they reach the core area. The buffer zone can be considered an ecological boundary, a zone of interaction and change between the core reserve and the surrounding matrix. (Schoenwald-Cox 1988). In these areas the rates or magnitudes of ecological transfers (e.g., energy flow, nutrients, species) change abruptly in relation to those on either side of the boundary. Boundaries may vary in their permeability or resistance to these transfers as a consequence of the nature of the boundary itself and the responses of different materials, organisms or abiotic factors to the boundary. (Wiens et al., 1985). The effectiveness of the buffer zone as a transition area depends on its size, ecological characteristics and human activities within it.

Connectivity. Many writers on the design of nature reserves advocate the use of corridors which may act as dispersal routes, facilitating recolonization, gene flow and increasing the effective size of small populations. In fact, empirical evidence on the use of corridors is meager. (Simberloff et al., 1992; Shafer 1990). What actually constitutes a corridor or a barrier is mostly based on assumption. (Forman and Godron 1986). Both are scale dependent and will be perceived differently by different species. (Hanson and Angelstam 1991). It is probably reasonable and prudent to assume however, that connectivity at all scales is more desirable than little or no connectivity. This is particularly true in those cases where a reserve is irregularly shaped and the connection of adjacent parts may add to the areas conservation value.

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