Agalinis acuta - Pennell
Sandplain Gerardia
Other English Common Names: Sandplain False Foxglove
Other Common Names: sandplain false foxglove
Synonym(s): Gerardia acuta (Pennell) Pennell
Taxonomic Status: Not accepted
Related ITIS Name(s): Agalinis acuta Pennell (TSN 33009)
Unique Identifier: ELEMENT_GLOBAL.2.140310
Element Code: PDSCR01010
Informal Taxonomy: Plants, Vascular - Flowering Plants - Figwort Family
Image 10396

© Alfred R. Schotz

 
Kingdom Phylum Class Order Family Genus
Plantae Anthophyta Dicotyledoneae Scrophulariales Scrophulariaceae Agalinis
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Concept Reference
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Concept Reference: Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
Concept Reference Code: B94KAR01HQUS
Name Used in Concept Reference: Agalinis acuta
Taxonomic Comments: Called Gerardia acuta in some older floras, but the generic name Gerardia, as used by Linnaeus, applies instead to some members of the Acanthaceae (Crow, 1982), rather than these plants of the Scrophulariaceae.  In a study using genetic and morphological analyses, Pettengill and Neel (2011) conclude that Agalinis acuta should be synonymized under A. decemloba, and also conclude that A. decemloba "is deserving of protection under the Endangered Species Act".  
Conservation Status
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NatureServe Status

Global Status: G1
Global Status Last Reviewed: 19Jan2005
Global Status Last Changed: 22Jun1990
Rounded Global Status: G1 - Critically Imperiled
Reasons: As of 2005, about 22-23 occurrences were known to be extant, but only a few of these support large populations or have large areas of occupied habitat, or even suitably managed potential habitat. The stronghold of the species may once have been New York where populations today occurs on a small, degraded remnants of a grassland that once covered about 20,000 ha in central Long Island. Six extant sites occur in small grassland areas of Massachusetts, two of which occur in old cemeteries that have probably been mowed regularly for over 100 years. Massachusetts has been successful in expanding the four rediscovered sites through management and augmentation, and it has introduced the species successfully at two additional protected sites now treated as separate EO's. Maryland has two populations, the largest being located on a well protected and managed serpentine barren ecosystem. Connecticut and Rhode Island each have one remaining population which they are attempting to manage and augment with modest success thus far. The species is threatened by the on-going loss of its coastal grassland habitats due to development. The loss of grazing animals and the suppression of fires have allowed woody vegetation to claim many of its historical sites.
Nation: United States
National Status: N1

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Connecticut (S1), Maryland (S1), Massachusetts (S1), New York (SNR), Rhode Island (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LE: Listed endangered (07Sep1988)
Comments on USESA: Agalinis acuta was recognized as a Category 2 candidate in the December 15, 1980 notice of review, and as a Category 1 candidate in the November 28, 1983 supplement to the 1980 notice of review and in a revised notice of September 27, 1985. It was proposed endangered on November 19, 1987 and determined endangered on September 7, 1988.
U.S. Fish & Wildlife Service Lead Region: R5 - Northeast

NatureServe Global Conservation Status Factors

Range Extent Comments: Agalinis acuta is known historically from five states: Connecticut, Rhode Island, Massachusetts, Maryland, and New York. It is primarily a species of the Coastal Plain, but in Maryland exists on the Piedmont. Connecticut is known to have supported two populations (Mehrhoff 1978), but now has one small population in a cemetery. Rhode Island harbored six populations, including one "on a sandy plain between Lime Rock and the Blackstone River" (Fernald 1906), and now has a single remnant population in a cemetery. Massachusetts once supported populations ranging from Nantucket, Marthas Vineyard, upper Cape Cod and at a few disjunct inland locations to Worcester County; It was thought to be extirpated from Massachusetts until the species was rediscovered on Cape Cod in 1980 and on Marthas Vineyard in 1994. None is currently extant on Nantucket or inland from Cape Cod. A large population occurs on a serpentine barren in Maryland, discovered in 1950, and reconfirmed in 1984. Montauk, Long Island, New York once harbored "untold millions" of Agalinis acuta plants (Taylor 1923). Agalinis acuta was thought to be extirpated from New York, but was rediscovered on Long Island and now is known from 12-13 EO's due to a combination of discoveries and restoration efforts.

Area of Occupancy: 1-25 4-km2 grid cells
Area of Occupancy Comments: Most of the occupied acreage for this taxon exists at one site, the 2000 acre Soldiers Delight Natural Area in Maryland, where there is approximately a couple hundred acres of open grassland and serpentine outcrop habitat. Most of the remaining sites consist of only a few acres or less of occupied grassland. Many, in fact, are less than an acre. The linear distance between Cape Cod and Soldiers Delight populations is estimated to be approximately 300-350 miles.

Number of Occurrences: 21 - 80
Number of Occurrences Comments: 22-23 (in 2004) Element Occurrences: 6 in Massachusetts, 12-13 in New York, 2 in Maryland, 1 in Rhode Island, 1 in Connecticut. Some of these eo's represent new introductions, reintroductions, and others have been augmented with seed sowing into managed grasslands.

Population Size Comments: Being an annual, population size varies considerably from year-to-year, but overall numbers have been increasing dramatically in response to active management at a number of sites. In 1994, the total Massachusetts population had risen to 3200 plants (representing three known sites) and in 2004 with continuing active management, augmentations and introductions, the estimate rose to 256,000 plants. Most of these plants (ca. 230K), however, occur in one dense population. Likewise, in New York, the number of sites and population sizes are increasing due to enhanced protection and management, but large fluctuations are occurring and most of the plants occur at one site where the population plummeted in 2004 (from ca. 80K to 3K). The Maryland population at Soldiers Delight increased from 200 in 1988 to ca. 10,000 (1989) and has been estimated to be well over 100K more recently under an active program of burning and mowing.

Number of Occurrences with Good Viability/Integrity: Few (4-12)
Viability/Integrity Comments: There are about 6 sites with reasonably large, stable, well managed populations. "Large" represents a population that has maintained an average of 3000+ individuals over the past 5 years. Many populations are still very small or occur on marginal habitat such as small expanses of grassland surrounded by development or unsuitably dense vegetation.

Overall Threat Impact: High
Overall Threat Impact Comments: Outright habitat destruction by human activities and habitat loss due to succession appear to be the most serious threats to most Agalinis acuta populations. Loss of grazing animals and the control of natural fires has caused the encroachment by pitch pine and scrub oak on the heathlands of Nantucket (Dunwiddie, pers. comm.). Many extant sites are on properties where some level of protection and/or management for the species has been accomplished though. At the large Maryland site, for example, where succession by Virginia pine and other woody perennials occurred due to fire suppression and cessation of grazing, protection as a natural area has occured and active management has restored habitat. Photographs taken in 1910 show no Virginia pine, but this subsequently became a dominant species over much of this serpentine barren. Aerial photographs taken in 1971 showed open vegetation in areas that by 1984 had developed a closed canopy (Knox 1984). Fortunately, management aimed at removing large areas of pine at this site has helped the population to increase substantially.

In general, the population decline of many plant species can usually be attributed wholly or to a large degree to habitat destruction caused by land development. This applies in the case of most Agalinis populations. However, more difficult to explain is the demise of a population despite the persistence of seemingly suitable habitat. The unsuccessful search for historic populations on Nantucket and Martha's Vineyard cannot be attributed entirely to habitat loss, although the alarming rate of land development on both islands makes this a real direct or indirect threat to the species. Old herbarium specimens from Nantucket are dated in the early 1900's and Martha's Vineyard specimens date from the late 1892's to 1940; a couple of these specimens notes it as frequent or common. It is not known why the species declined after this time period, but we do know that there were land use changes during this period, including the cessation of most sheep grazing, control of wildfires, and the reforestation of many open areas. Likely causes for the decline include loss of habitat to succession and lack of disturbance from grazing and burning. Except for continuing rapid development, many of these threats have run their course and the fire management is being applied selectively to reverse this trend. Small scale grazing by sheep as a conservation management tool is being used on Nantucket but is unlikely to return to a large scale again. Adequate grassland and sandplain habitat exists in protected coastal Massachusetts and New York conservation areas to restore this species to large local populations. Achieving metapopulation goals on a large landscape scale will be more difficult due to habitat fragmentation and ownership patterns.

Short-term Trend: Increase of >10%
Short-term Trend Comments: Due to protection and management efforts during the past decade, the short term trend is strongly upward. Numbers have increased dramatically at a few old sites, including several cemeteries, where remnant populations were discovered in New England. Under management and/or augmentation some populations have increased 500-1000 fold in the past 10-20 years. It is anticipated that there will be a modest or large increase, depending upon future conservation maintenance and restoration efforts. More open habitat will need to be created and maintained for the species at selected sites where the restoration potential is considered best. Some populations are unlikely to increase much more due to lack of available habitat or opportunities to restore habitat in the surrounding environment.

Intrinsic Vulnerability Comments: This species has several factors working against it. It requires frequent disturbance, such as that provided by burning, mowing or perhaps grazing, to maintain its early successional habitat conditions. Once dense cover or heavy litter builds up the species is apt to disappear. Despite being an annual that produces abundant seed, data from Massachusetts studies suggest that it typically disperses seed only over short distances (maximum of 3.5 m from previous year plots) and only seedbanks for short durations (up to 4 years only perhaps). It is also vulnerable to seed predation by insects, especially when populations become dense. Coastal populations face serious threats from severe storms that can cause wind sering, salt spray damage, overwashing or erosion of habitat by waves.

Environmental Specificity: Narrow. Specialist or community with key requirements common.
Environmental Specificity Comments: The species is a hemiparasite, but if it is like other members of the genus, there are many potential host plants ranging from grasses and forbs to woody angiosperms and gymnopsperms. Finding a suitable host may not be what limits its range, but being a hemiparasite, nevertheless, may help define a relatively narrow niche. To establish itself, the seed must germinate and quickly grow in suitably moist soil conditions to find a host root. In Massachusetts, soil testing revealed sandy loams to be typically present in sites where the species occurs naturally or where it was introduced successfully. In contrast, soils where the sand component was higher (i.e., loamy sands or coarse sands) failed to produce surviving Agalinis when seed was sown into them following burning, mowing or soil scarification pre-sowing treatments. Being a delicate annual, ground vegetation and litter must also be relatively low in cover or density. Areas that naturally maintain themselves in an early successional stage while still having soils rich and fine enough to successfully grow this delicate annual and its perennial hosts is not common on the landscape, especially with the suppression of wildfires and the reduction of grazing by large herds of herbivores, be they native or introduced. The species typically cross-pollinates when pollinators are available, otherwise it is capable of selfing to produce seed (Neel 2002).

Other NatureServe Conservation Status Information

Distribution
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Global Range: Agalinis acuta is known historically from five states: Connecticut, Rhode Island, Massachusetts, Maryland, and New York. It is primarily a species of the Coastal Plain, but in Maryland exists on the Piedmont. Connecticut is known to have supported two populations (Mehrhoff 1978), but now has one small population in a cemetery. Rhode Island harbored six populations, including one "on a sandy plain between Lime Rock and the Blackstone River" (Fernald 1906), and now has a single remnant population in a cemetery. Massachusetts once supported populations ranging from Nantucket, Marthas Vineyard, upper Cape Cod and at a few disjunct inland locations to Worcester County; It was thought to be extirpated from Massachusetts until the species was rediscovered on Cape Cod in 1980 and on Marthas Vineyard in 1994. None is currently extant on Nantucket or inland from Cape Cod. A large population occurs on a serpentine barren in Maryland, discovered in 1950, and reconfirmed in 1984. Montauk, Long Island, New York once harbored "untold millions" of Agalinis acuta plants (Taylor 1923). Agalinis acuta was thought to be extirpated from New York, but was rediscovered on Long Island and now is known from 12-13 EO's due to a combination of discoveries and restoration efforts.

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map

U.S. & Canada State/Province Distribution
United States CT, MA, MD, NY, RI

Range Map
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U.S. Distribution by County Help
State County Name (FIPS Code)
CT Hartford (09003)*, Windham (09015)
MA Barnstable (25001), Bristol (25005)*, Dukes (25007), Middlesex (25017)*, Nantucket (25019), Worcester (25027)*
MD Baltimore (city) (24510)*, Baltimore County (24005), Prince Georges (24033)*
RI Providence (44007)*, Washington (44009)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
01 Nashua (01070004)+*, Concord (01070005)+*, Farmington (01080207)+*, Cape Cod (01090002)+, Blackstone (01090003)+*, Narragansett (01090004)+*, Pawcatuck-Wood (01090005)+, Quinebaug (01100001)+
02 Gunpowder-Patapsco (02060003)+, Middle Potomac-Anacostia-Occoquan (02070010)+*
CA CAPE COD (CAPE COD)+*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A hemiparasitic, annual herb, (0.5-)1-2.5 (-3.5) dm tall, with small, linear, opposite leaves and showy flowers featuring deep pink corollas. Flowering occurs primarily in late August and early September, each flower lasting only a day.
General Description: A small, inconspicuous plant that grows from 1-4 dm tall, simple or sparsely branched. Branching is ascending, giving the plant a more erect rather than bushy appearance. It has a deep pink flower with darker spots toward a cream-colored center. The five petals are squared off and slightly indented at the ends.
Technical Description: Agalinis acuta has a slender stem, 7-30 (sometimes up to 40) cm tall, simple or sparsely branched, with four minute, low angles or wings. Branching is ascending, giving the plant a more erect rather than bushy appearance. Leaves are opposite, scabrous, linear, 0.5- 1.0 mm wide and 6-25 mm long. Flowers are borne on slender pedicels 10-20 mm long, usually 1-3 times the length of the subtending bracteal leaves. Calyx is cup-shaped by fusion into a short reticulate-veiny tube, with projecting lobes (teeth) about 0.5-1.0 mm long and narrowly triangular, fringed with short glandular hairs (calyx in one population was noted to be distinctly reticulate only after drying (Zaremba 1984). Corolla is usually 10-13 mm (sometimes as small as 5 mm) long, membranous and soft-textured, tubular, flared distally into spreading lobes which are retuse (rounded and shallowly notched) to rarely truncate. Corolla is pubescent on lobes and within tube, pink except for the white "throat" distinctly spotted with purple or red. Anthers are densely pubescent with white hairs; the style and stigma are glabrous. Capsule is 3.8-4.2 mm long and ovoid. Seeds are 0.4-0.6 mm, triangular-rounded, straw colored, with a conspicuously reticulate surface pattern. There are as many as 29 capsules per plant, although tiny individual plants sometimes bear only one or a few flowers and capsules.
Diagnostic Characteristics: The diagnostic characters of Agalinis acuta are long pedicels, shallowly notched petals, scalloped calyx border, and short calyx teeth fringed with hairs.

Four other northeastern species of Agalinis are similar in appearance to Agalinis acuta. They are A. skinneriana, A. setacea, A. decemloba, and A. tenuifolia. Characteristics which most clearly distinguish A. acuta from these species are described in the following paragraphs. (Bicknell 1915; Pennell 1929, 1935; Fernald 1950)

Agalinis acuta is found in the coastal plain region from eastern Massachusetts to Long Island, New York, with a disjunct population located in Maryland. Color of stems, leaves, and capsules is light green, drying green to straw color; stems may be angled or not, slightly scabrulous; leaves 0.6-2.5 cm long and ascending; pedicels 1-3X length; corolla lobes retuse to rarely truncate, 10-13 mm; calyx lobes reticulate with rounded sinus; capsule 3.8-4.2 mm; seed finely reticulate, straw-colored, 0.4-0.6 mm; 2n=26.

Agalinis skinneriana is found in the midwestern region of the United States and Canada, from southern Ontario and Ohio to northern Louisiana and west to Oklahoma. Color of stems, leaves, and capsules is yellow-green, with no change in color when dried; stem angles pronounced, very scabrulous; leaves 1-2 cm long, ascending; pedicels 1/2-1X length; corolla lobes not retuse, less than 13 mm long; calyx lobes reticulate with broad, flat sinus; capsule 4-5 mm; seed shallowly reticulate, yellow to tan, 0.7-0.9 mm; 2n= 26.

Agalinis setacea is found in the coastal plain and adjacent piedmont regions, from Long Island to Alabama. Color of stems, leaves, and capsules is tinged purple, drying blackish; stem angles not pronounced, slightly scabrulous; leaves 0.25-1.25 cm long, bushy and horizontal; pedicels 1-1.5X length; corolla lobes not retuse, less than 13 mm long; calyx lobes broad, flat sinus; capsule 3-4 mm; seed finely reticulate, dark brown, 0.5-0.6 mm; 2n=28.

A. decemloba is found in eastern region of the United States, from eastern Pennsylvania southwest to Kentucky, Mississippi, and Florida. Color of stems, leaves, and capsules is yellow-greenish; stem angles not pronounced, smooth; leaves 0.5-1 cm long, slender; pedicels 2-3X length; corolla lobes retuse; calyx lobes not reticulate; capsule 3-4 mm; seed finely reticulate, yellow-brown, 0.6-0.8 mm; 2n=26.

A. tenuifolia is found from Maine to Manitoba, south to Florida, and west to Texas. Color of stems, leaves, and capsules is dark green, drying blackish; stems angled, not or barely winged, not or minutely scabrulous; leaves 0.5-1.25 cm long; very narrow and bushy; pedicels 1/2-1X length; corolla lobes rounded, not truncate or retuse, not flared; calyx lobes not reticulate; capsule 3-5 mm; seed finely reticulate, brown, 0.6-0.9 mm; 2n=28.

Duration: ANNUAL
Reproduction Comments: Flowering in Massachusetts occurs between August 20th and September 30th, the peak being the first week of September (Lundgren 1983). Phenology of bloom has not been studied in Maryland populations, but flowering has been observed in early September (Ludwig, pers. comm.). Time of anthesis has been difficult to establish. The majority of fresh flowers have been observed in the morning, many dropping off by mid-afternoon. Maturation of plants continues well into October. Lundgren estimated from 0-29 fruits per plant, with a median of 8 and mean of 9. Mode of seed dispersal is unknown. If wind dispersed, seeds are not likely to travel far. Some dispersal may occur through ingestion by small animals such as meadow voles (Microtus pennsylvanica) or cottontail rabbits (Sylvilagus spp). Zaremba (1985) reported evidence of grazing on almost all of the plants (in a population of 1000-2000 plants) at Long Island site 4; herbivory was also noted at both Massachusetts populations (Zaremba and DiGregorio, pers. comm.).

Means of pollination are unknown, but fruit set at known sites suggests that pollination is not a problem. No pollinators were identified by Lundgren in her study (1984). A fly observed at one Massachusetts site was tentatively identified as Toxomerus marginatus, a common member of Syrphidae inhabiting salt marshes. Since adults of this species are known to feed on 200 plant species, it is not considered to be an effective pollinator (Maier 1985).

Ecology Comments: Agalinis acuta is an annual, so populations are dependent on seed production for continued survival. The germination and early growth of Agalinis acuta has not been monitored in natural populations.

William Brumback studied the germination requirements of Agalinis acuta in a controlled setting from 1983-1985. Four groups of seeds were sown in identical soil mixtures (one part rediearth, one part peat, one part sand) in flats and treated as follows: flats A and B were sown inside under lights at a temperature of 65 degrees F; flat C was sown in a solar greenhouse in early April; flat D was sown outside in November without a protective cover.

The indoor flats (A and B) germinated seedlings within one month, but died within seven to eight weeks of germination. Flat C produced one seedling within three weeks of planting, but no plants were evident by July. This flat was moved outside under a thin foam cover for the winter, and 14 seedlings germinated the following spring, on May 31. However, these plants were subsequently shaded out and died, and no further observations were made.

Flat D, sown outside in November 1983, produced 3 seedlings on May 2, 1984. 65 seedlings were noted by June 26. Only 10 plants remained by September, and these were in poor condition, although 2 were in flower. After wintering under the same conditions as during the previous season, 4 seedlings were noted on May 31, 1985. The flats were then transferred to the greenhouse for the winter, but no germination was observed the following spring.

From these preliminary data, Brumback concluded that although seed exposure to cold is not necessary for germination, a cold period appears to be beneficial, since the greatest number of seedlings germinated in the flat sown outside without a protective cover of foam. However, the population of plants grown under these conditions was not robust. A number of factors may have been responsible for the generally poor condition of the surviving plants, including the absence of a host species. The role of parasitism in the life cycle of Agalinis acuta needs further investigation (Brumback, pers. comm).

Populations of Agalinis acuta can fluctuate widely from year to year. Although a reduction is cause for concern, fluctuation in numbers might be a natural phenomenon in Agalinis acuta. Cutting of flowering stems does not necessarily preclude seed production, however. If severing occurs above the lowest axillary stems, the plant can produce new shoots that will successfully flower. This phenomenon was noted by Lundgren after mowing occurred in July, 1983. Herbivory has also occurred, but its effect is uncertain (Zaremba and DiGregorio, pers. comm).

Terrestrial Habitat(s): Grassland/herbaceous, Shrubland/chaparral
Habitat Comments: SUMMARY: Dry, sandy, short grass plains, roadsides, and openings in oak scrub. Dependent on periodic disturbance that maintains an open habitat. On-going disturbances at extant sites include mowing, fire, and use of a horse trail, which periodically exposes new soil. Historically, grazing was a common disturbance factor in areas where the species occurs. Grassland communities supporting Agalinis acuta are typically dominated by one or more of three grasses: little bluestem (Schizachyrium scoparium), Virginia broomgrass (Andropogon virginicus), or Indian grass (Sorghastrum nutans). Soils are nutrient-poor, usually acidic, and excessively drained. The more northerly Coastal Plain populations inhabit open sandy areas on moraines and sandplains, or glaciolacustrine terraces. The disjunct Agalinis acuta population center in Maryland occupies a nutrient-poor, circumneutral serpentine barren. An underlying factor common to all sites supporting Agalinis acuta is the lack of competition from other species - a factor imposed by conditions that include extremely nutrient-poor, and sometimes minerally toxic, soils and regular or sporadic disturbance. END SUMMARY. Wild oat grass (Danthonia spicata), panic grasses (Panicum spp.), fescue (Festuca rubra), and winter bent grass (Agrostis hyemale) occur to much lesser extent in the community. Other associated species are Maryland golden aster (Chrysopsis mariana), colicroot (Aletris farinosa), linear-leaved goldenrod (Solidago tenuifolia), toothed white-topped aster (Aster paternus), orange-grass (Hypericum gentianoides), spring ladies' tresses (Spiranthes vernalis), little ladies' tresses (Spiranthews tuberosa), several of the milkworts (Polygala spp.), purple gerardia (Agalinis purpurea), and pine-barren gerardia (Agalinis virgata). Common associates are also lichens (Cetraria and Cladonia spp.) and mosses (Polytrichum spp.), with scattered openings of bare soil.

NEW YORK: The Agalinis acuta sites on Long Island, New York range from small vulnerable populations to large and robust supporting nearly 2000 Agalinis plants. The following summaries are descriptions derived from field forms (Zaremba 1984, 1985):

1. Middle-aged Pinus rigida-Quercus ilicifolia barren is a mosaic of pines and grassy fields dominated by Andropogon gerardi, Schizachyrium scoparium, and Sorghastrum nutans. Other associates include Linum intercursum, Agalinis virgata, Scleria paucifolia, Spiranthes vernalis, Aster spectabilis, Aster solidagineus, Baptisia tinctoria, Polygala nuttallii, Rubus flagellaris, Aster linariifolius, Aster dumosus, Asclepias tuberosus, Hypericum gentianoides, Hieracium venosum, Tephrosia virginiana, Danthonia spicata, Lechea minor. The site is flat, open, xeric. Agalinis acuta occurs on the periphery of the property where mowing and disking has occurred in the past; use of herbicides is known from the 1960's. Other disturbance includes well-worn trails, and evidence of small confined fires.

2. Golf course supports a good quality maritime grassland interspersed with small wetlands and shrub thickets. Two subpopulations occur in the grassland, and a third on an adjacent maintenance area. The grassland is dominated by Schizachyrium scoparium, Andropogon gerardi, Sorghastrum nutans. Other species include Luzula bulbosa, Pycnanthemum muticum, Cirsium horridulum, Chrysopsis mariana, Anthoxanthum odoratum, Festuca rubra, Aster linariifolius, Aster paternus, Paniucm virgatum, Baptisia tinctoria, Rubus flagellaris, Lespedeza capitata. The golf course grasses are mowed frequently, and the maintenance area is moderately disturbed, with topsoil scarification and probable use of herbicides. 60, 26, and 150 Agalinis plants occur on this open, flat, dry site.

3. Schizachyrium scoparium-dominated grassy area along a roadside. 29 plants occur in this heavily disturbed area subjected to road maintenance (occasional mowing, litter, salt application). Other species here are Lechea minor, Linum intercursum, Lespedeza angustifolia, Agalinis setacea, Polygala nuttallii, Chrysopsis mariana, Lespedeza repens, Sorghastrum nutans, Aster solidagineus.

4. 10-meter strip between a major highway and railroad bed supports very small population (8 plants). Despite heavy disturbance along the track (scattered coal slag, herbicide use), the surrounding pine barrens are essentially undisturbed. Dominant grasses are Andropogon gerardi, A. virginicus, and other species include Aletris farinosa, Agalinis tenuifolia, Linum intercursum, Aster solidagineus, Liatris borealis. The surrounding pine barrens have been subject to sporadic fires due to proximity to the railroad track.

5. The largest population to date occurs within a 16-acre remnant of the once-extensive coastal plain grassland covering 40,000- 60,000 acres on central Long Island. The floristically diverse grassland was naturally maintained, unlike successional old fields. The Hempstead Plains grasslands were intensively botanized during the 1800's and early 1900's, but even at that time, pressure on land use was evident. The remaining site is moderately to severely disturbed, considered a degraded remnant (Zaremba 1984). Abandoned airfields are now populated by numerous exotic weedy species. 1000- 2000 Agalinis acuta plants occur within scraped areas in these Andropogon virginicum-dominated plains. Other species include Plantago aristata, Polygala verticillata, Hypericum gentianoides, Agalinis virgata, Rubus flagellaris, Helianthemum propinquum.

6. Rolling topography characterizes this sandy moraine located on the ocean shore. Several small Decodon wetlands occur on a perched water table underlain by clay deposits. Clethra alnifolia and Vaccinium corymbosum populate a slope in close proximity to the Agalinis site. Andropogon virginicus-dominated openings are surrounded by maritime shrubs. Associated species include Chrysopsis mariana, Aster paternus, Danthonia spicata, Solidago tenuifolia, Schizachyrium scoparium, Solidago nemoralis, Baptisia tinctoria, Anthoxanthum odoratum, Panicum depauperatum. Agalinis acuta occurs along a horse trail in exposed mineral soil caused by heavy trail use. Survey of the site in 1983 revealed about 150 plants; revisit in 1984 saw 132 plants. This site, originally discovered by Larry Penny, is probably the most natural grassland community of the 5 Long Island sites, and population numbers appear to be essentially stable.

MASSACHUSETTS: The two extant sites supporting Agalinis acuta in Massachusetts occur in small cemeteries on Cape Cod. Soils are sandy, and the grassland vegetation has probably been mowed regularly for over 100 years. Both are historic cemeteries with gravestones dating to the 1700's. The grassland openings are less than 1/2 acre in size, and surrounding vegetation is predominantly Pinus rigida and Quercus alba woodland. Both Massachusetts populations occur in Schizachyrium scoparium-dominated sites; associated species common to both sites include Arctostaphylos uva- ursi, Vaccinium angustifolium, Agrostis hyemalis, Aira praecox, Danthonia spicata, Festuca rubra, Carex emmonsii, Cerastium vulgatum, Gnaphalium obtusifolium, Helianthemum canadense, Hieracium venosum, Hypericum gentianoides, Polygala polygama, Potentilla canadensis, Spiranthes gracilis, S. tuberosa, Viola pedata, Aster paternus, Panicum sphaerocarpon, Aster linariifolius, and Aster undulatus (Sorrie 1985). MARYLAND: Agalinis acuta was first collected in Maryland in 1950 by Clyde F. Reed, although there are probably older specimens that are misidentified (Morse, pers. comm.). The species remains extant at Soldiers Delight, where it was noted in 1986 to to occur in an area of approximately 500 acres within a Pinus-Quercus woodland, associated with Schizachyrium scoparium, Aristida purpurea, Aristida dichotoma, Asclepias verticillata, Senecio smalli, Arabis lyrata, and Agalinis purpurea (Boone 1986). The 1987 field season revealed 500-1000 Agalinis acuta plants in an apparently stable population. The plants occurred as scattered groups of up to 100 plants each covering approximately 1/2 acre associated with openings of Schizachyrium scoparium within an open woodland of Pinus virginiana, Quercus marilandica, and Quercus stellata (Thompson, pers. comm.). Associated species populating the larger Soldiers Delight barren include Talinum teretifolium, Asclepias verticillata, Cerastium arvense, Phlox subulata, Oenothera fruticosa, Arabis lyrata, Polygala verticillata, Hypericum gentianoides, Viola pedata (Reed 1986). A second Maryland site, along a major highway, was last seen by Reed in 1965, and is now presumed extirpated (C. Ludwig, pers. comm.).

SOILS:

The soils supporting Agalinis acuta are nutrient-poor, usually acidic, and excessively drained. Two disjunct population centers inhabit very nutrient-poor habitats that are superficially somewhat similar, but the underlying bedrock and other ecological factors are quite different in the two regions. The more northerly coastal plain populations inhabit open sandy areas on moraines and sandplains, or glaciolacustrine terraces. Although the excessively drained sandplain soils are usually dry, the water table is often not far from the surface (Dowhan and Craig 1976). Further analysis of the soils at the two Massachusetts sites revealed acidic (pH 5.1) silty sandy soil which was low in macro nutrients and high in the trace metals aluminum, zinc, and iron (DiGregorio 1986).

The more southerly and disjunct Agalinis acuta population center inhabits a serpentine formation in Maryland. Soils are xeric and nutrient-poor, but are generally circumneutral, with a pH of 6-7 (Ludwig, pers. comm).

The following is a summary of the soil characteristics at Agalinis acuta populations in Massachusetts and New York (County Soil Surveys). For each site, five characteristics are mentioned: a) parent material, b) drainage, c) slope, d) surface soil, and e) subsoil.

Windsor (Massachusetts): a) deep sand deposits on terraces, b) excessively drained, c) 0-3%, d) loamy sand, loamy fine sand e) fine or medium sand.

Enfield (Massachusetts): a) silty mantle over stratified sand and gravel, b) well drained, c) 0-25%, d) very fine sandy loam and silt loam, e) stratified sands and gravel.

Hempstead (New York): a) outwash plains, silty mantle over stratified sand and gravel, b) excessively drained, c) 0-8%, d) black silt loam, e) gravelly loamy sand or stratified sand and gravel.

Plymouth (New York): a) sand plains, glacial outwash or lake laid materials, b) excessively drained, c)--, d) loamy sand, e) loamy sand or gravelly coarse sand.

Chrome (Maryland): a) serpentine bedrock, b) excessively drained, c) 0-15%, d)--, e) serpentine bedrock.

HABITAT SYNTHESIS: An underlying factor common to all sites supporting Agalinis acuta is the lack of competition from other species, imposed by one or both of the following conditions:

1. Extreme harshness of the habitat: soils at all sites are extremely nutrient poor. At site 6 on Long Island, the small openings of bare mineral soil were so harsh that the Schizachyrium scoparium and Chrysopsis mariana occupying these openings with Agalinis acuta were stunted (Rawinski 1983). Proximity to the ocean imposes an additional stress of salt spray. Adverse conditions are also noted to occur at the Maryland serpentine outcrops; vegetation is sparse in these low-nutrient soils. Low incidence of weedy exotics (Ludwig, pers. comm.) at a site with exposed mineral soil suggests that only those species tolerant of mineral toxicities and very low nutrients can survive.

2. Regular or sporadic disturbance. Some form of disturbance has been noted to occur at nearly every site supporting Agalinis acuta; at least occasional mowing takes place at both the Massachusetts cemetery populations and at the Long Island site 2 described above. Herbicide use has been known from Long Island sites 2 and 3; small fires, topsoil scarification, and other forms of human activity have occurred on all the Long Island sites except site 6, where use of a horse trail periodically exposes new soil for colonization. Most of the Agalinis plants occurring within the Maryland serpentine barren are also found on disturbed soils.

Fire has played a role in maintaining open habitat at a number of Agalinis acuta sites. Long Island site 4, located along a railroad track, as well as the surrounding pine barrens, have been subjected to occasional fire outbreaks caused by passing trains (Zaremba 1984). The open natural grassland supporting Long Island site 5 has probably been maintained in part due to fire (Reschke 1984).

The Maryland serpentine site at Soldiers Delight has been subjected to frequent fires; when the region was intensively mined for chromium, the area was occasionally burned over when fires were intentionally (and sometimes accidentally) set. Development in and around the Soldiers Delight region has also contributed to fires (Reed 1986). A study of forest age structure in these barrens has documented in tree rings the frequent disturbances caused by fire or drought (Knox 1984).

Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Major stewardship needs include: 1. Careful monitoring of extant populations. 2. Minimizing deleterious human disturbance (littering, dumping). 3. Educating landowners about appropriate management, and implementing management strategies. 4. Gathering information leading to an effective management plan for larger grassland populations.
Restoration Potential: Little is known about the potential of sites to recover from critical population declines or extreme disturbance. Most populations presently known have been discovered only within the past few years, and where intensive monitoring is being conducted, vast fluctuations in numbers have been noted (Lundgren 1983). The two Massachusetts sites have in the past two years seen notable population declines, and whether they can recover their previous numbers is not known at present.
Management Requirements: Most of the Agalinis acuta populations may require active management such as periodic burns, soil scarification, or mowing. Because of the critical rarity of this species, all efforts should be made to protect its habitat from development, and to institute an appropriate management regime. Attempts should be made to permanently protect all extant populations. Both Massachusetts sites are currently registered in the The Nature Conservancy Registry Program, and the only Maryland site is protected by the state government.

Maintenance of open habitat appears to be essential to most populations. Management approaches should be geared to the individual sites. For example, the two Massachusetts populations are located in cemeteries, and are already subject to mowing in the process of grounds maintenance. The land use and small area of these populations necessitates a management approach within these confines. On the other hand, some of the larger Long Island habitats, particularly remnant natural grasslands, may be suitable for prescribed burns, grazing, or soil scarification.

Area size and land use will dictate appropriate management procedures. Small populations already subjected to mowing should follow these guidelines: Mowing should not occur between July and October to ensure adequate seed production. If mowing must occur during this period in grounds maintenance, adjust the blade to the highest setting to avoid cutting the lowest axillary stems.

Large grassland populations of Agalinis acuta will require further research and monitoring before the most effective management approaches can be determined. The following tasks are offered by Reschke and Zaremba (1984):

1. Gather information on standard grassland management techniques (burning, mowing, removal of non-native species, collection and sowing of native plant seeds, and improving soil conditions).

2. Design a management plan including: a) small scale experimental burn plots and experimental mowing plots monitored and evaluated for effectiveness; b) development of policy for prescribed burns; c) plans for removal of weedy alien species.

Monitoring Requirements: Monitoring suggestions include the following (Sorrie and Swain 1987): 1. Annual population size and distribution, including number of flowers and capsules per plant. 2. Phenology of blooming. 3. Time of anthesis. In reference to the above monitoring needs, the following procedures are suggested: 1. Population counts should be conducted at least twice yearly: late August for peak flowering, and late September to note fruit-set. 2. In particular, watch for cessation of flowering. 3. Flowers are known to blossom mostly in the morning, but actual time of opening has not been observed. Check for a pattern.

Management Programs: Yearly population counts have been conducted since 1981 by the Massachusetts Natural Heritage Program and the Massachusetts/ Rhode Island Field Office of The Nature Conservancy. Active management of Massachusetts populations currently consists of attempting to restrict mowing during flowering and fruiting. These attempts have not been entirely successful, as mowing had occurred at a critical time in 1986. Miscommunication between owners and groundskeepers appears to have been the culprit. Bruce Sorrie and Caren Caljouw have met with all parties to avoid such communication problems in the future. In view of the notable population declines in recent years at these sites, more active management in the form of soil scarification should also be considered.

NEW YORK: a subpopulation of Long Island site 2 has declined from 150 in 1985, 2 in 1986, to 0 in 1987. Removal of some of the surrounding Japanese black pine and the accumulating pine needle duff was done in order to expose the soil. Intensive monitoring of all populations on Long Island has been in progress as a preparatory step to active management (Zaremba, pers. comm).

MARYLAND: the population in this state is being monitored, and active management has not been deemed necessary for the present.

Monitoring Programs: Annual monitoring of both Massachusetts populations has been pursued since their discovery. Coordinated by Caren Caljouw, Director of Stewardship (TNC, MA/RI Field Office, 294 Washington St., Boston, MA 02108) and Bruce Sorrie, Botanist (MA Natural Heritage Program, Division of Fisheries and Wildlife, 100 Cambridge St., Boston, MA 02202), monitoring was begun in 1983 by Julie Lundgren (University of Massachusetts, Morrill Science Center, Botany Department, Amherst, MA), and continued by Mario DiGregorio, Conservation Administrator (Town Hall, Brewster, MA 02631).

NEW YORK: Monitoring of Long Island populations has been conducted by Dr. Robert Zaremba, Botanist (The Nature Conservancy, New York Field Office, 1736 Western Avenue, Albany, NY 12203). Population counts are conducted annually at all sites. Additional baseline studies were conducted for all Long Island populations generating:

1. General community descriptions by noting percent cover of all species found. The community is defined by the area within 1 meter of outermost Agalinis acuta plants.

2. Lists of immediately associated species from line-intercept transects at each population.

MARYLAND: The population in Maryland is monitored annually by Daniel Boone, Ecologist, and Chris Ludwig, Botanist, both of the Maryland Heritage and Threatened/Endangered Species Project, Dept. of Natural Resources, C-3, Tawes State Office Bldg., Annapolis, MD 21401.

Management Research Programs: William Brumback, New England Wildflower Society, Framingham, Massachusetts, plans to continue germination experiments to determine growth requirements and role of parasitism in Agalinis acuta. In addition, he plans to begin and maintain a seed bank program using seeds from the Massachusetts and Maryland populations.

The following individuals are also involved in research and monitoring of Agalinis acuta: Dr. Judith Canne, Department of Botany and Genetics, University of Guelph, Guelph, ON Canada N1G2W1; and Sarah Davison, Director, The Nature Conservancy, South Fork/Shelter Island Chapter, PlO, Box JJJJ, East Hampton, NY 11937

Population/Occurrence Delineation Not yet assessed
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Population/Occurrence Viability
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Excellent Viability: Size: A population consisting of greater than 10,000 ramets and occupying greater than 2 acres (0.8 hectares) of habitat is recommended. Ideally the plants should be distributed in more than one discrete subpopulation. Because Agalinis acuta is an annual and numbers may fluctuate considerably from year to year, the size figures should be based on five consecutive years of survey data whenever possible.Condition: A very vigorous population that is relatively stable or increasing in size due to abundant sexual reproduction and quality of habitat. The healthy population should show minimal damage to ramets (<5% on average) from trampling, herbivory, seed predation by insects, disease or damage from weather related elements (e.g., drought, wind, salt spray). It should be growing in an open habitat comprised mainly of native species, such as those typically found in sandplain or serpentine barren habitats within the species range. Competition for light, water, or nutrients from other plant species should be low. Non-native species, especially invasive species, should be absent or, if present, at easily manageable levels. Landscape Context: The supporting landscape should have limited fragmentation and development. It should have the potential for expanding the population or metapopulation to a goal of 2 (0.8 ha) acres or greater of occupied habitat. Populations of invasive species that would easily invade and compete with Agalinis acuta should be at low or manageable levels in the immediate surroundings (up to ca. 0.5 km from the site in all direction).
Good Viability: Size: The population should consist of greater than 3000 ramets and occupy more than 1 acre (0.4 hectare) of habitat. Because Agalinis acuta is an annual and populations fluctuate considerably from year to year, the size figures should represent the average of five years of consecutive survey data whenever possible. Condition: The population should be vigorous with good annual seed production and on average only a low level of damage to ramets (< 10%) from trampling, herbivory, predation from insects, diseases, or damage due to weather related elements such as drought, wind, or saltspray. The habitat should be relatively free of non-native species, especially invasive species, with native species comprising greater than 75% of the ground cover. Landscape Context: The supporting landscape is one of some fragmentation or development, but with portions of it consisting primarily of natural vegetation for the local area. It should have the potential of expanding the population or metapopulation to a goal of >1 acre (0.4 hectare) of occupied habitat. Populations of invasive species that would easily invade and compete with Agalinis acuta should be at low or manageable levels in most of the immediate surroundings (up to ca. 0.2 km from the site in all directions).
Fair Viability: Size: The population consists of between 6 and 2999 ramets and occupies up to an acre (<0.4 ha) of habitat with at least 100 square meters of open, suitable habitat for the species to successfully flower and fruit annually. Because Agalinis acuta is an annual and populations fluctuate considerably from year to year, the size figures should represent the average of five consecutive years of data whenever possible. Condition: The plants should have at least fair vigor and show modest signs of damage to ramets (<20%) from trampling, herbivory, predation by insects, disease or weather related elements such as drought, wind or saltspray. The habitat may show signs of disturbance and the presence of many non-native plants, but natives should comprise > 50% of the cover. Landscape Context: The supporting landscape may have considerable fragmentation and development but should have the potential for expansion of suitable habitat for the population or metapopulation to a goal of 500 square meters to 1 acre (0.4 ha). Populations of invasive species that would invade and compete with Agalinis acuta may be abundant in portions of the immediate surroundings but still manageable as they invade the occupied habitat.
Poor Viability: Size: The population consists of < 6 individuals in a very small area of habitat (e.g., <100 square meters to about 0.2 ha). Because Agalinis acuta is an annual and numbers may fluctuate cnsiderably from year to year, the size figures should be based on five consecutive years of survey data whenever possible. Condition: The population is not vigorous and may have small plants producing few flowers or fruits. Plants may show considerable signs of damage to ramets (up to 20% or greater) from trampling, herbivory, predation by insects, disease, or damage from weather related elements such as drought, wind, or saltspray. The occupied habitat may show considerable signs of disturbance such as the presence of many non-native species, including ones that are considered invasive. Native plants may represent <50% cover in the habitat. Landscape Context: The supporting landscape has considerable fragmentation and development and has very limited natural vegetation or potential suitable habitat for the establishment of new subpopulations of the species (< 0.2 ha). Persistent encroachment by invasive plant species may represent a serious threat to the occupied habitat.
Justification: Typically, ramets and genets of Agalinis acuta are easy to distinguish in the field. Ramets, which nearly equal the number of genets in Agalinis acuta, represent the easiest units to count though. Rarely branching will occur at or below the soil level and result in confusion, so it is probably more accurate to say one is counting ramets. Being an annual, populations size can fluctuate due to poor seed production or germination, thus it is advisable to look at trends over at least a five year period. Population crashes have been observed at New York and Massachusetts sites due to insect predation on seed capsules, storm events and mismanagement (e.g., mowing at inappropriate times). Averages based on the population sizes and areas occupied over the prior five years are recommended to assess the true size of each population. A C/D threshold of only 6 individuals was chosen for a viable population size because it has been demonstrated at a number of the New England cemeteries that populations can be recovered from extremely low numbers where there is a small amount of suitable habitat and good management practices (e.g., mowing regimes) are applied consistently from year to year. Plant numbers at a cemetery on Cape Cod, for instance, stayed below 45 plants for 10 years and were even reported as one individual one of the years, and 8 another year, but rapidly rebounded to 3-4 thousand once mowing schedules were adjusted to allow flowering and fruiting to regularly occur. It has also been demonstrated that large populations can be achieved in a relatively short time period at a population in Falmouth, Mass., where seed was sown for the first time in 1994 and ten years later over 200,000 plants were estimated to be present in a 1-2 acre acre. This latter example along with the exemplary population in Maryland, helped set the parameters for an "A" Occurrence.<br><br>Threats to population are perceived as often related to competition or the lack of adequate management to maintain the early seral stage required to allow successful germination and survival of Agalinis. Population crashes from factors such as insect predation may be density dependent and are unlikely to totally eliminate a population. Landscape context seems less important for this species that size and condition. The fact that a large (100-200K) population can be maintained in 1-2 acres relatively high quality sandplain habitat amidst a somewhat developed residential environment on Cape Cod leads to the suggestion that the factors should be weighted approximately as follows: size (50%); condition (35%), and landscape context (15%)
Key for Ranking Species Element Occurrences Using the Generic Approach (2008).
Date: 19Jan2005
Author: P. Somers (2005)
U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 17Jan2005
NatureServe Conservation Status Factors Author: Caren Caljouw, Bruce Sorrie, Lesley Sneddon: Eastern Heritage Task Force, rev. Maybury/Somers (1996); Somers (2005)
Management Information Edition Date: 07Nov1988
Management Information Edition Author: Caren Caljouw, Bruce Sorrie, Lesley Sneddon.
Element Ecology & Life History Edition Date: 23Jun1992
Element Ecology & Life History Author(s): CAREN CALJOUW, BRUCE SORRIE, LESLEY SNEDDON: EASTERN HERITAGE TASK FORCE

Botanical data developed by NatureServe and its network of natural heritage programs (see Local Programs), The North Carolina Botanical Garden, and other contributors and cooperators (see Sources).

References
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Patterson, B.D., G. Ceballos, W. Sechrest, M.F. Tognelli, T. Brooks, L. Luna, P. Ortega, I. Salazar, and B.E. Young. 2003. Digital Distribution Maps of the Mammals of the Western Hemisphere, version 1.0. NatureServe, Arlington, Virginia, USA.

Acknowledgement Statement for Mammal Range Maps of North America:
"Data provided by NatureServe in collaboration with Bruce Patterson, Wes Sechrest, Marcelo Tognelli, Gerardo Ceballos, The Nature Conservancy-Migratory Bird Program, Conservation International-CABS, World Wildlife Fund-US, and Environment Canada-WILDSPACE."

Citation for Amphibian Range Maps of the Western Hemisphere:
IUCN, Conservation International, and NatureServe. 2004. Global Amphibian Assessment. IUCN, Conservation International, and NatureServe, Washington, DC and Arlington, Virginia, USA.

Acknowledgement Statement for Amphibian Range Maps of the Western Hemisphere:
"Data developed as part of the Global Amphibian Assessment and provided by IUCN-World Conservation Union, Conservation International and NatureServe."

NOTE: Full metadata for the Bird Range Maps of North America is available at:
http://www.natureserve.org/library/birdDistributionmapsmetadatav1.pdf.

Full metadata for the Mammal Range Maps of North America is available at:
http://www.natureserve.org/library/mammalsDistributionmetadatav1.pdf.

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