Geocarpon minimum - Mackenzie
Tiny Tim
Other English Common Names: Earthfruit
Other Common Names: tinytim
Synonym(s): Mononeuria minima (Mack.) Dillenb. & Kadereit
Taxonomic Status: Accepted
Related ITIS Name(s): Geocarpon minimum Mack. (TSN 195411)
Unique Identifier: ELEMENT_GLOBAL.2.153730
Element Code: PDCAR15010
Informal Taxonomy: Plants, Vascular - Flowering Plants - Pink Family
Image 11973

© Arkansas Natural Heritage Comission

 
Kingdom Phylum Class Order Family Genus
Plantae Anthophyta Dicotyledoneae Caryophyllales Caryophyllaceae Geocarpon
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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: Geocarpon minimum
Taxonomic Comments: Kartesz & Kartesz (1980) had placed this genus in Aizoaceae but Kartesz (1994 checklist) subsequently placed it in Caryophyllaceae.
Conservation Status
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NatureServe Status

Global Status: G2
Global Status Last Reviewed: 16Nov2006
Global Status Last Changed: 18Sep1990
Rounded Global Status: G2 - Imperiled
Reasons: Extant at about 40 sites in Missouri, Arkansas, Louisiana and Texas. Some sites are protected and appear to have good viability. Total population size is difficult to estimate.
Nation: United States
National Status: N2

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 Arkansas (S2), Louisiana (S2), Missouri (S2), Texas (S1)

Other Statuses

U.S. Endangered Species Act (USESA): LT: Listed threatened (16Jun1987)
U.S. Fish & Wildlife Service Lead Region: R4 - Southeast

NatureServe Global Conservation Status Factors

Range Extent Comments: Found in southwestern Missouri (Dade, Polk, Greene, and Lawrence Counties). Historically found in St. Clair & Jasper Counties, Missouri. Found in three southeastern counties in Arkansas (Cleveland, Drew, and Bradley) and one Northwestern County (Franklin). Also found at two locations in Louisiana (Wynn Parish). This species was discovered in Texas in 2004 in Anderson County (Keith et al. 2004).

Number of Occurrences: 21 - 80
Number of Occurrences Comments: Arkansas: 4 macrosites with a number of subpopulations. 3 sites in Louisiana. Missouri: twenty-two extant occurrences, one historical occurrence.

Population Size Comments: Fluctuates greatly so estimates are difficult.

Number of Occurrences with Good Viability/Integrity: Few to some (4-40)
Viability/Integrity Comments: There are at least 13 occurrences with good viability.

Overall Threat Impact: Very high - medium
Overall Threat Impact Comments: Changes in hydrology from fire line digging (creates a bearm along the fire plow line) is a threat as is very heavy ORV use. However, some disturbance is necessary. The salt slicks may have originally been herbivore salt source; trampling by by animals may create the necessary natural disturbance ((T. Witsell, pers. obs. 2006).

Short-term Trend: Decline of 10-30%
Short-term Trend Comments: It is declining at least to a certain extent due to threats such as development, resevoir creation and salt mining (pers. comm. E. Keith). Succession and suspected hydrologic changes have caused declines.

Long-term Trend: Decline of <30% to increase of 25%
Long-term Trend Comments: Although the long term trend of this species isn't documented, it is suspected that is has been stable (pers. comm. E. Keith).

Intrinsic Vulnerability Comments: Survives fairly harsh environmental conditions. Early-successional habitat held in stasis by soil composition. This annual species is also known to have good seed set, with lots of capsules (pers. comm. E. Keith).

Environmental Specificity: Very narrow. Specialist or community with key requirements scarce.
Environmental Specificity Comments: This species is a specialist only growing in salt praries of one kind. For example, in Texas this species is only known from one salt praire type and even though the other type is very similar, the Geocarpon is not found there. It is suspected that the salinity may be one factor driving the presence/absence of the species. Geocarpon is found on the saltier salt praire type (pers. comm. E. Keith).

Other NatureServe Conservation Status Information

Distribution
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Global Range: Found in southwestern Missouri (Dade, Polk, Greene, and Lawrence Counties). Historically found in St. Clair & Jasper Counties, Missouri. Found in three southeastern counties in Arkansas (Cleveland, Drew, and Bradley) and one Northwestern County (Franklin). Also found at two locations in Louisiana (Wynn Parish). This species was discovered in Texas in 2004 in Anderson County (Keith et al. 2004).

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 AR, LA, MO, TX

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AR Bradley (05011), Cleveland (05025), Drew (05043), Franklin (05047), Sebastian (05131)
LA Caddo (22017), De Soto (22031), Winn (22127)
MO Cedar (29039), Dade (29057), Greene (29077), Henry (29083), Jasper (29097)*, Lawrence (29109), Polk (29167), St. Clair (29185)
TX Anderson (48001), Harrison (48203), Panola (48365)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
08 Lower Saline (08040204)+, Castor (08040302)+
10 Harry S. Missouri (10290105)+, Sac (10290106)+, South Grand (10290108)+
11 Spring (11070207)+*, Frog-Mulberry (11110201)+, Dardanelle Reservoir (11110202)+, Bayou Pierre (11140206)+, Caddo Lake (11140306)+
12 Middle Sabine (12010002)+, Upper Neches (12020001)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: A small (1-4 cm tall), ephemeral, succulent winter annual that usually completes its life cycle within a 4-week period in the spring. Young plants are grayish; mature plants reddish-purple. Flowers are inconspicuous. This unusual species comprises the monotypic genus Geocarpon.
Technical Description: Diminutive, smooth, ephemeral annual from a small, slender taproot. Stems simple or branching only at very base, the branches few, at most 4, erect or spreading-ascending, mostly 3-4 cm high, less than 0.5 mm thick, terete, greenish-brown or strongly suffused with red; leaves oppposite, to 3-4 mm long, margins entire, apices acute; flowers usually 1/node and alternate, subsessile, erect, 3-4 mm long, bisexual, regular, funnelform-campanulate; sepals 5, 3-4 mm long, reddish or reddish-green; petals absent; stamens 5; staminodes 5; ovary superior, lance-ovoid, somwehat trigonous, about the length of the sepals, greenish, the narrow apex at anthesis 3-lobed, minutely glandular-toothed or retuse, the style lacking, the stigmas minute, recurved; fruit a capsule not much longer than the ovary, splitting along the 3 wirelike valve margins about halfway down the fruit from the apex to expose the numerous, long-funicular seed; seeds yellowish-green, transluscent, ovoid to nearly round, ca. 0.5 mm long, the rounded backs muriculate (minutely pebbled), the sides minutely and narrowly stirate-cancellate.
Diagnostic Characteristics: Geocarpon minimum is quite diminutive and has a succulent appearance. It is the only species of a monotypic genus. Originally placed in the family Aizoaceae, Palmer and Steyermark (1950) later transferred the genus to the Caryophyllaceae based on the following characters: staminodial rudiments, apetalous flowers, lack of stipules, gamophyllous calyx, 5 perigynous stamens, 1-celled ovary, and free-central placentation. Chemotaxonomic studies by Bogle et al. (1971) revealed the presence of anthocyanins, which provided further support for its placement in the Caryophyllaceae.
Duration: ANNUAL
Ecology Comments: Geocarpon minimum has been variously described as an annual (Tucker 1983, Palmer and Steyermark 1950) and as a winter annual (Mackenzie 1914, Bridges 1986, Morgan 1986), and this question has not been fully resolved to date. Small winter rosettes (1-2 mm wide) have been found on the Bona Glade site in Missouri during the month of November (Morgan 1986). Although winter rosettes have been carefully searched for in Arkansas, none have been found (Pittman pers. comm.). Shephard (1987) suggests that in Arkansas, Geocarpon exhibits characteristics of both annuals and winter annuals, with depauperate annual plants producing 1 flower each and more robust, branching winter annual plants producing several flowers each. The hypothesized 1 flowered annual plants predominate on Arkansas sites. The timing of emergence in the spring can vary widely from year to year with temperature having the greatest influence on initiation of growth.

Winter rosettes at Bona Glade begin producing flowering stems anywhere from March to mid April (Morgan 1986). Tucker (1983) lists March 23 to April 23 as flowering dates for Geocarpon in Arkansas; however, following a warm winter, Bridges (1986) found plants on the Warren Prairie site producing stems in late January, flowering by mid-February, and dead or dying by April 1. Although the initiation of growth in the spring is controlled by temperature, Morgan (1986) suggests that the vigor of plants and the number of flowers and seeds produced is probably dependent on soil moisture and temperature conditions experienced after the initiation of growth in the spring. Bridges (1986) reported a major deviation from typical phenological patterns due to unusual weather conditions in 1986. Following a warm winter, Geocarpon emerged early (late January) and began flowering by mid February. Subsequent dry weather conditions caused a decrease in growth and flower production until rainfall in mid March revitalized the population producing a peak of Geocarpon growth around March 20-23. Tucker (1983) states that the life cycle of individual plants is completed in a four week period, and Morgan (1986) indicates that a period of 4-6 weeks passes from the initiation of growth in spring to senescence. To date, no insect pollinators have been observed; however, Shephard (pers. comm.) has observed numerous small centipedes within Geocarpon populations and suggests that they be considered as possible pollinators. Flower production varies from plant to plant with some plants producing 5-7 stems with 2-4 flowers per stem and other plants being single stemmed with only 1-2 flowers produced (Morgan 1986). Years in which favorable conditions exist produce more robust plants with greater flower production. Shephard (pers. comm.) observed that even in apparently good years, a high percentage (ca. 90%) of the plants produced on the Warren Prairie site were one stemmed, one flowered individuals. Morgan (1986) collected 16 capsules and found from 9 to 72 seeds produced per capsule with a mean of 27 seeds per capsule. Seeds remain in dead capsules and are most likely dispersed near the parent plant by wind or rain or by plants simply falling over. Groups of seedlings have been observed growing from capsules produced the previous season (Morgan 1986). Tucker (1983) suggests that ants be considered as a possible secondary dispersal mechanism since ant mounds are often found in the vicinity of Geocarpon populations in Arkansas. Results of monitoring in Missouri (Morgan 1986) and Arkansas (Shephard 1987) indicate that Geocarpon appears in the same areas from year to year and has low vagility. As with many annual species, the size of Geocarpon populations varies greatly from year to year. Shephard (1987) indicates that 2033 plants were found within plots in 1986 (Warren Prairie, Arkansas) and 6761 plants were found within the same plots in 1987. Similarly, Morgan (1986) found 1900 plants in plots (Bona Glade, Missouri) in 1984 and 4055 plant present in the same plots 1986. Although it is apparent that seeds do survive in seed banks since plants are produced in years following poor seed production, little is known about seed viability and longevity. Geocarpon is very sensitive to competition from other species of vascular plants and occurs only in very open areas. Rettig (1983) found plants growing in the shade of other vascular plants but noted that they were less vigorous than plants observed on the same day growing in open areas. The role of disturbance in the maintenance of Geocarpon populations is not well understood. Rettig (1983) noted plants in lightly vegetated areas which he described as appearing to have been disturbed in the last year or two. In contrast, Bridges (1986) suggests that Geocarpon responds negatively to recent disturbance. He noted that recently disturbed areas supported greater growth of competitive species and a less vigorous growth of Geocarpon. Tucker (1983) suggests that slight disturbance of the soil surface is necessary for Geocarpon seed germination and seedling establishment. The role of disturbance in slick spot formation and maintenance is also not totally understood. It appears to be a factor in slick spot formation on those saline soils which have a deep subsurface horizon (i.e. Wing soil series) but slick spot formation on saline soils in which the subsurface horizon is close to the soil surface (i.e. Lafe soil series) does not appear to be dependent on erosional forces (Pittman 1988). More research needs to be done in this area. Bridges (1986) lists the following species as frequently associated with Geocarpon in Arkansas: Aristida spp., Oenothera linifolia, Plantago hybrida, Agrostis elliottiana, Hedyotis australis, Ambrosia bidentata, Hedeoma hispida, Coreopsis grandiflora, Hypericum drummondii, Talinum parviflorum, Dicanthelium sp., Scirpus koiolepis, Sagina decumbens, Centunculus minimus, Krigia occidentalis and Aster pratensis. Species frequently associated with Geocarpon in Missouri include: Plantago pusilla, Saxifraga texana, Selenia aurea and Talinum sp. (Morgan 1980).

Palustrine Habitat(s): Riparian
Terrestrial Habitat(s): Bare rock/talus/scree, Barrens, Grassland/herbaceous, Savanna
Habitat Comments: Sandstone glades and saline prairies. In Missouri, Geocarpon grows on moist, sandy soils on exposed sandstone outcrops or glades, where ledges of fine sandstone, interbedded with shale, are exposed along small streams. The surrounding area, where deeper soils prevail, is savannah. Sites in Arkansas and Louisiana are characterized by very thin soils that are high in sodium and magnesium. Woody plants are nearly absent. In these saline prairies, the species occurs mostly in very thinly vegetated, barren-like areas.

To date, Geocarpon minimum has been found on 13 sites in 7 counties in southwestern Missouri (Federal Register 1987) and on 4 sites in 4 counties in southern and western Arkansas (Franklin, Drew, Bradley, and Cleveland Counties) (Shephard pers. comm.). Geocarpon occurs in a distinctly different habitat in each of the two states. In Missouri, Geocarpon is found on sandstone glades which occur on "channel sand" formations of Pennsylvanian Age or near contact zones between Mississippian and Pennsylvanian rocks (Kurz, D. cited in Pittman 1988). Within the glades, Geocarpon is found in seasonally (winter and spring) wet, shallow depressions which contain a thin layer of sandy soil. Steyermark et al. (1959) described these depressions as being moss covered, with Geocarpon growing in areas where the moss cover is thin or broken. Morgan (1986) found Geocarpon most commonly growing on soils with a depth of 1-5 cm to bedrock. These thin soils support few other species, providing Geocarpon an environment relatively free of interspecific competition. During most years, Geocarpon is found on the thin soil present in the interior of these shallow depressions; however, in years in which environmental conditions are highly favorable for Geocarpon germination and development, the species can be found growing on deeper soils near the edges of these depressions in association with mosses, lichens and perennial vascular species (Morgan 1986).

The following description of Geocarpon habitat in Arkansas is condensed from Pittman (1988), which contains a thorough discussion of what is currently known of the habitat requirements of Geocarpon in Arkansas. To date, Geocarpon has not been found on any sandstone glades in Arkansas. Although the Ozark Highlands of Arkansas contain many sandstone glades which appear superficially similar to the Geocarpon supporting glades of Missouri, there are no known sandstone glades in Arkansas with the same mode of formation and chemical composition as the Missouri channel sand glades. Pittman notes that the Missouri channel sands have a relatively high concentration of magnesium, which is not the case for Arkansas sandstone glades. The presence of high concentrations of magnesium in the soil may be a critical factor in Geocarpon distribution, since the soils of Geocarpon sites in Arkansas are characteristically high in magnesium and/or sodium. All of the Geocarpon sites currently known from Arkansas occur on saline soil prairies on natric or saline soils. These soils possess a silt tom silt loam surface horizon underlain by a sodic subsurface horizon with a high concentration of sodium and magnesium salts. Physical and chemical properties of the subsurface layer include high runoff, low field capacity, low organic matter, high root toxicity, poor aeration, and low osmotic potential, producing soils characteristically poor for plant growth. Although there are six saline soil series in Arkansas, Geocarpon has been found only on the Lafe (Glossic Natrudalfs), Foley (Albic Glossic Natraqualfs), and Wing (Aquic Nastrustalfs) soil series to date, with the Lafe series supporting the best known populations. The vegetation of the saline soil "prairies" is maintained primarily by soil chemistry and depth to the subsurface horizon rather than by fire as in true prairies. Only the A-horizon of these soils is well suited for most root growth and the vegetation is distributed throughout the prairies according to the depth to the subsurface sodic horizon. This vegetation consists of low grasses and forbs with woody vegetation being confined to scattered low mounds which possess a deeper A-horizon. The result is a savanna-like community with scattered wooded areas interspersed with open "prairies". Geocarpon occurs only within the open areas.

Depth to the subsurface horizon is one of the most crucial factors in the distribution of Geocarpon. The closer the sodic horizon is to the surface, the more pronounced are the detrimental effects on vascular plant growth. In highly localized areas on the saline soils, the subsurface horizon can be extremely close to or at the soil surface producing areas virtually devoid of vegetation known as "slick spots". All of the Geocarpon found to date in Arkansas is associated with these slick spots on saline soils. Distribution of Geocarpon within the slick spots is highly dependent on the microtopography of the slick spots. Very few individuals grow in the harsh environment at the center of the slick spots and most are confined to the silt loam lip of the slick spot perimeter. The slick spot lip is characteristically covered by a layer of lichens, liverwort thalli, and moss protonemata and has been labeled the "cryptogamic lip" by Pittman. The soil here consists of a mixture of the surface horizon soil and the bodies of the lower plants. This mixture of lower plant bodies and mineral soil provides a moist, pliable substrate favorable for Geocarpon seed germination which is not present in the rest of the slick spot.

Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Protection: Identify and protect sites. Monitoring: Monitor the condition of known sites for signs of habitat destruction and population trends. Research: Continue research on biology and habitat requirements to determine if there are any management practices which could be implemented to better insure the survival of this species. Research on the biology of this species would increase the likelihood that management plans developed would be effective. A major threat to Geocarpon is the destruction or adverse modification of its habitat. In Missouri, some sites have been damaged by trampling and grazing by cattle. It has been suggested that physical disturbance may actually benefit Geocarpon at some sites by maintaining bare substrate for seedling establishment. A more serious threat is from pasture improvement with the subsequent invasion by prairie species. ORVs have also damaged Geocarpon habitat. In southern Arkansas much of the habitat suitable for Geocarpon has been heavily disturbed by silviculture, pasture, agriculture, and road expansion.
Restoration Potential: Recovery of Geocarpon to sites on which it has been extirpated appears unlikely at this time. Results of monitoring in Missouri (Morgan 1986)and Arkansas (Shephard pers. comm.) indicate that populations on suitable habitat are maintaining themselves well. Extirpation of Geocarpon from sites appears to be due to habitat destruction and once this has occurred it is unlikely that managers will be able to recreate suitable conditions and reintroduce the species.

In addition, natural recolonization of Geocarpon to areas of suitable habitat is not likely due to the low vagility of the species. Perhaps as more is learned of the specific requirements necessary for germination and establishment of this species attempts can be made to introduce or reintroduce this species to areas which appear suitable.

Preserve Selection & Design Considerations: On glade sites, it is important to protect all land up-slope from the glade to guard against siltation or pollution of the site from up-slope grazing or development.
Monitoring Requirements: So as a minimum, the overall quality of Geocarpon sites needs to be monitored on an annual basis to determine if the site is being detrimentally affected by grazing, off-road vehicle traffic, or any other destructive activity. Population trends and long term successional changes on Geocarpon sites are additional monitoring concerns. The monitoring of long term successional changes is of special concern on Arkansas sites where the role of disturbance in the maintenance of Geocarpon habitat is not fully understood.

Overall habitat quality of sites can be monitored by simple visual inspection of the site or by photo monitoring. Population trends and long term successional trends are best monitored by the establishment of plots along permanent transects which run through known Geocarpon populations. Information recorded should include the number, location, and vigor of individual Geocarpon plants within the plots and species associated with Geocarpon.

Monitoring Programs: Population trends at a site in Missouri were actively monitored from 1983 to 1985 using a system of permanent transects and plots. Contact: Don Kurz, Missouri Department of Conservation, P.O. Box 180, Jefferson City, Missouri, 65102-0180.

A site in Arkansas is currently being monitored with the information outlined under Biological Monitoring Procedures being collected. Contact: Bill Shephard, Arkansas Natural Heritage Commission, Suite 200, Heritage Center, 225 East Markham, Little Rock, AR 72201

Management Research Needs: Research on seed germination is currently being conducted by the Nebraska Statewide Arboretum and at Southwest Missouri State University.

Contacts: Nebraska Statewide Arboretum, 112 FSL, University of Nebraska, Lincoln, NE 68583-0283. (402)472-2971. Dr. Wallace R. Weber, Dept. of Life Sciences, Southwest Missouri State University, Springfield, MO 65802. (417)836-5883.

Population/Occurrence Delineation Not yet assessed
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Population/Occurrence Viability
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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: 20Nov2006
NatureServe Conservation Status Factors Author: T. Phillips, C. Reid, J. Singhurst, and T. Witsell
Management Information Edition Date: 26Jun1989
Management Information Edition Author: ROBERT F. STEINAUER
Element Ecology & Life History Edition Date: 04May1990
Element Ecology & Life History Author(s): STEINAUER, ROBERT F.

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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  • Behnke, H.D. 1982. Geocarpon minimum: Sieve-element plastids as additional evidence for its inclusion in the Caryophyllaceae. Taxon 3(1): 45-47.

  • Bogle, A.L., T. Swain, R.D. Thomas, and E.D. Kohn. 1971. Geocarpon: Aizoaceae or Caryophyllaceae? Taxon 20(4):473-477.

  • Bridges, E. L. 1986. Population Inventory and Monitoring of Geocarpon minimum at Warren Prairie Natural Area. Arkansas Natural Heritage Commission, Little Rock, Arkansas.

  • Dillenberger, M.S., and J.W.  Kadereit 2014. Maximum polyphyly: Multiple origins and delimitation with plesiomorphic characters require a new circumscription of Minuartia (Caryophyllaceae). Taxon 63(1): 64-88.

  • 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.

  • Keith, E. L., J. Singhurst and S. Cook. 2004. Geocarpon minimum (Carophyllaceae) new to Texas. Sida 21(2): 1165-1169.

  • Kral, R. 1983c. A report on some rare, threatened, or endangered forest-related vascular plants of the South. U.S. Dept. of Agriculture Forest Service Technical Publication R8-TP2, Athens, GA. 1305 pp.

  • Mackenzie, K.K. 1914. A new genus from Missouri. Torreya 14:67

  • McInnis, N.C., L.M. Smith, and A.B. Pittman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisiana. Phytologia 75(2):159-162.

  • Morgan, S. 1980. Status report on Geocarpon minimum in Missouri. Missouri Department of Conservation, Jefferson City, Missouri. 16 pp.

  • Morgan, S. 1986. A study of a population of Geocarpon minimum in Missouri. Missouri Department of Conservation, Jefferson City, Missouri.

  • Orzell, S. L., and E. L. Bridges. 1987. Further additions and noteworthy collections in the flora of Arkansas, with historical, ecological, and phytogeographical notes. Phytologia 64(2):81-144.

  • Palmer, E.J., and J. Steyermark. 1950. Notes on Geocarpon minimum MacKenzie. Bull. Torrey Bot. Club 77:266-273.

  • Pittman, A. B. 1988. Identification, survey and evaluation of potential habitats of Geocarpon minimum MacKenzie in Arkansas. Arkansas Natural Heritage Commission, Little Rock, Arkansas.

  • Poole, Jackie M., W. R. Carr, D. M. Price, and J. R. Singhurst. 2007. Rare plants of Texas. Texas A&M University Press, College Station. 640 pp.

  • Rettig, J.H. 1983. A New Arkansas station for Geocarpon minimum MacKenzie (Caryophyllaceae). Bull. Torr. Bot. Club. 110(2):213.

  • Shephard, W. 1987. Monitoring of Geocarpon minimum at Warren Prairie Natural Area in the Spring of 1987. Arkansas Natural Heritage Commission, Little Rock, Arkansas.

  • Steyermark, J., J.W. Voigt, and R.H. Mohlenbrock. 1959. Present biological status of Geocarpon minimum MacKenzie. Bull. Torrey Bot. Club 86: 228-235.

  • Steyermark, J.A. 1963. Flora of Missouri. Iowa State Univ. Press, Ames. 1728 pp.

  • Thurman, C.M. 1989. Final Report. A Missouri survey of six species of federal concern. Missouri Dept. of Conservation. 99 pp.

  • U.S. Fish and Wildlife Service (USFWS). 1987. Endangered and Threatened Wildlife and Plants; Threatened Status for Geocarpon minimum. Federal Register 52(115):22930-22933.

  • U.S. Fish and Wildlife Service. 1987. Threatened status for Geocarpon minimum. Federal Register 52(115): 22930-22933.

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