An annotated checklist of the liverworts and hornworts of the Delmarva Peninsula

1,2 William A. McAvoy Delaware Natural Heritage and Endangered Species Program, Delaware Division of Fish and Wildlife, 4876 Haypoint Landing Rd., Smyrna, DE 19953

Wesley M. Knapp Wildlife and Heritage Services, Maryland Dept. of Natural Resources, PO Box 68, Wye Mills, MD 21679

Lance T. Biechele 14011 Cooley Rd., Princess Anne, MD 21853

Oldest Botanical Journal in the Western Hemisphere

Journal of the Torrey Botanical Society 138(2), 2011, pp. 239–251

An annotated checklist of the liverworts and hornworts of the Delmarva Peninsula

1,2 William A. McAvoy Delaware Natural Heritage and Endangered Species Program, Delaware Division of Fish and Wildlife, 4876 Haypoint Landing Rd., Smyrna, DE 19953

Wesley M. Knapp Wildlife and Heritage Service, Maryland Dept. of Natural Resources, PO Box 68, Wye Mills, MD 21679

Lance T. Biechele 14011 Cooley Rd., Princess Anne, MD 21853

MCAVOY, W. A. (Delaware Natural Heritage and Endangered Species Program, Delaware Division of Fish and Wildlife, 4876 Haypoint Landing Rd., Smyrna, DE 19953), W. M. KNAPP (Wildlife and Heritage Services, Maryland Dept. of Natural Resources, PO Box 68, Wye Mills, MD 21679), AND L. BIECHELE (14011 Cooley Rd, Princess Anne, MD 21853). An annotated checklist and atlas of the liverworts and hornworts of the Delmarva Peninsula. J. Torrey Bot. Soc. 138: 239–251. 2011.—In order to increase the understanding of the diversity, distribution and abundance of liverworts and hornworts on the Atlantic Coastal Plain, a six year study (2004–2009) was conducted focusing on the Delmarva Peninsula (a land mass comprising portions of the states of Delaware, Maryland, and Virginia). Our previous knowledge of the liverworts and hornworts of the Delmarva were minimal. Collections were few and usually localized, and published and unpublished reports were scarce. Results from this study are presented in the format of an annotated checklist and represents the first comprehensive study on the liverworts and hornworts of the Delmarva Peninsula. A total of 65 species, subspecies, and varieties were documented during this study with 22 being reported as new to the bryoflora of the Delmarva. Key words: bryophytes, checklist, Delmarva Peninsula, hornworts, liver worts.

Of all the land plants, bryophytes, particu- research are presented here in the format of larly liverworts and hornworts, are one of the an annotated checklist. most understudied groups. In an effort to Prior to this study, earlier investigations and expand the overall knowledge of the diversity, collections of liverworts and hornworts on the distribution, and abundance of liverworts and Delmarva Peninsula were limited and usually hornworts of the Atlantic Coastal Plain local in terms of the areas studied or collected. physiographic province, research focusing on The earliest collections were made by Albert the Delmarva Peninsula was conducted from Commons in the 1890s and some of his 2004 to 2009. Research involved field surveys, collections, many of which are housed at the as well as herbaria and literature searches to Academy of Natural Sciences (PH, herbarium document the liverwort and hornwort flora of acronyms follow Index Herbariorum 2010) in the Delmarva. Data resulting from this Philadelphia represent significant discoveries for the Peninsula, such as Jungermannia gracillima, Ptilidium pulcherrimum, and Riccia 1 We thank Dr. Raymond Stotler for his time in membranacea. All three of these species are helping to confirm identifications and for providing considered historical on the Delmarva, with insightful comments and suggestions that helped to the only known collections being those made improve the quality of this manuscript. We greatly appreciate the time and efforts of Carl Darigo who by Commons. Edward Uebel (2000) published reviewed this manuscript and provided comments. an annotated list of the bryophytes collected in We are also thankful to Bruce Allen, Charlie Davis, Maryland by Dr. Elmer Worthley. Primarily and Steven Timme and Ann Wheeler for providing during the 1960’s, Worthley collected 12 assistance related to various aspects of this study. We thank the following curators who helped arrange species of liverworts (10 leafy, 2 thalloid) from loans and provided access to specimens: Arthur the Maryland Eastern Shore counties of the Tucker (DOV), Alina Freire-Fierro (PH) and Rusty Delmarva, one of which includes Blepharos- Russell (US). toma trichophyllum (Uebel 2000). This species is 2 Author of correspondence. E-mail william. now considered to be historical on the [email protected] Received for publication November 8, 2010, and in Peninsula, with the only collection being made revised form February 1, 2011. by Worthley. While a student at Salisbury

239 240 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 138

University (previously Salisbury State Col- The Peninsula lies south of the fall line (a lege), B. L. Davis (1976) produced an unpub- term applied to the boundary between the lished report titled: A Survey of the Bryophytes Appalachian Piedmont province and the of Wicomico County, Maryland. Davis listed Atlantic Coastal Plain) in New Castle County, 10 species of liverworts (7 leafy, 3 thalloid) Delaware and Cecil County, Maryland, and is based on a single season of field work within bordered on the east by the Delaware River, 18 acres near her home, and at ‘‘a dozen other Delaware Bay and the Atlantic Ocean, and on areas of the county’’ where ‘‘spot checks’’ were the west by the Elk River and Chesapeake done (Davis 1976). Though Davis’s report was Bay. It includes the Coastal Plain province of the result of an undergraduate project and Delaware (New Castle, Kent, and Sussex unpublished, it is included in this study in Counties), the Eastern Shore of Maryland order to be as thorough and comprehensive as (Cecil, Kent, Queen Anne’s, Caroline, Talbot, possible and to alleviate possible questions Dorchester, Wicomico, Somerset, and from future researchers. In 1991, Lance Worcester Counties), and the Eastern Shore Biechele began collecting liverworts and horn- of Virginia (Accomack and Northampton worts on the Delmarva, mostly in the south- Counties). Its length north to south is about west corner of the Peninsula (Somerset, 200 miles (320 km), its greatest width is about Wicomico and Worcester Counties), and 70 miles (110 km), its narrowest width is about Lance’s efforts were the impetus for this study. 10 miles (16 km), and the total land area is William McAvoy (1999) published the results of about 5,800 mi2 (15,000 km2). The climate of a bryophyte study of the Great Cypress the Peninsula is moderated by the Delaware Swamp Conservation Area in Sussex Co., Bay, Chesapeake Bay, and the Atlantic Ocean Delaware and Worcester Co., Maryland, and is characterized by cool winters and warm where he listed 12 species of liverworts (10 humid summers. The landscape of the Del- leafy, 2 thalloid). While doing bryological marva is mostly rural, on flat to gentle sloping studies on Assateague Island National Sea- sandy plains with slow-flowing rivers and shore in Worcester Co., Maryland, Shanda streams that are bordered by extensive swamp King (2000) reported in an unpublished forests and tidal marshes. In the coastal areas, document 6 species of liverworts (5 leafy, 1 barrier islands, salt marshes, tidal flats and thalloid). While doing field surveys related to inland bays are well developed. The Delmar- this paper, McAvoy, Biechele and Knapp va’s Coastal Plain soils of sands, silts, clays (2006) published their discovery of Calypogeia and gravel support forests primarily composed peruviana, which was new to the Delmarva of mixed evergreen; Pinus taeda L. (loblolly and a northern range extension. pine), Ilex opaca Aiton (American holly), and Results presented here mark the first focused deciduous tree species; Quercus spp. (oaks), and comprehensive study on the liverworts and Carya spp. (hickories), Fagus grandifolia Ehrh. hornworts of the Delmarva Peninsula. (American beech), Nyssa sylvatica Marsh. (black gum), Liquidambar styraciflua L. (sweet The Study Area. The Delmarva Peninsula gum), Acer rubrum L. (red maple), and (Fig. 1) has historically been defined as Liriodendron tulipifera L. (tulip poplar). The including the entire state of Delaware (both Delmarva Peninsula lies within the Chesa- the Piedmont and Coastal Plain physiographic peake Bay Lowlands Ecoregion as defined by provinces of New Castle Co.), the entirety of The Nature Conservancy (2002), and within the Eastern Shore of Maryland and both the the Outer Coastal Plain Mixed Forest Prov- Piedmont and Western Shore regions of Cecil ince as mapped by Bailey (1995). Co., Maryland, and the Eastern Shore of Virginia (Tatnall 1946). This definition, like so Methods and Materials. The study area (the many other geopolitical boundaries, is prob- Delmarva Peninsula of the Atlantic Coastal lematic from a natural history perspective Plain, United States) was regularly sampled because it includes very different physiograph- ic from 2004–2009 (usually from April to Octo- provinces. For the purposes of this study the ber) resulting in surveys that captured all Delmarva Peninsula will be referred to as an habitat types known from the Delmarva area located entirely within the Atlantic Peninsula. Surveys were conducted in all Coastal Plain physiographic province, thus habitat types (natural and anthropogenic) to creating a more ‘‘natural’’ unit of study. capture the full diversity of Liverworts and 2011] MCAVOY ET AL.: LIVERWORTS AND HORNWORTS 241

FIG. 1. Location map, Delmarva Peninsula. 242 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 138

Table 1. Summation of the Live rwort and Table 2. Families represented on the Delmarva Hornwort flora of the Delmarva Peninsula. Peninsula with the greatest number of taxa.

Totals Family Genera Taxa

Taxa 65 Cephaloziaceae 3 7 Species 56 Lejeuneaceae 3 7 Subspecies and Varieties 9 Ricciaceae 2 6 Native Taxa 64 Lophocoleaceae 2 4 Non-native Taxa 1 Frullaniaceae 1 5 Families 28 Calypogeiaceae 1 4 Genera 39 Leafy Liverworts 45 Thalloid Liverworts 18 (4 taxa). The largest genera include: Frullania Hornworts 2 Rare Taxa 22 and Riccia both with 5 taxa, and Calypogeia Southern Taxa 8 and Cephalozia both with 4. The majority of Northern Taxa 1 the taxa are leafy liverworts with 45, followed

by thalloid liverworts with 18, and hornworts with two (Table 1). There are 22 taxa consid- Hornworts. The seasonality of some taxa ered to be rare on the Delmarva, or 35% of the required multiple surveys over the course of flora (Table 3), and of that total, 5 are the study. Voucher specimens were made for historical (not documented for 20 or more all new county and Delmarva records. Collec- years, Table 3). The Delmarva county sup- tions made by the first and second authors are porting the highest number of taxa is Worces- deposited at the Claude E. Phillips Herbarium ter, with 46 (Table 4). The Delmarva county (DOV), Delaware State University, Dover, supporting the lowest number of taxa is Cecil, Delaware and the third author deposited with 13 (Table 4). Four species were docu- specimens at the Charles O. Wingo Jr. Herbar- mented from all 14 Delmarva counties: ium, Salisb ury University, Salisbury, Maryland. Lophocolea heterophylla, Odontoschisma pros- A variety of manuals were consulted for this tratum, Pallavicinia lyellii, and Scapania ne- project because no single treatment contained morea. Eight taxa occurring on the Delmarva all the species encountered. The following are more southern in their geographic distri-

manuals were referred to: Ammons (1940), Schuster (1969, 1974, 1980, 1992), Breil (1970, Table 3. Taxa considered to be rare (known 1996), Crum (1991), Hicks (1993), Lincoln from 10 or fewer populations) on the Delmarva (2008). Questionable identifications, or diffi- Peninsula. Those marked with an * are considered to cult to determine collections were sent to Dr. be historical (not documented for 20 or more years).

Raymond Stotler (Southern Illinois Universi- Taxon ty) for confirmation. A literature review was conducted in order to compile a list of Blepharostoma trichophyllum* Cephalozia macrostachya Chiloscyphus materials that may have included Delmarva polyanthus var. rivularis Cololejeunea liverworts and hornworts. Specimen searches cardiocarpa* Conocephalum conicum were done at various herbaria (PH, DOV, US, Gymnocolea inflata NY, DUKE) to confirm the identification of Jungermannia gracillima* Lejeunea maxonii literature reports and to obtain additional Leucolejeunea conchifolia county and Delmarva records. Leucolejeunea unciloba Lophocolea bidentata Results. Collectively, the liverworts and Lophocolea minor Lophozia bicrenata hornworts of the Delmarva Peninsula consists Lophozia capitata of 65 taxa (species, subspecies and varieties), Porella platyphylla which includes 64 native taxa and one non- Ptilidium pulcherrimum* native taxon, and is represented by 28 families Radula complanata and 39 genera (Table 1). The families with the Radula obconica Reboulia hemisphaerica largest number of taxa are (Table 2): Cepha- Riccia membranacea* loziaceae (7 taxa), Lejeuneaceae (7 taxa), Ricciocarpos natans Ricciaceae (6 taxa), Frullaniaceae (5 taxa), Trichocolea tomentella Calypogeiaceae (4 taxa) and Lophocoleaceae 2011] MCAVOY ET AL.: LIVERWORTS AND HORNWORTS 243

Table 4. Total number of taxa documented by Table 6. Taxa documented and reported as new county on th e Delmarva Peninsula (highest to the Liverwort and Hornwort flora of the to lowest). Delmarva Peninsula.

County Taxa Taxon

Worcester 46 Calypogeia fissa subsp. neogaea Sussex 45 Calypogeia peruviana (McAvoy, Biechele and Wicomico 32 Knapp 2006) Accomack 27 Cephaloziella byssacea var. byssacea Queen Anne’s 26 Cephaloziella rubella var. rubella New Castle 27 Conocephalum conicum Caroline 26 Frullania eboracensis subsp. virginica Northampton 25 Frullania ericoides Dorchester 24 Gymnocolea inflata Kent (DE) 23 Kurzia sylvatica Somerset 22 Lejeunea maxonii Talbot 20 Leucolejeunea conchifolia Kent (MD) 17 Lophocolea bidentata C ecil 13 Lophozia bicrenata Lophozia capitata Lunularia cruciata bution and are at or near the northern limits of Metzgeria furcata their range on the Peninsula (Table 5). Only Notothylas orbicularis one species has a more northern distribution Radula complanata and is at or near the southern limits of its Radula obconica Riccia bifurca range on the Delmarva (Lophocolea minor). Riccia hirta Twenty-two taxa documented during this Ricciocarpos natans study are reported here for the first time (Table 6). Calypogeia trichomanis (L.) Corda (Calypo- Excluded Species. The following species geiaceae). Calypogeia trichomanis is a northern have been excluded from the liverwort and species (Canada and New England), and was hornwort flora of the Delmarva Peninsula. cited for Worcester Co., Maryland by Worth- Blasia pusilla L. (Blasiaceae). This species ley (Uebel 2000) and for Wicomico Co., was cited for Wicomico Co., Maryland by Maryland by Davis (1976). The Worthley Davis (Davis 1976), but no specimens for this collection (1967, MO) has been annotated as species from Wicomico Co. could be found at Calypogeia muelleriana (pers. comm., Carl MO (pers. comm., Carl Darigo, Missouri Darigo, Missouri Botanical Garden, 3 April Botanical Garden, 3 April 2008). Though 2008). Concerning the Wicomico Co. collec- Blasia pusilla is not improbable for the tion, if Davis made a specimen, it would have Delmarva, the literature (Breil 1996, Hicks been deposited at MO (pers. comm., Dr. 1992, Lincoln 2008, Manville 2001, Plitt 1908, Ronald A. Pursell, Penn State University, 9 Schuster 1992) suggests that it may be more July 2004). The collection at MO has been northern in its range and more frequent in the searched and no specimens of this species from Piedmont and Mountain provinces then on the Wicomico Co., Maryland could be found Coastal Plain. (pers. comm., Carl Darigo, Missouri Botanical Garden, 3 April 2008). The Davis citation is Table 5. Taxa documented from the Delmarva likely C. muelleriana. Peninsula with a more southe rn distribution that are Cephalozia bicuspidata (L.) Dumort. (Ce- at or near their northern limits of natural dis tribut ion. phaloziaceae). According to Schuster’s Vol. III (1974), this species is essentially northern in its Taxon distribution, becoming rare farther south. This Calypogeia peruviana species was cited for Wicomico Co., Maryland Cololejeunea cardiocarpa by Davis (Davis 1976), but no specimens for Cololejeunea minutissima subsp. minutissima Lejeunea maxonii this species from Wicomico Co. could be Leucolejeunea conchifolia found at MO (pers. comm., Carl Darigo, Leucolejeunea unciloba Missouri Botanical Garden, 3 April 2008). Sphaerocarpos texanus Lophozia laxa (Lindb.) Grolle [syn. 5 L. T elaranea nematodes marchica (Nees) Steph., (Scapaniaceae)]. This 244 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 138 species was cited for Delaware [unknown if based on Schuster (1969, 1974, 1980, and Piedmont or Coastal Plain (i.e., Delmarva)] by 1992) and Hicks (1992). Evans and Nichols (1908). Shuster in Vol. II (1969) makes note of this citation and points DELMARVA HABITAT . Brief habitat descrip- out that this species was ‘‘Reported from tions refer to the habitat preferences of a taxon Delaware and West Virginia (Evans & Nichols on the Delmarva Peninsula, and are based on 1908). It is highly unlikely that either of these the collection record, the literature, and the reports represents this northern species, which field experience of the authors. is almost strictly confined to areas which ADDITIONAL INFORMATION. Comments are underwent Pleistocene glaciation.’’ No speci- varied and are meant to provide useful men can be found which supports the claim of information related to a taxon. Evans and Nichols (1908). If a taxon is listed for a state and county, then it has been vouchered with a modern day Checklist Format and Arrangement. For the (within the last 20 years) collection by the most part, nomenclature follows Stotler & authors, unless otherwise noted [e.g., A. Crandall-Stotler (1977). The checklist begins Commons (collector’s name), ## (collection with Phylum Marchantiophyta, the liverworts number), 1892 (year of collection), PH (her- and concludes with Phylum Anthocerotophyta, barium where deposited)]. A county record the hornworts. Within each Phyllum, arrange- has been vouchered by at least one collection, ment is by Class and then alphabetically by but in most cases several collections exist, both Family, Genus, and Species. The scientific name for each species includes author(s), modern day and historical. Since the majority followed by synonyms where appropriate. of county voucher specimens were collected by Author abbreviations follow Brummitt and the authors, collection data (year, number and Powell (1992). In addition, for each taxon listed herbarium) are not listed in order to save the following attributes are also provided. space. Collections made by the first two authors are deposited at the Claude E. Phillips STATUS. The status of a taxon refers to its Herbarium (DOV), Delaware State Universi- relative rarity on the Delmarva Peninsula and ty, Dover, Delaware. Collections made by the is based on the collection record, the literature, third author are deposited at the Charles O. and the field experience of the authors. Wingo Jr. Herbarium, Salisbury University, However, the level of rarity is applied with a Salisbury, Maryland. Specimens pertaining to a certain level of uncertainty. Continued field specific county of occurrence that were studies in the future will undoubtedly provide collected by the third author are distinguished new data that will add to our overall as (SUHC), following the county name. If a knowledge of the distribution and abundance taxon has been reported from a county based of liverworts and hornworts on the Delmarva. on the literature and no voucher specimen has The following terms are used to describe the been seen by the authors, then the literature status of a taxon on the Delmarva: Rare citation is given (e.g., Davis 1976), followed by (known from only 10 or fewer populations, or nss, (no specimen seen). For example, Md.: habitat is limited), Historical (has not been Wicomico, (Davis 1976, nss). reported for 20 or more years), Uncommon (evenly distributed, but not plentiful), Com- An Annotated Checklist of the Liverworts and mon (plentiful throughout). Hornworts of the Delmarva Peninsula DISTRIBUTION. Listed first by state [Del. Phylum Marchantiophyta (Liverworts) Class (Delaware), Md. (Maryland), and Va. (Vir- ginia)], then by the counties within each state Marchantiopsida (Complex Thalloid going from north to south (e.g., Delmarva Liverworts)

Status and Distribution: Del.: New Castle; Aytoniaceae Kent; Sussex. Md.: Cecil; Kent; Queen Anne’s; Caroline; Talbot; Dorchester; Wicomico; Reboulia hemisphaerica (L.) Raddi Somerset; Worcester. Va.: Accomack; North- Delmarva Status and Distribution: Rare. ampton). For some entries, it is useful to Md.: Kent. provide the overall North American distribu- Delmarva Habitat: Calcareous marl and tion and in those cases, distributional data are shell-bearing soils. 2011] MCAVOY ET AL.: LIVERWORTS AND HORNWORTS 245

Conocephalaceae wetland depressions, sandpits and fresh water impoundments. Conocephalum conicum (L.) Dumort. Additional Information: The aquatic Riccia Delmarva Status and Distribution: Rare. fluitans is nearly always sterile, but plants Md.: Queen Anne’s. stranded on mud may produce fertile capsules. Delmarva Habitat: Forms dense patches on These fertile populations are possibly a distinct organic substrates and gravel in groundwater species, Riccia canaliculata Hoffm. (Ammons seepage slope wetlands. 1940, Breil 1970, Crum 1991). Lunulariaceae Riccia hirta (Aust.) Underw. Delmarva Status and Distribution: Uncom- Lunularia cruciata (L.) Dumort. mon. Del.: New Castle; Sussex. Md.: Somerset. Delmarva Status and Distribution: Non- Delmarva Habitat: Develops rosettes on native, a native of Europe. Currently vouch- bare mineral soils in moist poorly drained ered only from Accomack Co., Va. (SUHC), corners of agricultural fields. but widespread throughout North America Riccia huebeneriana Lindenb. subsp. sullivantii and is probably found in most greenhouses on (Austin) R.M. Schust. Riccia sullivantii Austin the Delmarva. Delmarva Status and Distribution: Common. Delmarva Habitat: Not naturalized, but Del.: New Castle; Kent, Sussex. Md.: Kent; forms patches on soil in pots and trays in Dorchester; Somerset; Worcester. nurseries and greenhouses. Delmarva Habitat: Develops rosettes on

Marchantiaceae bare mineral soils in poorly drained corners of agricultural fields, and also appears in Marchantia polymorpha L. Coastal Plain seasonal ponds after draw- Delmarva Status and Distribution: Uncom- down. mon. Del.: New Castle. Md.: Dorchester, Riccia membranacea Gottsche et. Lindb. Wicomico (SUHC). Delmarva Status and Distribution: Species is Delmarva Habitat: Forms patches on bare historical on the Delmarva, known only from mineral soil, on stream banks and rivers, both the two collections cited. Del.: Sussex (A. tidal and non-tidal, and on disturbed ground. Commons, s.n., 1892, PH). Va.: Northampton (E. Wherry, s.n., 1925, PH). Ricciaceae Delmarva Habitat: Bare, poorly drained Riccia bifurca Hoffm. mineral soils. Delmarva Status and Distribution: Uncom- Ricciocarpos natans (L.) Corda mon. Del.: New Castle. Md.: Queen Anne’s; Delmarva Status and Distribution: Rare. Caroline; Worcester. Del.: New Castle. Delmarva Habitat: Develops rosettes on Delmarva Habitat: Coastal Plain seasonal bare mineral soils in moist to poorly drained ponds. corners of agricultural fields, and also appears Additional Information: This species takes in Coastal Plain seasonal ponds after draw- on two forms: aquatic and terrestrial. The down. aquatic form is a heart or fan-shaped thallus Additional Information: This species, as well with long, purplish-black scales on the under- as R. hirta, and R. huebeneriana subsp. side. The thallus floats on the surface of the sullivantii are characteristically found in sum- water in spring when flooding occurs, in mer on soils that have been flooded in the habitats such as seasonally flooded wetland spring. Plants are seasonal and appear to grow depressions (known as Coastal Plain seasonal annually from water-borne or soil-borne ponds). The aquatic form produces sex organs, spores. and spores mature by mid-summer. As the Riccia fluitans L. season progresses and water levels fall and Delmarva Status and Distribution: Common. sites become dry, thalli will become stranded Del.: New Castle; Kent; Sussex. Md.: Cecil; on exposed bottoms and take on a terrestrial Kent; Queen Anne’s; Dorchester; Wicomico form of a 1-sided rosette with narrow, forked (SUHC); Worcester. Va.: Northampton. segments. The terrestrial form will overwinter Delmarva Habitat: Aquatic, floating and and when wetland sites become flooded once stranded in Coastal Plain seasonal ponds, again in winter/early spring, the tips of the 246 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 138 thalli become detached, float to the surface, Delmarva Habitat: Usually on moist mineral then grow into the floating aquatic form. soil on banks, wooded slopes and along streams, infrequently on humus. Sphaerocarpaceae Calypogeia muelleriana (Schiffn.) Mull. Frib.

Sphaerocarpos texanus Austin subsp. muelleriana Delmarva Status and Distribution: Uncom- Delmarva Status and Distribution: Common. mon. The species occurs from Texas, east to Del.: New Castle; Sussex. Md.: Cecil; Kent; Florida and north to the Delmarva Peninsula, Queen Anne’s; Talbot; Dorchester; Worcester. its northern limit. Md.: Wicomico, (Davis Va.: Accomack; Northampton. 1976, nss); Worcester. Va.: Accomack. Delmarva Habitat: Usually on peaty humus Delmarva Habitat: Bare, moist mineral soils in swamps, infrequently on mineral soil. of agricultural fields, usually on the perimeter in Calypogeia peruviana Nees et Mont. unplanted strips adjacent to woodlands and Delmarva Status and Distribution: Uncom- thickets. Spores germinate and gametophytes mon. The species extends north on the Coastal develop in early spring. Plain (from South and Central America, Mexico, and the Gulf Coastal Plain) to the Class Jungermanniopsida (Simple Thalloid and Delmarva Peninsula (McAvoy, Biechele and Leafy Liverworts) Knapp 2006), where it reaches its northern

Aneuraceae limit. Del.: Sussex. Md.: Dorchester; Wico- mico; Somerset; Worcester. Va.: Accomack; Aneura pinguis (L.) Dumort. Northampton. Delmarva Status and Distribution: Uncom- Delmarva Habitat: Forming creeping patch- mon. Del.: New Castle; Sussex. Md.: Queen es on humus in swamps, often in Atlantic Anne’s; Talbot; Dorchester; Wicomico; white cedar swamps. Worcester. Calypogeia sullivantii Austin Delmarva Habitat: Forming patches on Delmarva Status and Distribution: Uncom- organic substrates in wet places. mon. Md.: Queen Anne’s; Dorchester Riccardia latifrons (Lindb.) Lindb. (SUHC); Worcester. Delmarva Status and Distribution: Common. Delmarva Habitat: On mineral soil of banks Del.: Sussex. Md.: Queen Anne’s; Caroline; and wooded slopes. Dorchester; Wicomico; Somerset; Worcester Cephaloziaceae (SUHC). Va.: Accomack; Northampton. Delmarva Habitat: Forms crowded, tiered Cephalozia catenulata (Huebener) Lindb. patches that are irregularly branched on Delmarva Status and Distribution: Uncom- decorticated branches and logs in shady, cool mon. Del.: Kent; Sussex. Md.: Wicomico swamps. (SUHC); Worcester. Va.: Accomack; North- Riccardia multifida (L.) Gray ampton. Delmarva Status and Distribution: Uncom- Delmarva Habitat: All of Delmarva’s species mon. Del.: New Castle; Kent (B. Allen, 7756, of Cephalozia form dense patches on humus, 1989, NY); Sussex. Md.: Caroline; Dorchester; decorticated logs and other organic substrates in Worcester. Va.: Northampton. swamps, and they are often mixed with Delmarva Habitat: Forming crowded patch- es other bryophytes. that are regularly branched (ultimate Cephalozia connivens (Dicks.) Linb. segments are parallel sided) on organic sub- Delmarva Status and Distribution: Common. strates in swamps. Del.: New Castle; Kent (B. Allen, 7756, 1989, NY); Sussex. Md.: Talbot; Wicomico (SUHC); Calypogeiaceae Worcester. Va.: Accomack. Calypogeia fissa (L.) Raddi subsp. neogaea Cephalozia lunulifolia (Dumort.) Dumort. R.M. Schust. Delmarva Status and Distribution: Common. Delmarva Status and Distribution: Common. Del.: Sussex. Md.: Kent; Queen Anne’s; Del.: New Castle; Sussex. Md.: Cecil; Kent; Caroline; Talbot; Dorchester; Wicomico; Queen Anne’s; Caroline; Talbot; Dorchester; Somerset; Worcester. Va.: Accomack; North- Wicomico; Somerset; Worcester Va.: Accom- ampton. ack; Northampton. Cephalozia macrostachya Kaal. 2011] MCAVOY ET AL.: LIVERWORTS AND HORNWORTS 247

Delmarva Status and Distribution: Rare. Frullaniaceae Md.: Worcester (SUHC). Frullania asagrayana Mont. Frullania tamarisci Nowellia curvifolia (Dicks.) Mitt. (L.) Dumort. subsp. asagrayana (Mont.) S. Hatt. Delmarva Status and Distribution: Common. Delmarva Status and Distribution: Common. Del.: New Castle; Kent; Sussex. Md.: Cecil; Del.: Sussex. Md.: Cecil; Caroline; Talbot; Queen Anne’s; Caroline; Talbot; Wicomico; Wicomico; Somerset; Worcester. Va.: Accom- Somerset; Worcester. Va.: Accomack. ack; Northampton. Delmarva Habitat: On decorticated logs in Delmarva Habitat: Epiphytic, forming swamps, often forming wide mats or carpets. creeping patches on the bark of trees in woods Odontoschisma denudatum (Mart.) Dumort. and swamps. Delmarva Status and Distribution: Common. Frullania eboracensis Lehm. subsp. eboracensis Del.: Kent; Sussex. Md.: Caroline; Talbot; Delmarva Status and Distribution: Common. Wicomico; Worcester. Va.: Accomack. Del.: New Castle; Kent; Sussex. Md.: Cecil; Delmarva Habitat: Forming creeping patch- Kent; Caroline; Queen Anne’s; Dorchester; es on humus and other organic substrates. Worcester. Odontoschisma prostratum (Sw.) Trevis. Delmarva Habitat: Epiphytic, forming Delmarva Status and Distribution: Common in all counties of the Delmarva. creeping patches on the bark of trees in woods and swamps. Delmarva Habitat: Forming creeping patch- es on mineral soil, humus, decorticated logs and Frullania eboracensis Lehm. subsp. virginica other organic substrates. (Gottsche) Schust. Delmarva Status and Distribution: Common. Cephaloziellaceae Del.: Sussex. Md.: Talbot; Dorchester; Wico-

Cephaloziella byssacea (Roth.) Warnst. var. mico (SUHC). Va.: Northampton. byssacea Delmarva Habitat: Epiphytic, forming Delmarva Status and Distribution: Uncom- creeping patches on the bark of trees in woods mon. Del.: Sussex. Md.: Queen Anne’s; and swamps. Caroline; Talbot; Dorchester; Wicomico. Frullania ericoides (Nees) Nees. Frullania Delmarva Habitat: Forming patches on squarrosa (Reinw; Blume & Nees) Dumort. bare, sandy soil on slopes and banks. Delmarva Status and Distribution: Common. Cephaloziella rubella (Nees) Warnst. var. ru- Del.: Sussex. Md.: Caroline; Wicomico bella (SUHC); Worcester. Va.: Northampton. Delmarva Status and Distribution: Uncom- Delmarva Habitat: Epiphytic, forming mon. Del.: Kent; Sussex. Md.: Queen Annes; creeping patches on the bark of trees in woods Talbot; Somerset; Worcester. Va.: Accomack. and swamps. Delmarva Habitat: On moist, peaty sand Frullania inflata Gottsche and decorticated logs, or as strands among Delmarva Status and Distribution: Common. other bryophytes. Del.: New Castle; Kent. Md.: Cecil; Kent; Queen Anne’s; Caroline; Talbot; Wicomico Fossombroniaceae (Davis 1976, nss); Somerset. Va.: Accomack. Fossombronia cristula Austin Delmarva Habitat: Epiphytic, forming Delmarva Status and Distribution: Uncom- creeping patches on the bark of trees in woods mon. Del.: New Castle; Sussex. Md.: Kent; and swamps. Queen Anne’s; Caroline; Wicomico; Somerset; Jamesoniellaceae Worcester. Delmarva Habitat: Forming seasonal patch- es Jamesoniella autumnalis (DC.) Steph. Delmarva on moist, often disturbed mineral soil; ditch Status and Distribution: Uncom- banks, tire ruts, puddles, Coastal Plain sea- mon. Del.: New Castle; Kent; Sussex. Md.: sonal ponds, and moist swales within the inner Somerset (SUHC); Worcester (SUHC). dunes along the coast. Delmarva Habitat: Decorticated logs in Additional Information: Species resembles a swamps. leafy liverwort, but dorsal sex organs place it Jungermanniaceae in the simple thalloid lineage. Pale-green, ruffle-lobed thalli and red-violet rhizoids are Jungermannia gracillima Sm. Solenostoma distinctive for the species. gracillimum (Sm.) Schust. 248 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 138

Delmarva Status and Distribution: Species is Delmarva Status and Distribution: Common. historical on the Delmarva, known only from Del.: Kent; Sussex. Md.: Kent; Caroline; the two collections cited. Del.: New Castle (A. Dorchester; Wicomico; Somerset; Worcester. Commons, s.n., 1894, NY; A. Commons, s.n., Va.: Accomack; Northampton. 1895, PH). Delmarva Habitat: Epiphytic, forming large, Delmarva Habitat: Forming patches on flat solid patches on tree trunks in woods and moist to dry mineral soil on banks and slopes. swamps, often found on American holly (Ilex

Lejeuneaceae opaca). Leucolejeunea conchifolia (A. Evans) A. Evans Cololejeunea biddlecomiae (Austin) A. Evans Delmarva Status and Distribution: Rare. Delmarva Status and Distribution: Common. Species occurs from Texas, east to Florida, Del.: New Castle; Kent; Sussex. Md.: Cecil; then north to the Delmarva Peninsula, its Caroline; Talbot; Wicomico; Somerset; northern limit. Md.: Dorchester. Worcester Va.: Accomack; Northampton. Delmarva Habitat: Epiphytic on the bark of Delmarva Habitat: Epiphytic on the bark of trees in woods. trees in woods and swamps, usually forming Leucolejeunea unciloba (Lindenb.) A. Evans minute patches in bark furrows and intermin- Delmarva Status and Distribution: Rare. gled with other species. Species is found in Texas and Mexico, east Cololejeunea cardiocarpa (Mont.) R. M. to Florida, then north to the Delmarva Schust. Peninsula, its northern limit. Del.: Sussex (A. Delmarva Status and Distribution: Species is Commons, s.n., 1893, NY). Md.: Wicomico. historical on the Delmarva, known only from a Va.: Accomack (E. Mears, s.n. 1891, NY). single collection from Accomack Co., Delmarva Habitat: Epiphytic on the bark of Virginia. The species is cited here based on trees in swamp. Schuster (1980): Vol. IV, page 1281, ‘‘at the northern most American station on Assatea- Lepidoziaceae gue Island, Virginia (R.M.S 78-301). Species Bazzania trilobata (L.) Gray occurs in the tropics and Florida, north on the Delmarva Status and Distribution: Common. Coastal Plain to the Delmarva Peninsula, its Del.: New Castle; Kent; Sussex. Md.: Caroline; northern limit. Talbot; Dorchester; Wicomico; Worcester. Delmarva Habitat: Epiphytic on the bark of Va.: Accomack. trees. Delmarva Habitat: Forming patches on Cololejeunea minutissima (Sm.) Schiffn. subsp. humus in deep swamps and less frequently minutissima on mineral soil of steep wooded slopes. Delmarva Status and Distribution: Uncom- mon. Species is found from Alabama and Kurzia sylvatica (Evans) Grolle Florida, north to the Delmarva Peninsula, its Delmarva Status and Distribution: Uncom- northern limit. Del.: Sussex. Md.: Worcester. mon. Del.: Kent; Sussex. Md.: Kent; Caroline; Va.: Accomack; Northampton. Talbot; Dorchester; Worcester. Va.: Accom- Delmarva Habitat: Epiphytic on the bark of ack; Northampton. trees in woods and swamps, usually forming Delmarva Habitat: Typically on moist min- minute patches in bark furrows and intermin- eral soil of steep wooded slopes and less gled with other species. frequently on humus and other organic Lejeunea maxonii (Evans) X.-L. He Rectole- substrates in swamps. jeunea maxonii A. Evans Telaranea nematodes (Gottsche ex Austin) M. Delmarva Status and Distribution: Rare. Howe Species is tropical, being found in Puerto Delmarva Status and Distribution: Com- Rico, Jamaica and Florida, then north along mon. This species is near its northern limit the Atlantic coast to the Delmarva Peninsula, on the Delmarva; it is found from Central its northern limit. Del.: Sussex. Md.: Worces- America and the Caribbean, north along the ter. Va.: Northampton. Coastal Plain to Long Island, New York. Delmarva Habitat: Epiphytic on the bark of Del.: Sussex. Md.: Kent; Queen Annne’s; trees in swamps, often mixed with other Dorchester (Biechele); Wicomico; Somerset; bryophytes. Worcester (Biechele). Va.: Accomack; North- Leucolejeunea clypeata (Schwein.) A. Evans ampton. 2011] MCAVOY ET AL.: LIVERWORTS AND HORNWORTS 249

Delmarva Habitat: Intermingled with other Delmarva Status and Distribution: Common bryophytes on humus and other organic in all counties of the Delmarva. substrates in swamps; easily overlooked. Delmarva Habitat: Forms mats on organic substrates in shady swamps. Lophocoleaceae Pelliaceae Chiloscyphus polyanthus (L.) Corda var. rivu- laris (Schrad.) Nees C. rivularis (Schrad.) Pellia epiphylla (L.) Corda Loeske Delmarva Status and Distribution: Common. Delmarva Status and Distribution: Rare. Del.: New Castle; Sussex. Md.: Kent; Queen Del.: New Castle. Md.: Queen Anne’s; Anne’s; Dorchester; Somerset; Worcester. Va.: Worcester. Accomack. Delmarva Habitat: Developing patches on Delmarva Habitat: Forms patches on or- gravel, humus and soft soil in cool, hillside ganic substrates in swamps, forested seeps and seepage wetlands, and submersed in shallow along streams. streams that originate from hillside seeps. Lophocolea bidentata (L.) Dumort. Chiloscy- Porellaceae phus chlorophyllus (Hook. f. & Taylor) Mitt. Porella pinnata L. Delmarva Status and Distribution: Rare. Delmarva Status and Distribution: Uncom- Md.: Queen Anne’s. mon. Del.: Kent; Sussex. Md.: Caroline; Delmarva Habitat: On gravel, humus and Queen Anne’s; Talbot; Worcester (SUHC). other organic substrates in cool, hillside Va.: Northampton. seepage wetland. Delmarva Habitat: Forming dense, hanging Lophocolea heterophylla (Schrad.) Dumort. patches on roots, bases of trees, and other Chiloscyphus profundus (Nees ) J.J. Engel & organic substrates along the edges of freshwa- R. M. Schust. ter tidal and non-tidal streams and rivers; Delmarva Status and Distribution: Common plants are submersed at high tides and during in all counties of the Delmarva. flooding events or periods of high water. Delmarva Habitat: On bark on the base of Porella platyphylla (L.) Pfeiff. trees, mineral soil, humus, and other organic Delmarva Status and Distribution: Rare. substrates in woods and swamps. Md.: Cecil; Somerset (Biechele); Worcester. Lophocolea minor Nees Chiloscyphus minor Delmarva Habitat: Epiphytic on the bark of (Nees) J.J. Eng. & R.M. Schust. deciduous trees in moist woods and swamps. Delmarva Status and Distribution: Rare. Del.: Sussex. Md.: Queen Anne’s; Caroline; Worces- Pseudolepicoleaceae ter (S. King, 196, 2000, DOV). Blepharostoma trichophyllum (L.) Dumort. Delmarva Habitat: Forming creeping patches Delmarva Status and Distribution: Species is on bark at the base of trees and rarely on historical on the Delmarva, known only from mineral soil. the single collection cited. Md.: Worcester (E. Metzgeriaceae Worthley, 85208, 1967, MO).

Metzgeria furcata (L.) Dumort. Delmarva Habitat: Decorticated logs in Delmarva Status and Distribution: Uncom- swamps, usually mixed with other bryophytes. mon. Del.: Kent; Sussex. Md.: Caroline; Ptilidiaceae Wicomico; Worcester. Delmarva Habitat: Epiphytic on the bark of Ptilidium pulcherrimum (Weber) Hampe trees in swamps, or along streams; often on Delmarva Status and Distribution: Species is smooth barked tree species, such as Carpinus historical on the Delmarva, known only from caroliniana. the single collection cited. This species could Additional Information: A form of this be considered disjunct on the Delmarva; it species with thalli that are copiously gemmi- occurs on the Atlantic Coastal Plain from parous (treated by some authors as variety Massachusetts south to Long Island, New ulvula Nees), has been found on the Delmarva. York and then is scattered widely through the Piedmont and Appalachian Mountains south to Pallaviciniaceae North Carolina. Del.: Sussex (A. Commons, s.n., Pallavicinia lyellii (Hook.) Carruth. 1893, PH). 250 JOURNAL OF THE TORREY BOTANICAL SOCIETY [VOL. 138

Delmarva Habitat: Bark of trees. Historical Phylum Anthocerotophyta (Hornworts) collection is on the bark of loblolly pine (Pinus Class Anthocerotopsida taeda). Anthocerotaceae Radulaceae Phaeoceros carolinianus (Michx.) Prosk. An- Radula complanata (L.) Dumort. thoceros carolinianus Michx.; Phaeoceros laevis Delmarva Status and Distribution: Rare. (L.) Prosk. subsp. carolinianus (Michx.) Prosk. Del.: Kent; Sussex. Md.: Caroline; Wicomico; Delmarva Status and Distribution: Uncom- Worcester (SUHC). mon. Md.: Wicomico; Somerset; Worcester. Delmarva Habitat: Epiphytic on the bark of Delmarva Habitat: On bare, moist mineral trees in swamps. soils, usually disturbed soils. Radula obconica Sull. Delmarva Status and Distribution: Rare. Notothyladaceae

Del.: Sussex. Md.: Caroline; Wicomico Notothylas orbiculatus (Schwein.) Sull. (SUHC); Worcester (SUHC). Delmarva Status and Distribution: Uncom- Delmarva Habitat: Epiphytic on the bark of mon. Del.: Kent; Sussex. Va.: Northampton. trees in swamps. Delmarva Habitat: On wet mineral soil, in

Scapaniaceae puddles, tire tracks in fields, and ditch banks.

Gymnocolea inflata (Huds.) Dumort. Literature Cited Delmarva Status and Distribution: Rare. AMMONS, N. 1940. The Liverworts of West Virginia. Del.: Sussex. Am. Mid. Nat. 23: 3–164. Delmarva Habitat: Forming creeping patch- es BAILEY, R. G. 1995. Ecoregions of the United States. on moist, peaty sand in wetland depressions USDA Forest Service, Ft. Collins, CO. 108 p. dominated by cranberry (Vaccinium macro- BRUMMITT, R. K. AND C. E. POWELL. 1992. Authors carpon) within the inner dunes along the coast. of Plant Names. Royal Botanic Gardens, Kew, London, UK. 732 p. Lophozia bicrenata (Schmid. ex Hoffm.) Du- BREIL, D. A. 1970. Liverworts of the Mid-Gulf mort. Jungermannia bicrenata Schmidel Coastal Plain. J. Am. Bry. Lich. Soc. 73: 486. Delmarva Status and Distribution: Rare. BREIL, D. A. 1996. Liverworts and Hornworts of the Del.: New Castle (A. Commons, s.n., 1895, Virginia Piedmont. Banisteria 8: 3–28. CRUM, H. 1991. Liverworts and Hornworts of NY). Md.: Queen Anne’s; Worcester. Southern Michigan. University of Michigan Delmarva Habitat: Forming small creeping Herbarium, Ann Arbor, MI. 233 p. patches on bare, dry sandy soils; plants are DAVIS, B. L. 1976. A survey of the bryophytes of closely appressed to soil. Wicomico Co., Maryland, unpublished report. Lophozia capitata (Hook.) Macoun Junger- Salisbury State College, Salisbury, MD. 10 p. EVANS, A. W. AND G. E. NICHOLS. 1908. mannia capitata Hook. The bryophytes of Connecticut. Conn. Geol. Delmarva Status and Distribution: Rare. Nat. Md.: Worcester (SUHC). Hist. Surv. Bull. 11: 1–203. Delmarva Habitat: Forming small creeping HALEY, J. A. AND D. L. MARSH. 2002. Abstract: A distinctive leafy liverwort in the Arkansas Qua- patches on moist sandy soils. chitas, Arkansas. Academy of Science, 2002 Scapania nemorea (L.) Grolle Annual Meeting, University of Arkansas at Little Delmarva Status and Distribution: Common Rock. in all counties of the Delmarva. HICKS, M. L. 1992. Guide to the Liverworts of North Carolina. Duke University Press, Dur- Delmarva Habitat: On moist, bare mineral ham, NC. 239 p. soil in woods and on slopes, and on soil and INDEX HERBARIORUM. 2010. Retrieved 2 March 2010 organic substrates in seepages. from the New York Botanical Gardens. ,http:// sweetgum.nybg.org/ih/. Trichocoleaceae KING, A. 2000. Assateague Island National Sea- shore, Bryological Survey, unpublished report. Trichocolea tomentella (Ehrh.) Dumort. Assateague Island National Seashore, Berlin, Delmarva Status and Distribution: Rare. MD. 13 p. Del.: New Castle, Kent; Sussex. Md.: Wor- LINCOLN, M. S. G. 2008. Liverworts of New cester. England. Memoirs of the New York Botanical Delmarva Habitat: Forming patches on hu- Garden, Vol. 99. The New York Botanical Garden Press, Bronx, NY. 161 p. mus in swamps and along streams, and is per- MANVILLE, G. C. 2001. An annotated checklist of haps Delmarva’s most pulchritudinous liverwort. liverworts and hornworts (Hepatics and Antho- cerotes) of Pennsylvania. Pennsylvania Dept. of 2011] MCAVOY ET AL.: LIVERWORTS AND HORNWORTS 251

Conservation and Natural Resources, Harris- SCHUSTER, R. M. 1980. The Hepaticae and Anther- burg, PA. 58 p. ocerotae of North America. Vol. IV. Columbia MCAVOY, W. A. 1999. The Bryoflora of the Great University Press, New York, NY. 1334 p. Cypress Swamp Conservation Area, Sussex SCHUSTER, R. M. 1992. The Hepaticae and Anther- County, Delaware and Worcester County, Mary- ocerotae of North America, Vol. V. Chicago Field land. The MD Nat. 43: 9–17. Museum of Natural History, Chicago. 854 p. MCAVOY, W. A., L. T. BIECHELE, AND W. M. KNAPP. STOTLER, R. E. AND B. J. CANDALL-STOTLER. 1977. 2006. New records for the leafy liverwort Calypo- A checklist of the liverworts and hornworts of geia peruviana Nees et Mont. (Calypogeiaceae) on North America. The Bryologist 80: 405–428. the Delmarva Peninsula. Evansia 23: 19–20. PLITT, TATNALL, R. 1946. Flora of Delaware and the Eastern C. C. 1908. A Preliminary list of Hepatics Shore: An Annotated List of the Ferns and found in the vicinity of Baltimore. The Bryologist Flowering Plants of the Peninsula of Delaware, 11: 100–104. Maryland and Virginia. The Society of Natural SCHUSTER, R. M. 1969. The Hepaticae and Antho- History of Delaware, Wilmington, DE. 313 p. cerotae of North America, Vol. II. Columbia THE NATURE CONSERVANCY . 2002. Chesapeake Bay University Press, New York, NY. 1062 p. Lowlands Ecoregion Plan. Arlington, VA. 180 p. SCHUSTER, R. M. 1974. The Hepaticae and Anther- UEBEL, E. C. 2000. Maryland bryophytes collected ocerotae of North America, Vol. III. Columbia by Elmer G. Worthley. Maryland Native Plant University Press, New York, NY. 880 p. Society, Silver Springs, MD.