B AN I STE R I A A JOURNAL DEVOTED TO THE NATURAL HISTORY OF VIRGINIA L Y Number 8 ISSN 1066-0712 1996 BANISTERIA A JOURNAL DEVOTED TO THE NATURAL HISTORY OF VIRGINIA ISSN 1066-0712 Published by the Virginia Natural History Society The Virginia Natural History Society (VNHS) is a nonprofit organization dedicated to the dissemination of scientific information on all aspects of natural history in the Commonwealth of Virginia. Membership in VNHS includes a subscription to Banisteria. Annual dues are $15.00 (per calendar year); library subscriptions to Banisteria are $30.00. Subscribers/members outside the Linked States should add $3.00 for additional postage. Checks should be made payable to the Virginia Natural History Society. Membership dues and inquiries should be directed to the Secretary-Treasurer; correspondence regarding Banisteria to one of the co-editors. Banisteria is a peer-reviewed journal. Editorial staff: Banisteria Co-editors Joseph C. Mitchell, Department of Biology and School of Continuing Studies University of Richmond, Richmond, Virginia 23173 Richard L. Hoffman, Virginia Museum of Natural History, Martinsville, Virginia 24112 Associate Editors Botany Thomas F. Wieboldt, Department of Biology, Virginia Polytechnic Instiaite c Vail, 1893). Thomas Kearney, a vascular plant taxonomist, conducted a botanical survey of the Dismal Swamp in 1901, incidentally picking up six species of liverworts (Patterson, 1949). During the 1930s, M. L. Fenrald made a number of collecting trips to southeastern Virginia, accompanied by Bayard Long of the Academy of Natural Sciences of Philadelphia. Mr. Long collected many bryophytes, including 34 hepatics of which A nthnceros adscerulms was found for the first time north of South Carolina (Patterson, 1951). Hepatics were collected around the Mountain Lake Biological Station during the decades of 1940-1970. .Aaron J. Sharp of the University of Tennessee was the first of many bryologists to conduct field courses on these plants at the station. His collections for the summer season of 1940 alone included 80 different species, 61 of them leafy, 19 rhallose (Sharp 1944). At different intervals, Paul M. Patterson, Rudolf M. Schuster, David A. Breil, and Susan Smdlar also taught courses at Mountain Lake and added substantially to our knowledge of the Hepaticae of southwestern Virginia. While formal regional lists of these collections have not been published, some have been documented in generic revisions and in notes on distribution and ecology. [Tiring the summer of 1944, Irma Schnooberger and Frances Wynne (1944) collected bryophytes including 31 hepatics in the Blue Ridge Mountains of the Shenandoah National Park. Lire majority of their finds were widely distributed boreal species. H. H. litis (1950) reported a number of hepatics for the vicinity of Fredericksburg, Virginia, including the first state records for southern species of leafy Hepaticae. ITiis pajx^r is also significant in recording the first comprehensive collection of bryophytes from the Piedmont section ofVirginia. LXiring the mid 1950s, Rudolf Schuster and Paul Patterson foraged for liverworts in the Dismal Swamp and in the mountains of southwestern Virginia. Jointly they 4 BANISTERIA NO. 8, 1996 reported (Schuster & Patterson, 1957) 16 taxa new to the Virginia Coastal Plain and a similar number for , the mountains. .All but five of the liverwort species were leafy. Since coming to Longwood College in 1968, 1 have collected bryophytes throughout the state with primary emphasis on the plants of the central and southern Piedmont. The accumulated material forms the basis of the following treatment. THE VIRGINIA PIEDMONT The Piedmont Topographic Province extends in a NE - SW direction throughout the length of Virginia and is about 96 km wide at the northern end, broadening to about 192 1cm wide along the North Carolina border. Tire eastern edge of the Piedmont is formed by the Fall Line, a series of rapids occurring in rivers (James, Rappahannock, Potomac, Appomattox and Roanoke) draining to the east. Eire western boundary of the Piedmont is marked by the base of the Blue Ridge Mountain escarpment, about 300 m elevation. The Piedmont is underlain by ancient crystalline rocks mainly covered by residual soils which are somewhat acidic (pH 5.0 - 6.0). Tire area is hilly, with elevational differences not usually exceeding 15 m. Occasional resistant ridges or monadnocks occur as solitary outliers of the Blue Ridge Mountains. Precipitation averages about 45 inches (114 cm) per year occurring throughout tire year except during the drought season during late summer, usually August. Braun (1950) described the outer Piedmont as occurring in the pine-oak region of the Eastern Deciduous Forest. Mature upland deciduous forests are composed of populations of oaks (white, red, post, Spanish, chestnut, scar¬ let), hickories (sweet pignut, pignut, shagbark, mockernut), and mixtures of other hardwood species (red maple, sweetgum, tulip poplar, ironwood, beech, black gum, dogwood, sourwood), often with old successional pines scattered throughout. North slo[>e forests are dominated by American beech with white oak, red or Horida maples, tulip poplars and ironwood. Successional community stages range from old fields to conifer forests (loblolly, Virginia scrub, red cedar), and some hardwood types ( including sweetgum and tulip poplar). Wetland communities include small streams (with hazel alder, sycamore), rocky river shorelines, floodplain forests (with river birch, sycamore, willow oak, American elm, box elder) and grassland marshes. Most reservoirs, lakes and ponds were created in the last hundred years but strongly influence the vegetation of this region. Microhabitats of soil hummocks, rock ledge, rocky ravines, logs, stumps, tree trunks and roots are especially important to the liverworts with the greatest diversity always being found in the more moist shaded areas. THE SURVEY AREA Tire Virginia Piedmont has been virtually unsurveyed for the presence of bryophytes prior to this study. The central axrd southern part of the Virginia Piedmont from Louisa county in the northern part to the North Carolina border was utilized. A buffer zone of about one Piedmont county to the east and west was maintained in order to diminish the direct influence of plants from the mountains and the coastal plain. The counties included in this study are .Amelia, Appomattox, Buckingham, Campbell, Charlotte, Cumberland, Huvanna, Goochland, Halifax, Lunenburg, Louisa, Mecklenburg, Nottoway, Pittsylvania, Powhatan, and Prince Edward (see Figure 88, p. 28). BRYOPHYTE CHARACTERISTICS Bryophytes consist of hornworts, liverworts, and mosses, all of which are small (normally less than two inches long) and have a similar life cycle. Hornworts (anthocerofes) generally occur on the soil and are flat, dark green thalloid plants producing narrowly cylindric saprophytes. Liverworts have two growth forms: thalloid (flat, scale-like, ribbon-shaped, sometimes branched) and leafy: with stem and leaves, occurring in all habitats but most commonly on tree trunks and roots. Leafy liverworts may be confused with mosses but differ from mosses in leaf and sporophyte structure. Leafy liverworts have leaves in two or three distinct rows, each leaf possessing two or more lobes that lack midribs. Moss leaves occur in more than three rows and have singly pointed unlobed leaves with midribs. lire sporophytes of liverworts are short-lived and produce black cylindric or ovate s{>orangia (capsules) which split info four valves to release the spores. Mosses usually develop persistent green to brown sporophytes with sporangia that are ovate, cylindric, spherical, or oblong and allow' the escape of spores through the release of a terminal cap. KEY TO THE SUBDIVISIONS la. Plants thallose, each cell with a single large chloroplast bearing a central pyrenoid; sporophytes linear . . Anthocerotae lb. Plants thallose or leafy, each cell with several small chloroplasts, pyrenoids absent; sporophytes producing stalked spherical or ovoid capsules . Hepaticae BREIL: LIVERWORTS Anthocerotae (Homworts) Key to Genera la. Sporophytes erect, narrowly cylindric, several times longer than enclosing collar of tissue at base (Fig. 1) . . Anthoceros lb. Sporophytes barely emerging from the thallus collar (Fig. 2) . Notothylas Hepaticae (Liverworts) Key to Genera la. Plants thallose, thallus simple, lobed or branched . 2 lb. Plants with stem and leaves . 15 Plants thallose 2a. Thallus surface almost obscured by ovoid sheaths surrounding sex organs or sporophytes (Figs 23, 24) .... . Sphaerocarpos 2b. Thallus surface not obscured . 3 3a. Midrib distinct, cordlike, remainder of thallus thin (Figs 12, 14) . 4 3b. Midrib indistinct, often covered with rhizoids or scales, tapering to margins . 5 4a. Hairs emerging from margins of thallus; on trees and rocks . M etzgeria 4b. Hairs lacking; on organic soil . Pallavicinia 5a. Thallus margins rounddobed, black-spotted at bases; gemmae produced on dorsal surface or in flask-like structures (Fig. 19) . Blasia 5b. Thallus margins straight or wavy, not round-lobed . 6 6a. Plants aquatic, floating or submerged in quiet water, often stranded on shore . 7 6b. Plants terrestrial . 8 7a. Thallus heart-shaped, thick, often in rosettes, with long- toothed ventral scales . Ricciocarpus 7b. Thallus linear, with open forked branching, lacking scales . Riccia, in part 8a. Thallus 1.5 cm or less in diameter, rarely of slender branched ribbons . Riccia, in part 8b. Thallus 1.2 - 6.5 cm long, broadly ribbon-like, some¬ times branched . 9 9a. Plants with green gemmae cups on upper surface (Fig. 7) . Marchantia 9b. Plants lacking gemmae cups . 10 10a. Pores and net-pattern present on upper surface (some¬ times indistinct); scales on lower surfaces (Fig. 8) . 1 1 10b. Pores or pattern on surface of plant lacking; lower surface lacking scales . 13 1 la. Thallus very large, 1 - 3 cm wide; a distinct net-like surface pattern developed around pores . . Conocephalum lib. Thallus smaller, less than 0.8 cm wide; surface pattern present but not distinct; plants often in rosettes . 12 12a. Thallus 5-8 mm wide; epidermal cells with thin walls and large trigones; capsules of elevated receptacles lacking white, scale-like fringe . RebouUa 12b. Thallus up to 3 mm wide; epidermal ceUs with thin waUs, lacking trigones (the epidermis must be mounted separately on slide and examined); capsules fringed by white scales (Fig. 4) . A sterella 13a. Lower surface densely covered with brownish rhizoids along center line; sporophytes developing from a flap or low cup-like structure near apex of thallus (Figs. 17,18) . PeJlm 1 3b. Lower surface sparsely covered with rhizoids (or absent); sporophytes developing from lateral branches . 14 14a. Plants 3-10 mm wide, sparingly branched; yellowish green; always on wet rotten logs and stumps . Aneura 14b. Plants less than 2 mm wide, abundantly branched; deep green; on wet rock, soil or moist rotten logs ....Riccardia Plants leafy, with stem and leaves 15a. Leaves composed of hair-like filaments 1-2 celT wide, or leaf blades fringed with long hairs; underleaves similar to leaves (Figs. 30, 32, 34, 36) . 16 15b. Leaves entire, lobed or toothed, but not with marginal hairs . 19 16a. Plants delicate, less than 1 mm wide; leaves divided nearly to base into 2-4 slender filaments, 1-2 cells wide at base; underleaves of filamentous segments . 17 16b. Plants robust; leaves divided to middle or beyond into 2-5 lobes, the lobes soon divided into many hair-like segments . 18 BREIL: LIVERWORTS 7 17a. Plants pinnately branched; underleaves 2-3 lobed with 1 or more lobes often very short . Kurzia 17b. Plants irregularly branched; underleaves of (3V4 well- developed, filamentous segments . Blepharostonui 18a. Leaf blades 6-10 cells wide at base; cells with large trigones; plants normally reddish-brown . Ptiluhum 18b. Leaf with very little blade, lobes 1-4 cells wide at base, divided into filaments; cells thin walled, without trigones; whitish-gjeen . Trichocolea 19a. Stems lacking underleaves, or under leaves so small as to be inconspicuous . 20 19b. Stems with conspicuous underleaves (Figs. 37, 53, 70, 83,87) . 33 20a. Leaves with lobules (the lobule a smaller portion of the leaf folded back against itself, thus complicate-bilobed) (Figs. 50, 67, 72,78) . 21 20b. Leaves without lobules thus, leaves entire, toothed, lobed or indented at apex . 24 21a. Lobule located above (dorsal to) the leaf, the leaf mar¬ gins usually toothed; on soil (Figs. 50, 51) . 22 21b. Lobule located below (ventral to) the leaf, the leaf margins entire or toothed; plants on roots, rocks or trees (Figs. 71, 72, 77, 87) . 23 22a. Leaves elongated and pointed at apex; lobules strajv shaped, directed towards the stem tip; gemmae sharply angular . Diplophyllum 22b. Leaves broadly ovate to round, barely longer than wide; lobules ovate to rectangular, pointing at an oblique angle to the stem; gemmae ovoid, smooth . Scapanui 23a. Lobules rectangular to rounded; rhizoids on lobule; leaf cells and keel of lobule smooth; perianths flattened; plants large (Fig. 67) . Rcukda 23b. Lobules ovate or reduced to a very narrow fold; rhizoids on stem; leaf cells and lobule papillose; perianths inflated, plants minute (Fig. 7 2) . C ololejeunea 24a. Leaves wavy and irregular in shape, frequently pro ducing small distant hairs along margins; perianth broadened outward, bell-shaped; stem hairs purple (Fig. 9) . Fossombronia 24b. Leaves regular, rarely wavy, lacking marginal leaf hairs; perianths spherical, cylindrical, or flattened contracted or closed at the apex; stem hairs colorless, pink, or brown . 25 25a. Leaves toothed along margins or leaf apex 2-4 lobed . . 26 25b. Leaves entire, round, rounded-rectangular, or ovate, occasionally shallowly notched at tips . 30 26a. Leaf margins toothed all around (sometimes entire margined) . Plagiochila 26b. Leaves 2-4 lobed, the lobe tips sharply pointed . 27 27a. Leaves 2-3 lobed, the lobes toothed on margins . . Lophozia 27b. Leaves 2 lobed, the lobes not toothed along margins .... . 28 28a. Leaf lobes drawn into a fine hairpoints, each several single cells in length; the deeply concave leaf resembling a billowing sail (Fig. 57) . Nowellia 28b. Leaf lobes acute to obtuse, never drawn into a hairpoint; leaves mostly flat to moderately concave ... 29 29a. Leaves attached transversely to stem; plants minute; cells with small oil bodies (Figs. 63, 65) . Cephaloziella 29b. Leaves inserted obliquely on stem; plants larger and easily seen; cells lacking oil-bodies (Figs. 58, 60) . . . . C ephabzia Figures 1-13. 1. Anthoceros laevis subsp. caroliniana. Thallus with erect sporophyte, X3. 2 Notothylas orbicularis. Thallus with emergent sporophytes (mature), X3 3-4. Asterella tenella. 3. Thallus with receptacle, male organs behind, X5. 4 Detail of female receptacle showing papery sheaths around sporophytes, X7 5. Reboulia han.ispha.mca. Thallus with elevated female receptacle bearing sporophytes beneath, male pad behind receptacle, X3. 6-7- Marchantia polymorpha. 6. Habit with female receptacle and gemma cup, X2. 7- Gemma cup, gemmae within, X4 8. Conocephalum amicum. Dorsal surface of thallus, X3. 9-10. Fossombronia brasiliensis . 9. Ventral surface, X6. 10. Spore, X280. 1 1 . Fossombronia wondraczckii. Spore, X2 80. 12. Metzgeria conjugata . Ventral surface of thallus, XlO. 13. Met zgeria crassipilis. Ventral surface of thallus bearing gemmae and rhizoids on wings, XlO. BREIL: LIVERWORTS o 30a. Stems producing long white leafless branches from the lower surface (a long leafy stem must be examined); leaves mostly orbicular (Figs. 84, 86) . Odontoschisma 30b. Stems not developing white branches from lower surface; leaves mostly rectangular to rounded-quadrate (orbicular in some species of Solenostmrui) . 31 31a. Perianths tapered toward a narrow ciliate mouth; bracts beneath perianth ciliate at tips; plants often dull reddish-brown (Figs. 44, 45) . Jarnnsonielh 31b. Perianths not tarred toward mouth and not ciliate; bracts not ciliate; plants green . 3 1 32a. Leaves rectangular on mature sboots; leaf cells with bulging trigones; perianths smoothly cylindric, abruptly constricted to a smooth beak (Fig. 47) . Jungermannia 32b. Leaves circular, elliptical, or ovate; perianths cylindric, longitudinally creased (Figs. 40, 42, 48) . Solenostoma 33a. Leaves producing lobules, the lobules rectangular, strap¬ shaped, or like a smooth Viking helmet (Figs. 69, 71, 78) . 34 33b. Leaves not producing lobules . 37 34a. Underleaves distinctly bilobed for 1/3 to 1/2 their length, lobule helmet-shaped (occasionally strap-shai>ed) . 35 34b. Underleaves entire (sometimes notched at apex), round or broadly ovate in shape . 36 35a. Leaf apex mostly rounded; plants green to shades of red- brown . Frullania 35b. Leaf apex pointed; plants black-green (Fig. 82) ... Jubula 36a. Lobule strap-shaped, sometimes recurved on margins, extending parallel to the stem; each cell with several small oil-bodies (Fig. 85) . Porella 36b. Lobule rectangular, extending parallel to leaf margin and broadly attached to it; one (rarely' 2) large oil-body(s) in each cell . Leucolejeunea 37a. Leaves 3-toothed at apex; underleaves multitoothed (Fig. 37) . Bazzania 37b. Leaves entire or bilobed; underleaves bilobed . 38 38a. Leaves entire (sometimes barely indented at apex) .... 39 38b. Leaves broadly bilobed (a few often entire) (Fig. 52) . . Lophocolea 39a. Leaves rectangular but rounded at the comers, broadly rounded at the apex; near or in water . Chiloscyphus 39b. Leaves ovate-pointed (often bidentate at apex); on soil (Fig. 38) . Gdypogeja Annotated Species Accounts 1. Aneura Dum. Plants thalloid, deep green, 3-10 mm wide, lacking a midrib although thickened in the center. Dioecious; male plants with many elongate antheridial branches arising laterally from thallus; female plants developing erect sporophytes within a fleshy, warty, translucent calyptra. Annum pingms (L.) Dum. (Figs. 20-21) Pioneers on moist rotten logs and stumps, less commonly on tree roots and humus in swamps and shaded margins of ponds and streams. Buckingham, Prince Edward counties. 2. Anthoceros L. Thalloid plants 0. 5-3.0 an broad, in green rosettes or clusters, the margins and upper surface smooth to mffled, lacking a midrib or scales on lower surface; cells each with a single large chloroplast with a central pyrenoid. Monoecious (our species); Figures 14-28. 14-17- Pallavicinia lyellii. 14- Female plant, X7- 15. Male plant, a ntheridia beneath flap, X7- 16. Sporophyte within perigynium on female plant, X7. 17. Pcllia cpiphylla. Monoecious thallus, sporophyte at tip with antheridial bumps behind it, X7. 18. Pellia necsiana. Female thallus with emerging sporophyte, X5 . 19. Blusia pusilla. Thallus with 2 kinds of gemmae and embedded clumps of blue-green bacteria, X7- Thallus, X9. 20-21. Aneura pinguLs. 20. Male plant, x4- 21. Female plant with sporophyte, X4- 22. R iccardia multifidu. 25-24- Sphacrocarpus texanus. 25. Female thallus with dorsal, occluding peri gym a (each later with a sporophyte), X7. 24 Male thallus, with small male organs, X7. 25-26. Riccia hunhenariana suhsp. sullwantii . 25. Dorsal view of thallus with embedded sporophytes, X7. 26. Lateral view of embedded sporophytes, Xl7. 27. Riccia sorocarpa. Thallus with embedded sporophytes, X5. 28. Riccia memhranacea . Thallus, X6. BREIL LIVERWORTS 11 anther id ia and archegonia develo[red in open cavities beneath the upper surface. Sporophyte resembling a single elongated horn, erect, long exserted beyond an enclosing collar from thallus tissue at base. la. Spores yellow . A. carolinianits lb. Spores black . (A. punctatus) 1. Anthoceros carolmianus Michx. (Fig. 1) Also known as Anthoceros laevis subsp. carolinianus, dark green, thalioid plants occurring in rounded clumps on mineral soil along ditches, streambanks and in old cornfields; fmiting from late fall through spring. Plants small in this area. .Amelia, Buck' ingham, Cumberland, Prince Edward counties 2. {Anthoceros punctatus L.) .Also known as Asprromitus punctatus, dark green thalioid plants, dichotomously lobed, occurring in disturbed habitats such as gardens, pathways and ditches, fruiting in winter and spring. Monoecious. Widely distributed through the influence of man’s activities thus expected in gardens. 3. Asterella Beauv. Plants thalioid, elongate, 2-3.5 mm broad, simple or forked, margins thin, wavy, often purple hut green if growing in shade; the upper epidermal cells lacking trigones, the surface indistinctly net-patterned; the lower surface with rhizoids and two rows of crescent-shaped scales. Monoecious; antheridia clustered on upper surface immediately behind the elevated umbrella-like receptacle bearing archegonia and, eventually, papery sheathed sporophytes. Asterella tenella (L.) Beauv. (Figs. 3, 4) Occurring on wet sandy soil in fields, roadside ditches, boulder crevices, rocks along streams; often with Riccia and Anthoceros species. Buckingham, Cumberland, Fluvanna, Prince Edward counties. 4. BazzaniaS. Gray Farge, dull green leafy liverwort, 3A mm wide, forking at the tips and producing white, minutely leafy shoots in axils of the underleaves, heaves densely overlapping, trapezoidal, the apices terminated by 3 (4) shallowly triangular lobes; lobules absent. Underleaves distant to adjacent, large, wider than stem, the margins multitoothed. Not seen with reproductive structures. Bazzcinia trilobata (F.) S. Gray. (Fig. 37) Occurring in moist shaded habitats over vertical granitic or gneissic rock, shaded banks on peaty soil; along rivers or hemlock bluffs. Campbell, Fluvanna, Halifax, Prince Edward counties. 5 . Blasia F. Plants thalioid, pale green, lobed, occurring in mats and forking more or less repeatedly to form rosettes; the thallus center thickened, often with a faint whitish line; rounded lobes with dark blotches at base; lower surface with pinkish scales; upper surface producing two kinds of multicellular gemmae: (1) clusters of star-shaped gemmae and (2) smooth elliptical gemmae within long-necked, flask-shaped structures. Retxrrted dioecious but not seen with sexual structures. Blasia pusilla F. (Fig. 19) Thalli occurring on moist eroding loamy slopes. Prince Edward County. 6. Blepharostoma (Dum. emend Findb.) Diun. Plants delicate, filamentous, 0.7-0. 8 mm wide, in pale green mats or strands among other bryophytes. Feaves inserted transversely on stems, divided into four filamentous lobes nearly to base; underleaves similar to leaves, 3- or 4-lobed. Monoecious. .Antheridia in axils of leaves along stem; perianth cylindrical, contracted towards the ciliate mouth, terminating a stem or branch. Blepharostoma tnchophylhon (F.) Dum. (Figs. 29,30) With other bryophytes or in thin mats on moist shaded rocks; logs, tree bases. Campbell, Huvanna, Prince Edward counties. 7. Calypogeja Raddi Pale green to green medium sized leafy hepatics, occurring singly or loosely intertwined in thin mats; shoots simple or Figures 29-43. 29-30. Blepharostoma trichuphyllum. 29. Shoot, dorsal surface, X 12. 30. Leaf, X64 31-32. Ptilidium pukherrimum. 3 1. Shoot, dorsal surface, Xl 5. 32. Leaf, X3 5. 33-34. Kurzia sylvatica. 33. Shoot, ventral surface, X98. 34. Leaf, X202. 35-36. T richocolea tomentella. 35. Plant, ventral surface, XlO. 36. Leaf, X36. 36. Bazzania trilobata. Shoot, ventral surface, Xl3. 38-39. Calypogeja fissa. 38. Shoot, ventral surface, Xl6. 39. Leaf tips, X99. 40-41. Solenostoma gracillimum. 40. Male plant, antheridia in axils of upper leaves, X23. 40. Leaf marginal cells, X93. 42-43. Solenostoma hyalinum. 42. Shoot, X23. 43. Leaf marginal cells, X99. 12 BANISTER1A NO. 8, 1996 sparsely branched, branches ventral. Leaves ovate, broadly pointed, the apex bilobed or bidentate, less commonly entire. Lobules absent; underleaves conspicuous, entire and slightly notched or bilobed, frequently with a lateral tooth on each outer margin; rhizoids abundant from base of underleaves; unicellular gemmae sometimes produced on outer margins of leaves and at apex of erect branches. Plants monoecious, the male branch short, ventral, with 4-6 pairs of overlapping hracts. The female inflorescence on a short ventral branch, developing a sporophyte within an ovoid fleshy perigynium rather than a leafy perianth as in other leafy hepatics. la. Leaves entire, blunt or narrowly rounded at tips; underleaves rounded, less than 1.3 times as wide as long, with lobes not strongly spreading, their lateral margins evenly rounded, only occasionally toothed; plants robust, 2.5 to 4 mm wide . C. muellenana lb. Leaves frequently bidentate or sharply pointed at apex; underleaves broad, 1.5 to 2.0 times as wide as long, with spreading lobes which have a single tooth; plants smaller, 1.5 to 2.5 mm wide . C.fissa 1 . G aiypogeja fissa (L.) Raddi subsp. neogaea Schust. (Figs. 38, 39) On roadside banks or moist clayey soil of ditches or shaded roadside embankments. Buckingham, Campbell, Cumberland, Fluvanna, Prince Edward, Spotsylvania counties. 2. C aiypogeja muelleriana (Schiffn.) K. Mull. On thin soil or humus in ravines, over shaded rocks, humus on steep slopes, peaty soil. Lunenburg County. 8. Cephalozia (Dum.) Dum. Small leafy plants (0.3 - 0.8 mm wide), in thin mats or mixed with other bryophytes; stems with a translucent outer cell layer; sparingly branched, the lower stem surface with rhizoids throughout; leaves ovate to round, deeply bilobed, distant to slightly overlapping; lobules and underleaves absent. .Asexual reproduction by clumps of gemmae formed at tips of erect branches. Monoecious (our species). Male inflorescence on short branches, with 2- 14 pairs of closely overlapping leafy hracts; perianths on short branches, ovate, obmsely 3-keeled, the apex constricted, toothed. la. Leaf lobes three to five cells wide at base; cell? at base of leaf 40 - 60 p long . G connwens lb. Leaf lobes five to nine cells wide at base; cells at base of leaf 15 05 p long . 2 2a. Leaf bases conspicuously extending down stem; leaf often with two lobes crossing; cells thinwvalled . . C. lunulifolia 2b. Leaf bases not or only slightly extending down stem; leaf lobes not crossing; cells slightly to strongly thick- walled . . C. catemdata 1. Cephalozia catemdata (Hub.) Lindb. (Figs. 60, 61) In moist woods or swamps on moist rotten logs. Often associated with Ncrwellia curvifoha and Odontoschisma denudation. Buckingham, Campbell Ruvanna, Prince Edward counties. 2. Cephalozia connivens (Dicks.) Lindb. (Fig. 62) On shaded moist silty soil along paths, sometimes on moist rotten logs, often with other bryophytes. Campbell County. 3. Cephalozia lunulifolia (Dum.) Dum. (Figs. 58, 59) Occur¬ ring in woods on humus, peaty or sandy soil, sometimes on rotten logs. Ruvanna, Prince Edward counties. 9. Cephaloziella (Spruce) Steph. Minute plants resembling Cephalozia, but without an outer stem layer of large celLs; leaves bilobed with cells containing minute oil-bodies; underleaves absent or minute and narrowly triangular. Perianths and associated bracts large compared to the normal leaves. la. Plants sterile, i.e., sexual structures usually absent; underleaves of robust shoots present, often large; leaf cells thick-walled, some with spines . C. byssacea lb. Plants usually fertile, with perianths and male branches; underleaves minute or absent on sterile shoots . 2 2a. Leaf lobes ovate-triangular, 6-9 cells wide at base of mat¬ ure sterile shoots; leaf cells thin-walled . C. hampeana 2b. Leaf lobes 4-6 cells wide on sterile shoots; leaf cells mostly thick-walled . G rubella 1. Cephaloziella byssacea (Roth.) Warnst. (Figs. 63, 64) Dioecious but usually sterile; plants green to brown; on wet BRE1L: LIVERWORTS 13 seepage on granitic boulders or often mixed with other bryo phytes such as Scapania nemorosa, Dicranum, or Poly trichum. Campbell, Nottoway counties. 2. C ephaloziella hampeana (Nees) Schiffn. - Monoecious; green to brownish plants growing over humus or logs, usually in swamps or other wet areas. Prince Edward County. 3. Cephaloziella rubella (Nees) Wamst. (Figs. 65, 66) Mono ecious; green to reddislvtinged hepatics growing on peaty or sterile soil or rock, often with Solenostoma or protonema of the moss Pogonatum. Prince Edward County. 10. Chiloscyphus Corda Green to pale-green leafy liverworts, the shoots T3 mm wide, irregularly branched; leaves squarish, rounded on the corners; underleaves as wide as the stems, the lobes elongate and triangular with often a thread-like tooth on either side. Monoecious. Antheridia occur in axils of leaves in small cup- like pockets next to the stem on the dorsal surface. Hie perianth occurs on a short ventral branch that is obscurely lobed and toothed. These hepatics possess characters similar to those found in the genus Lophocolea, a taxon which has been transferred to Chiloscyphus by Engle and Schuster (1984). Chiloscyphus pallescens (Ehrh.) Dum. (Fig. 5 3) Rat patches on moist shaded soil, on rock along streams, or in other wet areas. Amelia, Buckingham, Prince Edward counties. 11. Cololejeunea (Spmce) Schiffn. A minute, yellow green species with freely branched, zigzagging stems ; leaves triangular to ovate, the cells with conical papillae on the outer leaf surfaces; lobules well developed, ovate, papillose, with a conspicuous tooth on free margins; underleaves absent; rhizoids developing sparsely along the lower surface of the stem. Monoecious; antheridia occurring at bases of bracts along the stem; large, 5-keeled perianths occur at tips of branches, resembling gas-filled balloons with a terminal beak . Q>lolejeunea buhUecomiae (Aust.) Evans. (Fig. 72) In small patches on tree bark, roots, and rock in shaded moist habitats. Buckingham, Ruvanna, Mecklenburg, Prince Edward counties. 12. Conocephalum Wiggers The largest thallose species, 2-3 inches long, and short branched, occurring in deep green patches; abundant white rhizoids and 2 rows of large scales develop on the lower surface; upper surface with a conspicuous net-like pattern, each unit containing a single open pore leading to a green chamber beneath; gemmae cups and gemmae not produced. Dioecious. Male plants with a slightly elevated “pad” at the tip of the thallus in the spring. Female plants produce an elevated, umbrella-like, receptacle bearing archegonia (later, sporophytes) beneath. This plant is known as the “fragrant liverwort” because of its sweet, spicy odor. Conocephalum conicum (L.) Underwood. (Fig. 8) Occurring over rocks, wet sandy soil, and logs, usually along streams. Buckingham, Cumberland, Ruvanna, Mecklenburg, Prince Edward, Spotsylvania counties. 13. Diplophyllum Dumort. Small leafy plants occurring in green to reddish brown patches, seldom branched; leaves close, complicate-bilobed, the larger blade pointed outwardly, the smaller blade (lobule) pointed toward the stem tip, both lobes finely toothed; angular green gemmae often densely produced at stem apices. Monoecious; male bracts in a series along the stem, similar to leaves. Perianths on short to long branches, ovate, creased lengthwise, and contracted to a tooth-fringed mouth. Diploplv/llum apiculatum (L.) Underwood. (Fig. 51) Common on peaty banks, soil and rocks, especially along road cuts in forests or in partial shade, often occurring with Scapania, and “pigeon wheat moss,” Diphyscium. Appomattox, Bucking¬ ham, Campbell, Ruvanna, Mecklenburg, Prince Edward counties. 14. Fossombronia Raddi A thallose liverwort with deeply lobed wings resembling irregularly shaped leaves, the “leaves” producing spaced marginal hairs; stem rhizoids purple; lobules and underleaves lacking. .Antheridia and archegonia on upper stem surface near the tips of stems or branches. False bell-shaped perianths develop at the base of the sporophyte revealing the spherical black capsule. Most distinctions between species are based on spore ornamentation. Hie plants are seasonal, often appearing with “pygmy mosses” on moist clayey soil. Fossombronia brasiliensis Steph. (Fig. 9,10) Monoecious; occurring singly or in greenish patches on moist, clayey, compact soil in old fields, along streams or in swamps, often with Riccia species or Sphaerocarpos. Cumberland, Prince Edward counties. Fossombronia rwrulracz.ek.il (Corda) Dum. 14 BANISTERIA NO. 8, 1996 may be present. It differs mainly in spore ornamentation, the spores having essentially parallel ridges (Fig. 11). 15. Frullania Raddi Plants leafy, dark green, reddish, or brownish, highly branched, in thin mats closely attached to substrate; leaves entire, overlapping, producing a helmet-shaped lobule next to the stem from the underside of the leaf; occasionally some lobules flattened into a creased, strap-shaped “tongue” paralleling the stem; underleaves large and bilobed. Male branches with closely overlapping bracts in pairs; female perianths large, lantern-shaped and beaked at apex. Asexual reproduction, when present, occurs from dropped leaves which are able to develop into new plants. la. Leaves with a distinct oblique line of colored cells (ocelli); plants usually reddish brown . . F. tamansci subsp. asagrayana lb. Leaves lacking ocelli; plants green to brownish red . 2 2a. Some or all of lobules flattened, forming straps rather than sacs; cell walls lacking trigones and intermediate thickenings . Frullania inflata 2b. Lobules sac-like or helmet-shaped, rarely flattened; cells with large swollen trigones and walk with knot-like thickenings . 3 3a. Under leaves 2.5-4 times as wide as the stem; leaves strongly curving away from stem when moist; plants frequently brownish red . . . F. ericoides 3b. Underleaves 1 - 2 (3) times as wide as main stem; leaves only standing slightly away from stem when moist; plants nonnally green . 4 4a. I aider leaves slightly broader than long, with several teeth above midleaf; lobule sacs compressed near the mouth; plants not usually producing gemmae on leaf margins nor dropping leaves . F. bnttoniae 4b. Underleaves slightly longer than wide with margins entire or a single tooth on either side; lobules not com¬ pressed at mouth; leaves often developing gemmae or falling away from stem . F. eboracensis subsp. virginica 1. Frullania brittoniae Evans. (Figs. 73, 74) Dioecious; in patches on the bark of trunks and branches of trees in hardwood forests. Flalifax County. 2. Frullania eboracensis Gott. subsp. virginica Schust. (Figs. 77- 79) Dioecious, usually with perianths; occurring on bark of hardwood trees in oak forests but also along wooded streams and occasionally on red cedar trunks. Buckingham, Char¬ lotte, Fluvanna, Mecklenburg, Prince Edward counties. 3. Frullania ericoides (Nees) Nees (Figs. 80, 81) Dioecious; found in a variety of somewhat open areas, on bark of hardwoods, red cedars, downed logs. ( Frullania squarrosa) Appomattox, Buckingham, Cumberland, Halifax, Prince Edward counties. 4. Frullania inflata Gott. (Figs. 75, 76) Monoecious; in or near wet areas on trees, granite boulders and moist rotten logs in swamps, beech-maple slopes. Buckingham, Cumberland, Nottoway, Prince Edward counties. 5. Frullania tamarisci (L.) Dum. subsp. asagrayana (Mont.) Hatt. (Fig. 83) Dioecious; found on shaded boulders and rock faces; sometimes on tree trunks in humid shaded forests. Appomattox, Buckingham, Campbell, Cumberland, Prince Edward counties. 16. Jamesoniella (Spruce) Schiffn. Plants leafy, large, greenish to reddish-brown (in sun), growing in patches, 2-3.5 mm broad, sparingly branched with erect growing tips erect; rhizoids numerous along stems; leaves entire, squarish, rounded at comers, somewhat overlapping; cell walls thin with small trigones; lobules lacking. Finder- leaves lacking or very small and narrowly triangular. Di¬ oecious. Male plants producing short male branches with several pairs of bracts. Female plants producing cylindrical perianths, compressed near tips and producing cilia, perianths surrounded by ragged bracts at base. No asexual reproduction. Jamesoniella autumnalis (DC) Steph. (Figs. 44-46) On shaded granite boulders and decaying logs, rarely on humus and rich soil near wooded streams. Amelia, Nottoway counties. 17.Jubula (Steph.) Evans Plants dark green, leafy, medium sized, 1-2 mm broad, branching irregularly pinnately in prostrate mats; leaves overlapping, complicate-bilobed, the blade rounded and abruptly sharp-pointed at apex; lobule helmet-shaped, resembling those of Frullania, but much smaller; stylus absent; underleaves 2-3 times the width of stems, bilobed for 1/2 their length; rhizoids few from underleaf bases. Monoecious. Male branches with 4-5 sac-like bracts, overlapping; perianths lantern-shaped, large, constricted at apex to a small beak. Jubula Pennsylvania i (Steph.) Evans (Fig. 82) On wet rocks in and along streams. Appomattox, Campbell, Huvanna counties. BREIL LIVERWORTS 18. Jungennannia L. Plants leafy, medium-large, in flat green patches, 2- 2.5 mm wide, the stems sparingly branched bearing pale brownish (with age) rhizoids to apex; leaves entire, somewhat over- lapping, mostly rectangular and often notched slightly at apex; cells with bulging trigones; lobules and underleaves absent. Monoecious. Antheridia in the swollen bases of bracts im¬ mediately below the terminal perianth; perianth large, tubular, abruptly contracted at apex into a small beak in a flat or shallow depression. Asexual reproduction rare, by gemmae at tips of erect, small-leaved shoots; gemmae 2-celled, thin-walled. Jungermannia leiantha Grolle. (Fig. 47) .Also known as Jungennannia lanceolata; on soil or moist rocks in wet areas along streams, ditches, or in mixed oak forests; sometimes found with JamesomeUa autumnalis. Buckingham, Prince Edward counties. 19. Kurzia Martens Small filamentous plants often resembling large algae, green to brown, 0.5 mm wide and pinnately branched; leaves divided to base into 3-4 filaments; underleaves like leaves but slightly smaller. Dioecious. Antheridia in the axils of bracts on short lateral branches. Perianths on a short ventral branch, large, cylindrical tapering to a short ciliate mouth . Kurzia sylvatica (Evans) Grolle. (Figs 33, 34) In patches in shaded sites over peaty soil or in ravines; often with species of Caiypogeja, C ephalozia, or O dontoschisma prostratum. Buck¬ ingham, Campbell, Fluvanna, Lunenburg, Prince Edward counties. 20. Leucolejeunea Evans Medium sized leafy hepatics in yellow green patches, sparingly branched, the shoots ca. 1 mm wide; leaves complicate- bilobed, blades ovate, entire, slightly overlapping, each cell with a single large “grape-cluster” oil-body; lobules large, oblong with a folded keel about 1/3 the leaf length; underleaves distant and round, about twice as wide as the stem, bearing rhizoids at the base near attachment. Mon¬ oecious. Male branches short, lateral, with tightly over-lapping bracts; perianths on short lateral branches, ovate with five weak keeL abmptly contracted to a tubular beaked mouth. Leucolejeunea clypeata (Schwein.) Evans. (Fig. 88) In patches on granite boulders and base of trees (oak or hemlock) in 15 shaded sites. Buckingham, Campbell, Charlotte, Fluvanna, Nottoway, Prince Edward counties. 21. Lophocolea (EXim.) Dum. Pale green leafy plants in prostrate mats, occasionally branched. Leaves mostly bilobed, slightly overlapping; leaf cells thin-walled containing a few granular oil-bodies; lobules lacking; underleaves thin and transparent, bilobed, usually wider than the stem, often with a lateral tooth; rhizoids restricted to underleaf bases. Antheridia at base of bracts along the stem (if plants dioecious) or in sac-like bracts beneath the perianths. Perianths tubular, sharply 3-keeled, toothed at their tips. FFiere is the possibility that L bidenta (1.) Dum. will occur here (Schuster 1980, vol. 4) on moist soil. It is dioecious and the leaf lobes end in three to seven celL in a row. Engel > Records ot Anurans from Greensville County, Virginia Richard L. Hoffman Virginia Museum of Natural History Martinsville, Virginia 24112 Joseph C. Mitchell Department of Biology and School of Continuing Education, University of Richmond Richmond, Virginia 23173 Because of its location in southeastern Virginia, adjacent to the North Carolina state line and bisected by the Fall Zone into a western Piedmont half and eastern Coastal Plain half, Greensville County constitutes an important source area for biogeographic studies. Perhaps because of its location, 160 km inland from Virginia Beach and 96 km south of Richmond, the region has been generally overlooked by naturalists and collectors. One notable exception to this neglect was the renowned Harvard botanist Merritt Lyndon Fernald, who scoured parts of the county intensely during his search for rare plants during the 1930s and 1940s (e.g. Femald, 1937). However, from a zoological standpoint, the potential for discoveries has remained almost entirely untapped. Since 1952, the first author has been collecting various kinds of animals in Greensville County. For most of that time excursions were short and sporadic, but following estab- lishment of the Virginia Museum of Natural History in 1989, a fairly regular inventory program has been conducted. A drift fence array was operated from May 1993 to July 1994, with frequent (two week or one month) pickups. .After removal of the pitfalls, a Malaise trap was installed to capture flying insects at the same site. In addition to the pitfall servicing visits, numerous one or two-day forays have been made by museum staff during the past five years, entailing routine hand-collecting as well as operation of blacklight traps at a number of localities. .Although the focus of this activity has been the collection of arthropods, opportunities to observe amphibians have been numerous. Because frogs are so often found by nocturnal road cruising and vocalizing males are so audibly conspicuous, these amphibians have garnered the preponderance of this somewhat tangential attention. During the months of May and June, 1990, the second author conducted inventories of terrestrial vertebrates in the vicinity of Skippers in the southwestern quadrant of the county, and obtained numerous sight and sound records for frogs in addition to specimens taken in pitfall operations. He had earlier (May 1984) accumulated records for reptiles and amphibians in the same general area. On 29 - 30 May, 1960, W. Leslie Burger collected along several secondary roads west and northwest of Skippers. Presumably by road-cmising on the night of the 29th - on which date tropical storm "Brenda" passed through the region - Dr. Burger enjoyed a still unparalleled success with frog captures and obtained no fewer than 1 1 slides. This material was deposited for some years in the VPI&Sl reference collection, and later transferred to the American Museum of Natural History. Through the courtesy of R. G. Zweifel and M. W. Klemens, the second author was able to obtain the relevant collection data. Although coverage of the entire county’ has been inadequate (the northern and western parts in particular receiving almost no attention), we believe that our collective information is of some importance in providing a fairly complete baseline catalog of the local frog fauna, one which is being impacted by ongoing development (agricultural, resi¬ dential, silvicultural). Additional, more systematic, surveys seem appropriate and urgent, not only for anuran species bur animals in general. 30 THE REGION The major geographic features of Greensville County and the collection sites mentioned in the following account are represented on Figure 1 . Because the southwestern (south of US 58 and west of E95) and northern (north of US 58, west of E95) thirds are so disturbed by agricultural and logging activities, most work by RLH has been confined to the southeastern third: adjacent to the still forested floodplains of the Meherrin River and its major tributary, Fontaine Swamp. Collections and observations by JCM were made chiefly in an area southwest of Skippers and also to the northwest thereof, along Fontaine Creek, thus mostly on the Fall Zone itself. NO. 8, 1996 hand clearing goes on apace almost everywhere in the county, and both suburbanization and farming are spreading south from Emporia along county routes 730, 622, and 625. Virtually all of the county roads are now macadamized and carry a substantial load of high-speed traffic around the clock. Not only is the habitat becoming severely fragmented, but migration between the isolates is fraught with danger as well any small animal on the pavement more than a minute or two faces certain death. It may occur in just a few more years that only the sections of the floodplains subject to annual inundation will remain forested. This is very unfortunate because the Coastal Plain biotopes here are the inlandmost in the state and biogeographically interesting on that account. BANISTERIA figure 1. Greensville County, Virginia, showing main highways, settlements, and collecting areas mentioned in the text. The eastern edge of the "fall Fine (or Zone)" is indicated by the heavy dashed line. HOFFMAN & MITCHELL- ANURANS 31 PRIMARY COLLECTING SITES The numbered paragraphs correspond to numbers on the map (Figure 1). 1. Fontaine (or Fountain) Swamp. The lowermost several km of Fontaine Creek, east of US 301, constitute a true year- round swamp, up to 3 km wide in places, which merges into that formed by the Meherrin River as it approaches the North Carolina state line. It is accessible from county routes 624 and 624 which cross the lower third. The southern boundary is marked by the pronounced scarp of a terrace that stands about 20-30 m above the swamp level; definition of the northern side is much less distinct. Most of the original cypress ( Taxodium distichum ) has been logged out, leaving black gum ( Nyssa sylvatica) as the dominant emergent tree species. The higher regions are invested by pines (Pinus echiruita, P. Hie da), sweet gum ( Liquidambar styraafhui), beech (Fagus amencarux), and red maple (Acer rubrum). Co. Rt. 624 nins adjacent to the swamp at nearly water level for about 3.5 km. 2. VMNH pitfall site, 3 km east of Claresville, at the end of Co. Rt. 666. Beyond the end of state maintenance, the road continues as an access into privately owned land in the Meherrin River floodplain. There is a local mosaic of cultivated fields, pine woods, dense black gum swamp, and open areas occupied by water most of the year. The pitfall site is a small knoll high enough that it avoids inundation even during the highest spring floods. The eastern edge of this general area is the complexly meandering course of the river itself, entrenched about 3-6 m below the general level of the floodplain. In the last century the region had been substantially cleared and cultivated, with some dyking and channeling done to manage high water levels; more recently the surviving forest stands have been severely logged out except in the lowest and wettest places. Tire usual community of pine, sweetgum, and red maple is proceeding to recolonize the floodplain. The VMNH pitfall array coirsisted erf four plastic buckets sunk flush with the surface, arranged in a "Y" shape with one bucket at the center and the other three spaced from five to seven meters distant; all were connected by drift fences of sheet aluminum flashing 0.5 m in height. The contents were removed at approximately monthly intervals from May 1994 ter June 1995. 3. Garner's Millpond, located on the Piedmont in the extreme southwestern corner erf the county, is approximately 12 km SSW of Skippers, at the end of Co. Rt. 641. .About 0.7 bn in length, it is fonned by an impoundment of Beaverdam Creek. 4. Proposed site for Virginia Power generating plant, ca. five to ten bn SW of Skipjrers, a subtriangular area delimited by Co. Rts. 633, 632, and 621. Pitfall traps were installed in oak-pine and bottomland hardwood forest types, considered to represent the dominant woodland habitat types. Three pitfalls were placed in oak-pine stands at both the north and south ends of the study area, and six pitfalls were installed in a transect along the western boundary of Fontaine Creek and the eastern boundary of Cattail Creek, in bottomland forest. These traps operated during the period 9 May - 18 June, 1990. This study was conducted by the second author in connection with a contracted biological inventory, and he had previously (1984) obtained data from the same region by manual and visual procedures. Aside from these primary sampling areas, we obtained numerous individual records during road cruising chiefly in the southern half of the county, as specified in the following species accounts. Annotated species list 1 . Bufo americanus americanus Holbrook. In both call and cranial crest characters, the local population is clearly referable to this taxon rather than to Bufo terrestns. Lobey's maps for these two species (1985, p. 54) show Greensville County centered in a major hiatus for American toad records, which can now be corrected. Three adult males (VMNH 6766T3768) were collected at three points southeast of Emporia: on Co. Rt. 730 about 6 bn from the city limits, on Co. Rt. 666 about 1 bn east of Claresville, and on Co. Rt. 624 as it passes through Fontaine Swamp, on the night of 12 April 1995, during the first rainfall in six weeks. Many others were seen by road cruising on the same night. Two (AMNH 122615-16) were taken by Burger "within 3 mi. of Barley", presumably on Co. Rt. 629, on 29 July 1960. 2. Bufo fowleri I Imckley. Common and widespread, this is the species most often seen on the road after dark. Males advertise during May-June near Claresville. Calling and clasping at JCM site 4, on Co. Rt. 621 SE of Brink, 3 May 1984 (5 specimens). Tadpoles in logging road nits, 9 May 1990. Metamorphs 14 June 1990. One immature specimen (VMNH 6583), 1.6 bn E of Claresville, 6 Oct.- 12 Nov. 1993. 3. Bufo quercicus Holbrook. W. L. Burger obtained a single specimen (AMNH 122617) on Co. Rt. 627, 5 bn NE of Barley, on 29 July’ 1960. We have had no personal experience with this species, which must be scarce at this, its inlandmost locality in Virginia. 32 BANISTERIA NO 8, 1996 4. Scaphiopus holbroola (Harlan). One (USNM 135460) found on the road (Co. Rt.. 627, ca 6.8 kmSW of Emporia) after a thunderstorm, 3 May 1953 (RLH). Burger obtained three (AMNH 122387-89) further to the southwest and possibly on the same road ("within 3 mi. of Barley") on 29 July 1960. We have neither specimens nor sound records since then, despite plenty of midsummer road cruising during rainy weather. 5. Pseudacns tnseriata feriarum (Baird). This is an extremely unpredictable species in terms of its spring activities. On occasions, RLH heard the calls along every road in the southeast third of the county, virtually never being out of earshot, when a week earlier or later the roadside ditches and flooded fields were mute. 6. Pseudacris crucifer (W led). Ubiquitous, calling as late as mid-April, and again in late (October and early November, often during the day. 7. Hyla ckrysoscelis Cope. ITiquitous, but not always vocalizing even when climatic conditions seemed optimal to a human observer. The call period seems to be from late April to mid-July. On two occasions small specimens (VMNH uncatalogued) were captured by Malaise trap (1.6 km east of Claresville). 8. Hyla cinerea (Schneider). The green tree frog generally occurs in estaurine situations around the Chesapeake Bay and there are few truly "inland" records in Virginia (in contrast to the situation farther south). It had not been heard calling anywhere in the county prior to the summer of 1993, when many males were advertising in Fontaine Swamp. Capture was precluded by difficult access, but RLH obtained a single male (VMNH 6546) calling in a cypress/black gum swamp, near the intersection of Co. Rts. 730 and 624 on 14 June 1994. On 9 June 1995, males were calling here in large numbers, even from recently clear-cut areas. There is a large population at a pond about 0.5 km west of the intersection of Co. Rt. 624 and the North Carolina state line, but has not been recorded at the Rt. 625 crossing of Fontaine Swamp, only 2.5 km away, during many years of listening at that site. .Astonishingly, a visit to the drift fence site 1.6 km east of ( Jaresville on 9 June 1995 disclosed cinerea calling in great numbers from several separate sites, following a hot day (35° C) arid an evening thunderstorm. Ihe largest aggregation seemed to be in a permanent blackgum swamp, others from an open cattail (Typha latifolia) marsh half a mile away, with individuals and small groups generally distributed. .Again, on 7 August 1996, hundreds of males were calling at this locality, although not a single one was in voice at Fontaine Swamp where advertising males had been numerous in previous years. How could such a large and viable population have been missed in previous summers (e.g., 3 June 1993, 10 June 1994), when light-trapping was being done at the same place, nearly the same time, and under similar climatic conditions, with all of the other associated frog species actively vocalizing? The Claresville population is about 9 km NNW of that in Fontaine Swamp, and suggests that this species may occur throughout the lower Meherrin River floodplain east of Co. Rt. 730 - [xissibly even upstream as far as Emporia. The Greensville populations appear to be by far the most inland known for this species in Virginia. Ihe mating period seems to extend through May and June, hut calling is not continuous through this time. More than once, RLH monitored known population sites on warm nights prior to, during, or after thundershowers, and was met with silence although a week earlier or later, chonises were active. 9. Hyla femoralis Bose. The pinewoods treefrog seems to be widespread in southern Greensville County but nowhere occurring in large colonies, and, to date, not found east of Co. Rt. 730 in the Meherrin floodplain. A fairly extensive population exists in the region ca 5-10 km southwest of Skippers, where heard calling and collected by JCM on 3 May 1984; others calling along Co. Rt. 632, where it passes through a pine plantation, 20 June 1989 (RLH).. Hyla femoralis attains its inlandmost limits at this latitude, and occurs well up on the Piedmont in southern Brunswick County. On 20 June 1989 RLH captured three vocalizing males (VMNH 6403-6405) at an area recently demolished by logging along Co. Rt. 602, 1.6 km W of Triplet. Later that night he heard a large chorus in a pond beside Co. Rt. 603, 2.5 km SE of the intersection named "Fitzhugh" on the DeLorme Atlas, p. 31 (but "Poplar Mount" on the VDOT Bmnswick County map). This seems to he the inlandmost known Virginia locality for this species. No other populations were heard in the region despite a lot of road cruising during climatically optimal periods. HOFFMAN & MITCHELL- ANURANS 33 10. Hyla squrrella Bose. This is another treefrog that RLH missed for many years, until a large vocalizing population was located at Taylor's Millpond on 5 August 1993 (Hoffman, 1994). However, three decades earlier Dr. Burger had encountered a chonis "within 3 mi. of Barley" on 29 July 1960, and secured seven specimens (AMNH 123214-20). The species was heard calling along Fontaine Creek downstream from Co. Rt. 639 on 14 June, 1990 by the second author and C. A Pague. An additional locality was added on 19 August 1994, when a small chorus of about two dozen advertising males was heard beside Co. Rt. 730, at its intersection with Rt. 629 (ca. 6.5 km SE of Emporia). These frogs were calling sporadically in late afternoon following a day of heavy rain. Most recently (9 June 1995) several dozen were calling in a newly plowed flooded field (actually a vast mudpuddle) beside Co. Rt. 730 near Bryant's Comer, in company with Bufo fowlen and Gastrophryne carolinensis. A male was captured and examined but immediately escaped, and local conditions (there being no shoulder on which to get the car off the pavement) prevented stopping long enough to obtain a voucher. Both of these localities have been driven past for many years, without an audible trace of squirella being detected; later in the year they dry up completely and seem a poor choice for a breeding site. 1 1 . Acre crepitans crepitans Baird Sporadic throughout the county, but not always distin¬ guished by the first author from Acris gryllns. Burger captured one specimen (AMNH 122718), 30 July i960, on Co. Rt. 629, 1.5 km SW of Skippers, and the second author both heard and captured specimens at the Virginia Power locality (our site 4) on the 1st and 3rd of May, 1984. 1 2. Acris gryllns gryllns LeConte. The southern cricket frog is widespread in the south¬ eastern third of the county, calling from ditches, small ponds, and gum swamps. Vocalizing begins in mid-April and persists info August, during the day only as the summer wears on. A small voucher series was taken at Gamer's Millpond on 2 1 September 1994 (VMNH 6717-27), all specimens with the hind foot webbing and femoral strips exactly as stipulated for the species (e.g., by Conant & Collins, 1991). One (VMNH 6680) fell into a pitfall trap 1.6 km east of Claresville in the spring (25 March-25 May) of 1994. This site is about 100 m from the nearest w ater. 13. Gastrophryne carohnensis (Holbrook). Narrowmouth toads are obviously common in the county, as suggested by the number which found their way info our pitfall traps located about 1.6 km E of Claresville. VMNH has 3 3 specimens taken during the period 28 April-6 October 1993-1994 with the great majority taken from mid- May to mid-June. Burger found a single specimen (AMNH 122297) 1.5 mi. SW of Skippers on Co. Rt. 629 on 30 July’ 1960. JCM removed a male from a pitfall trap 2.0 mi. SW of Emporia and another 3.0 mi. SSW of Skippers on 18 June 1990. It is a remarkable fact that in spite of numerous visits made by RLH to Greensville County over several decades - before, during, and after thunderstorms, this species was heard calling only once prior to 1990 and even then (25 May 1989) only one male was vocalizing. But for some arcane reasons the evening of 9 June 1995 was apparently optimal for the species, and calling was heard along virtually all backroads between Skippers and Claresville, again at the drift site 1.6 km E of Claresville, and also along Co. Rt. 730 south to Bryant's Corner. On 30 May 1990 JCM heard about 10 vocalizing males in pooh in a clearcut area, 7.5 km SW of Skipjxrs, and saw fresh egg masses of this species. Collectively these data suggest that late May and early’ June is the time of maximum reproductive effort (as also implied by the peak of pitfall catches). Elsewhere in eastern Virginia, calling may extend well into August. 14. Rana catesbeiana Shaw. Calling east of Claresville and at Garner's Millpond 7 June 1990 and 9 May 1991 and a single, very large female on Co. Rt. 625 in Fontaine Swamp on a rainy night (17 July 1993). Two specimens preserved from Site 4, 1 May 1984, by JCM; one seen on Co. Rt. 632, 3 May 1984- 15 .Rana clamitans melanota (Rafinesque) Generally distributed but not abundant, calling through¬ out the summer. Several pitfall captures preserved (JCM) from the Virginia Power site, 1 May and 30 May 1984. 16. Rana palnstns LeConte. Calling in black gum swamp 2 km east of Claresville, also at Garners Millpond, March-April, in small numbers. Not taken by Burger, and no material has been collected by either of us. 34 BANISTERIA NO 8, 1996 17. Ram sphenocephala Cope [in recent literature as Ram utriculana Harlan]. lire species is generally distributed and common everywhere, calling as late as the end of May in 1989. Large numbers of recently metamorphosed specimens fell into the Claresville site pitfalls during the time interval 25 May-30 June 1994, apparently while dispersing from their transformation site. Possible .Additional Local Species In addition to the anurans heard and occasionally captured, we note the conspicuous absences of Rana virgatipes, Pseudacris ocularis, and Pseudacris brimleyi. Tire latter species is plotted on Tobey's map (1985, p. 60) for adjoining Sussex Co. and probably occurs in the northeastern fringe of Greensville. Possibly, the two local species of pseudacrids are competitive, and one excludes the other. The experience of RLH with brimleyi in Caroline County some years ago was that a single colony near Dawn was a tiny island in a sea of fenarum, emphatically verified by prolonged road cruising throughout a 16 km radius of the brimleyi site. Early season visits to Greensville County when both Pseiulacns triseriatu fenarum and P. crucifer were vocalizing en masse have so far been negative for brimleyi, with its simultaneous calling period. Possibly brimleyi is localized in the scarcely collected northern half of the county. The same may be said for P. ocularis, recorded from several sites in adjacent Southampton County to the east, and from Brunswick County to the west. Almost certainly, location of the specific biotope required by ocularis will disclose its presence in Greensville County. Tire apparent absence of Ram virgatipes is similarly inexplicable, considering its inland occurrences farther north in Virginia, and the abundance of apparently suitable local habitat. One noteworthy observation that has been dramatically impressed upon us by recent field experiences in Greensville County is that populations of various species may exist at particular sites without vocalizing during one or more breeding seasons. .Another is that vigorous activity may be in progress at one site, but not at another - just a few miles away - which is known to have been a hotbed of clamor and tumult at other times. Lhe accounts for Hyla cinerea and Hyla squirella provide examples of whar to the human mind may seem flagrant behavioral inconsistency. .An Unusually Diverse Breeding Congress It is not unusual for several species of anurans to utilize a particular breeding site simultaneously, yet in four decades of listening to frogs in Virginia by RLH, only once did he ever hear as many as eight species (half of the known Greensville County frog fauna) advertising at once from the same place. This event occurred beside Co. Rt. 625 about 100 meters north of the VA-NC state line, on the southern periphery of Fontaine Swamp, 25 May 1989. The site had been devastated a year earlier by a clear-cut operation that residted in a deeply- gouged and rutted wasteland of stumps, trunks, and slash, growing up with sweetgum, blackberries (Rubus spp.), and poison ivy (Rhus toxicodendron). Lhe depressions had filled with water and attracted frogs. It was not possible to penetrate the morass, but a half-hour period of listening provided a good idea of what species were vocalizing. Approximate rank order, based on calling frequency, was as follows: Acris gryllus (dozens calling), Hyla chrysoscelis (dozens), Bufo fowleri (many), Ram clamitans melanota (many), Ram catesbeiam (several), Ram sphenocephala (several), Hy la femoralis (only a few), and Gastrophryne caroUnensis (only one). These records were made on a warm, clear, very humid night after a hot day. No rain had fallen for at least a week. Recent visits to the site have been rather negative; it is both drying up and growing up, and only a few of the commonest species listed above were heard. FAUNISTIC SUMMARY Our present total of 17 confirmed species of anurans is apparently the greatest now recorded for any Virginia county or comparable city, using the maps in F. J. Tobey's atlas (1985) as resource. The City of Virginia Beach is there credited with 16 species, the City of Chesapeake with 14, Surry County with 15, and Fairfax, Charlotte, and .Alleghany counties with 12 each. In acmality Surry County may claim first place, since several very common species were not represented in Tobey's data, and when collected (if they have not already been taken) will raise the Surry total to 20 species or more, about the maximum to be expected for a Coastal Plain county. .As is evident from the foregoing list, the Greensville County anuran fauna basically reflects two distribution patterns, one being that of species widespread over much of eastern l Anted States such as Bufo amencanus, Bufo fmvlen, Acris crepitans, Pseudacris triseriata, P. crucifer, Scaphiopus holbrooki, Ram catesbeiam, R. clamitans, and R. palustris. The second pattern is that of the classical "Lower Austral" life zone, and includes species which, if not actually restricted to the Coastal Plain, are at least most abundant there and may exist only sporadically on the Piedmont. Species in this category include *Bufo quercicus, *Hyla cinerea, H. chrysoscelis, H. femoralis, *H. squirella, Acris , gryllus , and Gastrophryne carolinensLs (species marked by an asterisk reach their inlandmost Virginia localities on the Fall Line belt south and west of Emporia; and HOFFMAN & MITCHELL- AN! JRANS 35 H. femoralis extends only a few km further westward into Bnmswick County). If is a little surprising that the "American Toad" represented here is the northern B. americanns rather than the southern B. terrestris. CONSERVATION STATUS Documentation of amphibian, particularly anuran, population declines are being published at an alarming rate (e.g., Drost & Fellers, 1996; Gamradt & Kats, 1996; Laurance et ah, 1996). These reports and others demonstrate that amphibians worldwide are subject to a wide variety of perturbations (see reviews in Phillips, 1990, 1994; Livermore, 1992; Pechmann & Wilbur, 1994; Blaustein (Sc Wake, 1995; Stebbins c Gustavo R. Spmelli. 1995. Patagonian Odonata in Malaise traps. Argia 7(3): 22-23. Roble, Steven M. 1994. A preliminary checklist of the damselflies of Virginia, with notes on distribution and seasonality (Odonata: Zygoptera). Banisteria 4: 3-21. Roble, Steven M. 1995. [Note on Malaise trap records of Virginia Odonata]. Argia 7(2): 3. Roble, Steven M., 1996 by the Virginia Natural History Society PREDATION OF MARBLED SALAMANDER (AMBY STOMA OPACUM [GRAVENHORST]) EGGS BY THE MILLIPED UROBLANIULUS JERSEYI (CAUSEY) - Millipeds (class Diplopoda) are detritivores, feeding mainly on leaf litter and other forms of decomposing plant material. They have not hitherto been implicated in predation of amphibians or their eggs, although Schubart (1947) mentioned that a specimen of Heteropyge araguayensis (Schubart) (family Spirostreptidae) devoured an unidentified Brasilian hylid with which it had been confined overnight. It is unknown if the frog was killed by the conditions of its confinement or by defensive allomones produced by the milliped. It is therefore of interest to record our observations of apparent predation of amphibian eggs by millipeds in Virginia. On 7 October 1987 six adult Ambystoma opacum were found guarding their egg clusters in the Maple Flats region of the George Washington National Forest, 13 km S of Stuarts Draft, Augusta County, Virginia. Al were found under decaying logs in dry vernal ponds. The soil under the logs was moist and the salamanders were found in depressions with their eggs. In two of the nests we found several individuals of the milliped genus Uroblaniulus crawling among the eggs. Five millipeds were found in one clutch and three in another. Al eggs, attending females, and millipeds were collected for subsequent examination. Each egg was examined closely for indications of punctures and mutilation, however, none were found. We were also unable to see actual predation in the field. The stomachs of the six females contained only one beetle larva, earthworm remains, and what appeared to be a salamander egg, but no millipeds. This suggests that Uroblaniulus jerseyi may enjoy some immunity from predation by these females. Adult marbled salamanders are predators of a variety of invertebrates (Surface, 1913; Bishop, 1941) but millipeds have not yet been reported in their diets. Carnivory is an exceptional lifestyle among millipeds. The literature on this subject was reviewed by Hoffman r. Roble for his donation of this material (and thousands of other insects taken during inventory activities). References .Andersen, N. M. 1975. The Limnogonus and Neogerris of the Old World with character analysis and a reclassification of the Gerrinae (Hemiptera: Gerridae). Entomologica Scandanavica (Supplementum) 7: 1-96. Blatchley, W. S., 1926. Heteroptera or Tme Bugs of Eastern North .America. Nature Publishing Company, Indianapolis. 1116 pages. Bobb, M. L., 1974. The aquatic and semi-aquatic Hemiptera of Virginia. The Insects of Virginia: No. 7. Research Division Bulletin, Virginia Polytechnic Institute