SMITHSONIAN INSTITUTION

SCHOLARLY STUDIES PROGRAM

Name S.I. Address & Phone Principal Investigator:

Robert Hershler Invertebrate Zoology, NMNH,

Smithsonian Institution,-

Washington, DC 20560

(202) 786-2077

Co-Principal Investigator: Margaret Mulvey Savannah River Ecology

Laboratory, Drawer E, Aiken,

SC 29801

Collaborator(s):

Proposal Title:

Evolution of the North American Springsnail

Budget: Year 1 $14,640 Year 2 $11,326 Total $25,966

Start Date and Duration of Project: March 1, 199; two years

Signatures: Principal Investigator

Co-Principal Investigator NON-SPECIALIST SUMMARY

This proposal requests funding to continue research on the western American freshwater snails ("springsnails") of the family

Hydrobiidae; we plan a collaborative study of the evolution and biogeography of Tryonia, a diverse group of 21 described distributed throughout major drainages of western North America. Because they are obligate inhabitants of water and disperse slowly, the distributions and evolutionary histories of springsnails are closely tied to regional drainage histories.

This synthetic study of relationships, which builds from the previous research of the Principal Investigator, will yield a landmark contribution to the biogeography of western American freshwater mollusks. These mollusks are well suited for such an inquiry owing to their distribution in a highly disrupted and historically complex drainage, but the absence of corroborated phylogenetic hypotheses for these has prevented their utilization in such studies. Tryonia has been selected for this pioneer study because it is the only large genus of western

American springsnails that is well known in terms of scope and content, and includes endemic species with restricted distributions which constitute ideal units for biogeographic inquiry. The primary goals of this research are to reconstruct the evolutionary history of Tryonia using a multidisciplinary approach, employing both morphologic (shell and anatomy) and genetic (allozyme) data; to use the phylogeny as the basis for

1 generating and testing biogeographic hypotheses that explain the current distribution of these species; to compare these results with those of similar studies of western fishes; to evaluate whether co-occurring species form spring-dwelling species flocks

(i.e., groups of locally endemic species that are closely related) comparable to those identified in lacustrine fishes; and to study the evolution of parthenogenesis (asexuality) and of miniaturization among members of the genus. The resulting genetic database also will be of great use to management programs

for Tryonia; these snails are largely restricted to national parks, monuments, and wildlife refuges and include many species that are candidates for addition to the (Federal) List of

Endangered and Threatened Wildlife. Morphological data will be obtained by dissection, and study

of shell, , , and ciliary patterns on exterior body surfaces using Scanning Electron Microscopy. Protein electrophoresis will be performed on large samples from one or more populations of each species to provide allozyme data. These

data will be used to prepare a cladistic analysis of the

evolutionary history of Tryonia using several available phylogenetic software packages. The phylogeny will be compared

with information on the historical development of western American drainage based on the physical record using techniques of vicariance biogeography. Anticipated products of the project are a taxonomic revision

of Tryonia and a synthetic paper reviewing the evolution and biogeography of the genus and its species.

2 Introduction

This proposal requests funding to continue research on western American freshwater snails of the family (: ). Previous work by the Principal

Investigator has largely focused on taxonomic description and

documentation of diversity of these poorly known snails (see

Hershler 1985, 1989; Hershler and Longley 1986; Hershler and Sada 1987; Hershler and Landye 1988; Hershler and Pratt 1990), and

included a recent (1991-1993) field survey of the Great Basin fauna funded in part by the Scholarly Studies Program. Research objectives of the Great Basin project have led to a shift toward

synthetic studies of relationships, with the first such product

being a revision and phylogenetic analysis of the large genus (Hershler In Press). We request support to continue this increasingly synthetic program in the form of a

collaborative study on the evolution and biogeography of Tryonia, a diverse genus distributed throughout major hydrographic units

of western North America, including not only the Great Basin, but

also the Colorado River drainage and disrupted segments thereof (Death Valley "System", White River drainage), and Pecos-Rio

Grande drainages of New Mexico and Texas (Figure 1).

The western North American freshwater molluscan fauna, comprising about 65 genera and 300 species in 24 families (Taylor 1985:290) and comparable in size to the regional fish fauna, is

distributed in a highly fragmented and historically complex drainage that provides an ideal setting for study of the evolutionary consequences of vicariance. The small (1-8 mm shell

3 length), gill-breathing snails of the family Hydrobiidae are the

most diverse group of freshwater mollusks in the West, totalling

about 85 described species, and probably the most suitable subjects for biogeographic inquiry. These animals have direct development, are obligate inhabitants of water, and are

presumably slow to disperse, although they may be passively transported by floodwaters or birds. As a result of this combination of factors, western hydrobiids usually occur as narrowly distributed species restricted to single springs or spring systems. The modern fauna parallels regional fishes in predating geologic and climatic conditions under which they are found today (extant genera having minimum age of Late Tertiary; Taylor 1985) and thus their evolutionary histories can be correlated with the remarkable array of events associated with development of western drainage.

While this molluscan fauna has proven to be a rich subject for biogeographic speculation (Taylor 1960, 1966a, 1985, 1987, 1988; Taylor and Bright 1987; Hershler and Pratt 1990), explanatory scenarios were posed solely based on distribution patterns and "intuitive" opinions of evolutionary history. These lacked the phylogenetic approach critical to rigorous constructions of historical biogeography (Humphries and Parenti

1986). Preparation of these hypotheses has been limited by incomplete knowledge of the scope, content, and of this fauna, with the most significant factor the absence of species- level phylogenetic hypotheses for major western groups. The Principal Investigator recently completed the first such

4 L analysis, treating the genus Pyrgulopsis (Hershler In Press). This was necessarily preliminary as the 53 species studied

represented only the nominate fauna and perhaps as many undescribed congeners await study.

During the phase of research proposed herein, which capitalizes on the differing expertise of the two participants,

populations and species of Trvonia will be examined in detail using shell morphometric, anatomical, allozyme, and available paleontologic data in order to produce a clear picture of the

evolutionary history of this speciose clade. Cladograms will be

used to generate and test biogeographic proposals of the historical pathways resulting in the current distribution of these species. These will in turn be contrasted both with the known physical record pertaining to the hydrographic history of the region and the results of similar studies of western fishes

that live in association with these snails. Pertinent aspects of

geological history include accretion of allochthonous terranes

onto California and Nevada, development of the Gulf of California

and its inland embayment to the north, opening of the Rio Grande Rift, crustal extension, rotation and displacement of intra- continental subplates, tectonic formation of mountains and

basins, and development of basinal lacustrine habitats. The

rigorous phylogenetic hypothesis will permit one of the few

modern evaluations of the development of putative species flocks

in invertebrates and provide the framework necessary for the study of evolution of parthenogenic and miniature-sized species

in the genus. Documentation of the extent of genetic variation and differentiation among populations (or species) will greatly

5 aid ongoing management programs for these snails, which are largely restricted to national parks, monuments, or wildlife refuges and include many species that are as threatened or endangered as are associated fishes.

Study Group and Rationale

We selected Tryonia for detailed study since it is the only large genus of western American Hydrobiidae reasonably well known in terms of content and distribution. The genus is composed of 21 described Recent species (Table 1), providing diversity suitable for biogeographic inquiry, but is not so large as to pose problems for robust phylogenetic analysis (Sanderson and

Donoghue 1989). The study will also include two undescribed, locally endemic species from the northern Great Basin (Monitor Valley, Steptoe Valley). The Principal Investigator has worked on this group previously, having described many of its species

(Hershler and Sada 1987; Hershler and Landye 1988; Hershler

1989), and has accumulated a large collection (more than 125 lots) of anatomical material of most species in preparation for the morphological component of this study.

Tryonia is distributed along a broad arc sweeping across the major drainages of the Southwest, extending from the Pecos River west to the California Coast and from the northern Great Basin to an indeterminate southern limit in Mexico. In addition, a disjunct group of two eastern species occurs in peninsular

Florida (Hershler and Thompson 1987), providing opportunity to

6 examine relationships of taxa profoundly separated across the Continental Divide. Sixteen of the 21 described species of Tryonia are restricted to either single springs or spring systems, providing numerous areas of endemism suitable for

biogeographic inquiry. One species, Tryonia protea, has a uniquely disjunct distribution which has been interpreted as

indicating former connection between the Colorado River drainage

and Bonneville Basin (Taylor 1985; Taylor and Bright 1987), but the taxonomic status of these widely separated populations has

not been carefully studied. Although Tryonia species are usually distributed in an allopatric fashion, clusters of locally endemic

forms that appear closely related occur (often syntopically) in

both Death Valley (four species) and Ash Meadows (three species), thus providing an opportunity for study of development of

possible species flocks (sensu Greenwood 1984; Ribbink 1984), which heretofore has been largely restricted to lacustrine ichthyofaunas.

The extensive studies of mode and pattern of differentiation of western pupfishes (genus Cyprinodon) (Echelle and Dowling

1992; Echelle and Echelle 1992, 1993; and references cited therein), which parallel Tryonia both in terms of distribution and development of local species flocks (Miller 1981), provide a source of data for evaluating congruence of biogeographic pattern.

In the West, Tryonia are found in springs (with the exception of T. imitator, which lives in brackish coastal waters), often thermal and highly mineralized, thus indicating an unusual level of physiological tolerance. (The two eastern

7 species are found in lakes and springs.) Tryonia live in soft benthos, and usually are extremely common, permitting collection of large samples. Female Tryonia are ovoviviparous, brooding young in their distal gonoducts. One species, I. Drotea, is parthenogenic and others are characterized by either rareness and/or reduced body size of males. One New Mexican species of

Tryonia has been listed as Endangered by the U.S. Fish and Wildlife Service (USDI 1991a) and another 13 western species are candidates for addition to the List of Threatened and Endangered Wildlife (USDI 1991b). Members of the genus Tryonia vary in terms of size, sculpture and shape of shell, and include several "miniaturized" forms that mature at only about a third of typical adult size

(Figure 2). Anatomical variation other then penial ornament has been scarcely studied and even these details have not been recorded for some species. Tryonia has not been the subject of a modern review and no classification is available for members of the genus, which have been described as "... often a series of monotypic species-groups with no evident mutual relationships"

(Taylor 1987:6), although Taylor (ibid.) created the subgenus

Paupertryonia based on a single penial character. The sister group to Tryonia is unknown, although the genus is very similar to other genera of the subfamily Cochliopinae that also have penes bearing glandular papillae (Hershler and Thompson 1992). Monophyly of the genus, suggested by the unique combination of shell and genitalic features that define the group (Hershler and

Thompson 1992), has not been corroborated although such will be

8 sought in a phylogenetic analysis of cochliopine genera that is

in progress (with George Davis [Academy of Natural Sciences of Philadelphia]).

The two phylogenetic analyses thus far produced for genus-

level groups of hydrobiids (Ponder and Clark 1990; Hershler In Press) yielded polytomy-laden cladograms without adequate resolution for construction of detailed biogeographic hypotheses. This reflects the high levels of homoplasy and paucity of significant synapomorphies among the morphological characters employed. Morphological variation among species of Tryonia does not appear to be great and thus we will use biochemical data to help clarify intrageneric systematic relationships and supplement morphological characters in phylogenetic reconstruction. Starch gel electrophoresis provides a rapid method to obtain genetic information from many individuals and has proven a useful technique for recovering phylogenies at the intrageneric level (Avise 1975; Buth 1984; Hillis 1987). This technique has been been successfully applied to freshwater gastropods, including several studies pertaining to hydrobiids (Davis et al. 1988;

Ponder and Clark 1988; Klemm and Schlegel 1989; Ponder et al.

1989). Data obtained during a recent local Tryonia study in California (Hershler 1992) provide encouragement as the two species examined were very strongly differentiated in terms of allozymes (fixed-allele difference at 14 of 20 loci). We acknowledge that there are some problems inherent in using allele data for phylogenetic analysis, but they nevertheless represent a critical first step in a molecular genetic approach to the systematic and evolutionary problems of interest herein.

9 Materials and Methods

Fieldwork. A total of 53 collections of Tryonia will be made for electrophoretic study (see Table 1; note that single samples will also be obtained for each of the two undescribed species from the Great Basin)1. Most of the species have a restricted range and for these one to three samples will be taken. The few species having a wider distribution (imitator, protea) will be studied in greater detail (4-6 samples). An additional six populations will be sampled as potential outgroups

(Floridian species of Aphaostracon, Littoridinops, and Pyrgophorus). From each population sampled, 40 adult snails will be collected and frozen in the field (immersed in tris-buffer) using liquid nitrogen or dry ice. Samples will be shipped by express mail to Savannah River Ecology Laboratory (SREL) where they will be stored at -70° pending electrophoretic study. In the few cases where samples also need be taken for morphological studies, large series will be collected, relaxed with menthol

' Tryonia patzcuarensis (Pilsbry, 1891), living in a lake near Mexico City and questionably allocated to the genus

by Taylor (1966b:197), and a small undescribed fauna in northern Mexico (Chihuahua, Coahuila, Durango) will

not be included in this study because extensive fieldwork will be required to assess these taxa and current

problems in obtaining permits to collect in Mexico prevent confident assumption that this work can proceed.

Instead we will pursue study of these species following completion of the current project. Also excluded are the

following: Trvonia diaboli (Pilsbry and Ferriss 1906), which was not assuredly based on Recent material

(Fullington 1978); Tryonia hertleini (Drake, 1956), which probably went extinct following drying of its type

locality in Chihuahua, Mexico; and Hawaiian Tryonia, possibly referable to Paludina porrecta Mighels, 1845,

which is likely introduced from California and not assuredly extant.

10 crystals overnight, fixed in dilute formalin (10% of stock solution), and preserved in 70% ethanol for laboratory study.

Subsets of these collections will be dried to provide shell material. In most areas where fieldwork is to be done, permits are required by State and/or Federal agencies. Permits already have been sought for the first field trip to the Death Valley region (California Department of Fish and Game, National Park Service

[Death Valley National Monument], Nevada Department of Wildlife, United States Fish and Wildlife Service [Ash Meadows National

Wildlife Refuge, Moapa National Wildlife Refuge)), which already has been funded by the Smithsonian Institution Research Opportunities Fund (in conjunction with provision of "seed money" as explained below). Great Basin material (Utah Drotea, undescribed Nevada species) already has been obtained in conjunction with the earlier project mentioned above. Remaining field trips are summarized below.

1. California, Arizona, New Mexico, Texas (April-May, 1994). Collect adamantina, alamosae, brunei, cheatumi, circumstriata, gilae, kosteri, quitobaquitae. 21 days,

originating in Tucson.

2. California (July, 1994). Collect imitator, protea. 7 days, originating in San Diego.

3. Florida (January, 1995). Collect aequicostata,

brevissima, outgroup populations. 10 days, originating in

Orlando.

11 Laboratory Studies. Morphologic study will be made of one or more populations of each species (including outgroups) corresponding to the same demes sampled for electrophoretic study. Each population (or species) will be scored for a set of up to 175 potentially suitable characters from shell, anatomy, and reproductive biology (derived from a larger set prepared in collaboration with Winston Ponder [Australian Museum] and George Davis), although deletions or additions may occur as the study progresses. Shell shape parameters (Raup 1966) will be obtained from series of sexed specimens following methods of Hershler and

Sada (1987). Shells, radulae, and opercula are cleaned with liquid household bleach (CLOROX) and then studied and photograped using a Hitachi S-570 Scanning Electron Microscope (SEM). Whole animals, or parts thereof, are dried using a Denton DCP-1

Critical Point Drier and studied with SEM to elucidate details of ciliation. Small (3-6 individuals) sets of male and female animals are dissected to obtain information on anatomical characters. Inorganic shell material is removed by soaking specimens in Bouin's Solution or dilute hydrochloric acid.

Animals are dissected in dilute Bouin's Solution according to the protocol of Hershler (In press). Anatomical illustrations are prepared from camera lucida drawings of pinned out specimens. Protein electrophoretic methods described by Mulvey and Vrijenhoek (1981), and Hershler (1992) will be used to obtain allozyme data for 20 or more loci. Samples of 40 individuals from each of the 59 populations will be assayed. Samples of T. protea, a parthenogenic species, will be run on all gels as

12 reference material for comparison of allozyme mobilities between runs. Samples from multiple populations also will be run concurrently to allow for comparison of allozyme mobilities. Gels will be photographed to provide a permanent record of results.

Data Analysis. For the morphological data, continuous characters will be divided into states separated by gaps (Archie

1985). Multistate characters will be treated as unordered, and characters that are polymorphic within a species will be scored for the presumed derived state. Allozyme frequencies, percent polymorphic loci, heterozygosity, and genetic distance coefficients will be obtained using BIOSYS-1 (Swofford and Selander 1981). Genetic variation within and among populations and species will be defined.

Morphological and genetic data will be analyzed together, as well as separately, in order to evaluate congruence. All phylogenetic analyses will be rooted using a series of outgroups. We will employ two methods to investigate phylogenetic relationships using allozyme data. First, we will apply the distance Wagner method (Farris 1972), using Swofford's (1981) multiple addition criterion procedure and Rogers' genetic distance (Rogers 1972, 1984). In addition, we will treat allozymes as discrete characters and use PAUP version 3.0 (Swofford 1990) and HENNIG86 (Farris 1988) to derive cladograms. Only the single most common allozyme for each locus will be scored, as either present or absent. The morphological data and combined data sets also will be analyzed using the above

13 phylogenetic software packages. Character state evolution on preferred tree(s) will be studied using CLADOS (Nixon 1992). The resulting phylogenetic hypothesis will be used to generate area cladograms according to methodology of vicariance biogeography summarized in Wiley (1981) and Humphries and Parenti (1986). Area cladograms will be compared and contrasted with information on historical development of western American drainage (summarized in large part by Smith 1978; Hendrickson

1986; Minckley et al. 1986; Smith and Miller 1986), as well as with the emerging literature of similar studies of western American freshwater fishes (ibid.; also see Hocutt and Wiley 1986; Mayden 1992). Non-concordance between phylogeny and geological/hydrological history will be evaluated within the context of possible dispersal events.

Anticipated products of this project include a formal taxonomic revision of the genus Tryonia that will include description of the two new species from the northern Great Basin; and a major, synthetic paper reviewing the evolution and biogeography of the genus and its species.

Personnel

The principal investigator will devote approximately 30% of his time over two years to the project. He will perform all fieldwork and morphologic studies, and prepare the taxonomic revision of Tryonia. The Co-Principal Investigator will devote 15% of her time to the project over two years; she will perform all of the electrophoretic analyses at SREL and work together

14 with Hershler on data analysis and preparation of manuscripts

treating the evolution and biogeography of Tryonia.

Timetable

All fieldwork will be done during the first year of the

project, during which time slightly more than half of the

laboratory studies also will be completed. Laboratory studies will be entirely completed by mid-point of the second year, and the last six months will be spent compiling and analyzing data,

and preparing the two main manuscripts which will arise from the study.

15 Table 1. List of Recent species of Trvonia, with type localities, distributions, and number of populations to be sampled for electrophoretic study (in parentheses). Additional nomenclatural details are provided by Hershler and Thompson (1992).

Tryonia adamantina Taylor, 1987. Diamond Y Spring, Pecos County, Texas. Type locality and several nearby springs, Pecos River drainage (2).

Tryonia aequicostata (Pilsbry, 1889). Sumter County, Florida. (Type lot attributed to Lake Panasoffkee;

Thompson 1968:50.) Springs and lakes in upper Florida Peninsula, St. Johns and Withlacoochee River drainages, and Lake Okeechobee (Thompson 1968) (3).

Tryonia alamosae Taylor, 1987. Ojo Caliente, Socorro County, New Mexico. Type locality and nearby springs at head of Alamosae Creek, Rio Grande drainage (2).

Tryonia angulata Hershler and Sada, 1987. Fairbanks Spring, Nye County, Nevada. Type locality and two other springs, Ash Meadows, Amargosa River drainage (2).

Trvonia brevissima (Pilsbry, 1890). Haulover Canal, at head of Indian River, Florida. (Locality considered in error by Thompson [1968]). Lakes and springs in central Florida Peninsula (Thompson 1968) (3).

Tryonia brunei Taylor, 1987. Outflow of Phantom Lake Spring at Joe Kingston Ranch, Jeff Davis County,

Texas. Endemic to type locality, Pecos River drainage (1).

Trvonia cheatumi (Pilsbry, 1935). Phantom Lake, near Toyahvale, Reeves County (emended to Jeff Davis

County by Taylor [1987]), Texas. Phantom Lake and San Solomon Springs, Pecos River drainage (2).

Trvonia circumstriata (Leonard and Ho, 1960). Late Pleistocene terrace deposits along Pecos River near

Independence Creek, Terrell County, Texas. Endemic to spring in Diamond Y Draw, Pecos River drainage (1).

Tryonia clathrata (Stimpson, 1865). Basin of the Colorado Desert (Taylor [1966] suggested that type material instead came from Muddy River, Clark County, Nevada.) Springs in Moapa and Pahranagat Valleys

(2).

Tryonia elata Hershler and Sada, 1987. Point of Rocks Springs, Nye County, Nevada. Type locality and nearby springs, Ash Meadows, Amargosa River drainage (2).

16 Table 1, continued.

Tryonia ericae Hershler and Sada, 1987. "North Scruggs Spring," Nye County, Nevada. Type locality and and nearby spring, Ash Meadows, Amargosa River drainage (2).

Tryonia gilm Taylor, 1987. Unnamed spring on north side of Gila River about 2 miles north of Bylas,

Graham County, Arizona. Type locality and several nearby springs, Gila River drainage (2).

Tryonia imitator Pilsbry, 1899. Santa Cruz, California. California coast from Salmon Creek to Imperial

Beach (Taylor 1981) (4).

Trvonia kosteri Taylor, 1987. Sago Spring, Chaves County, New Mexico. Type locality and other springs near Roswell, Pecos River drainage (2).

Trvonia mareae Hershler, 1989. Grapevine Spring complex, Inyo County, California. Type locality area in

Death Valley (2).

Tryonia protea (Gould, 1855). Colorado Desert (Gran Jornado). Lower Colorado River drainage, upper

Owens River drainage, Bonneville Basin (Taylor 1985: fig. 35) (6).

Tryonia quitobaquitae Hershler Hershler and Landye, 1988. Quitobaquito Springs, Pima County,

Arizona. Type locality and two nearby springs (2).

Trvonia robusta Hershler, 1989. Nevares Springs, Inyo County, California. Type locaity and nearby

Travertine Springs, Death Valley (2).

Tryonia rowlandsi Hershler, 1989. Grapevine Spring complex, Inyo County, California. Type locality area in

Death Valley (2).

Tryonia sauna Hershler, 1989. Cottonball Marsh, Inyo County, California. Endemic to type locality (1).

Trvonia varieeata Hershler and Sada, 1987. Five Springs, Nye County, Nevada. Ash Meadows (numerous springs) and springs along lower course of Amargosa River (6).

17 0 100 200 300 400 I I I 1 1 I I I I Kilometers

Figure 1. Type locality areas for western American species of Trvonia. 1 Coastal drainage, California

; 2, Death Valley, California (margae, robusta rowlandsi salina). 3, Salton Sea area, California

(protea); 4, Ash Meadows, Nevada(angulata, elata, ericae varie2ata). 5, White River drainage, Nevada

(clathrata); 6, Rio Sonoyta drainage, Arizona (cmitobactuitae); 7, Gila River drainage, Arizona (at); 8, Rio

Grande drainage, New Mexico (alamosae); 9, upper Pecos River drainage, New Mexico (kosteri); 10, lower

Pecos River drainage, Texas (brunei cheatumi); 11, ibid. (adamantina); 12, ibid. (circumstriata). 18 Figure 2. Shells of Tryonia spp. Top row (left to right), adamantina, aequicostata, alamosae, angulata, brevissima; second row, brunei cheatumi, eimumstriata clathrata, data; third row, ericae giloe, imitator kosteri, margae; bottom row, protea Quitobaquitae robusta rowlandsi jj g, variegata. The shell to the uppermost left (adamantina) is 2.3 mm tall; all others are printed to the same scale.

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Mayden, R. L. (ed.). 1992. Systematics, Historical Ecology, and North American Freshwater Fishes. Stanford University Press, 969 pp.

23 Mighels, J. W. 1845. Descriptions of Shells from the Sandwich Islands, and Other Localities. Proceedings of the Boston

Society of Natural History, 2: 18-25. Miller, R. R. 1981. Coevolution of Deserts and Pupfishes (Genus Cyprinodon) in the American Southwest. Pp. 39-94 in Fishes in North American Deserts, R. J. Naiman and D. L. Soltz (eds.), John Wiley and Sons, New York, 552 pp. Minckley, W. L., D. A. Hendrickson and C. E. Bond. 1986.

Geography of Western North American Freshwater Fishes: Description and Relationships to Intracontinental Tectonism.

Pp. 521-613 [+ bibliography] In The Zoogeography of North

American Freshwater Fishes, ibid. Mulvey, M. and R. C. Vrijenhoek. 1981. Genetic Variation among Laboratory Strains of the Planorbid Snail Biomphalaria glabrata. Biochemical Genetics, 19: 1169-1182. Nixon, K. 1992. Clados Version 1.3. K. C. Nixon, L. H. Bailey

Hortorium, Cornell University, Ithaca.

Pilsbry, H. A. 1889. New and Little-known American Mollusks.

No. 1. Proceedings of the Academy of Natural Sciences of

Philadelphia, 41: 81-89.

Pilsbry, H. A. 1890. Notices of New Amnicolidae. Nautilus, 4: 63-64. Pilsbry, H. A. 1891. Preliminary Notices of New Mexican Shells. Nautilus, 5: 8-10.

Pilsbry, H. A. 1899. Catalogue of the Amnicolidae of the

Western United States. Nautilus, 12: 121-127.

Pilsbry, H. A. 1935. Western and Southwestern Amnicolidae and a New Humboldtiana. Nautilus, 48: 91-94.

24 Pilsbry, H. A. and J. H. Ferriss. 1906. of the Southwestern States. II. Proceedings of the Academy of

Natural Sciences, 58: 123-160.

Ponder, W. F. and G. A. Clark. 1988. A Morphological and

Electrophoretic Examination of 'Hydrobia buccinoides,' a Variable Brackish-water Gastropod from Temperate Australia (Mollusca: Hydrobiidae). Australian Journal of Zoology, 36: 661-689.

Ponder, W. F. and G. A. Clark. 1990. A Radiation of Hydrobiid I Snails in Threatened Artesian Springs in Western Queensland. Records of the Australian Museum, 42: 301-363.

Ponder, W. F., D. J. Colgan and G. A. Clark. 1991. The

Morphology, Taxonomy and Genetic Structure of Tatea (Mollusca: Gastropoda: Hydrobiidae), Estuarine Snails from Temperate Australia. Australian Journal of Zoology, 39: 447-497.

Raup, D. M. 1966. Geometric Analysis of Shell Coiling: General

Problems. Journal of Paleontology, 40: 1178-1190.

Ribbink, A. J. 1984. Is the Species Flock Concept Tenable? Pp.

21-25 In Evolution of Fish Species Flocks, ibid.

Rogers, J. S. 1972. Measures of Genetic Similarity and Genetic Distance. University of Texas Publication, Studies in Genetics, 7: 145-153.

Rogers, J. S. 1984. Deriving Phylogenetic Trees from Allele Frequencies. Systematic Zoology, 33: 52-63.

Sanderson, M. J. and M. J. Donoghue. 1989. Patterns of

Variation in Levels of Homplasy. Evolution, 43: 1781-1795.

25 Smith, G. R. 1978. Biogeography of Intermountain Fishes. Pp. 17-42 in Intermountain Biogeography: A Symposium, K. T. Harper and J. L. Reveal (eds.), Great Basin Naturalist Memoirs, 2, 268 pp. Smith, M. L. and R. R. Miller. 1986. The Evolution of the Rio Grande as Inferred from its Fish Fauna. Pp. 457-485 [+ bibliography] In The Zoogeography of North American Freshwater Fishes, ibid.

Stimpson, W. 1865. Diagnoses of Newly Discovered Genera of Gasteropods, Belonging to the Sub-fam. Hydrobiinae, of the Family Rissoidae. American Journal of Conchology, 1: 52-54. Swofford, D. L. 1981. On the Utility of the Distance Wagner

Procedure. Pp. 25-43 In Advances in Cladistics: Proceedings

of the First Meeting of the Willi Hennig Society, V. A. Funk

and D. R. Brooks (eds.), New York Botanical Garden, Bronx, 250 pp. Swofford, D. L. 1990. PAUP. Phylogenetic Analysis Using Parsimony, Version 3.0. Illinois Natural History Survey,

Urbana.

Swofford, D. L. and R. Selander. 1981. BIOSYS-1: a FORTRAN

Program for the Comprehensive Analysis of Electrophoretic

Data in Population Genetics and Systematics. Journal of

Heredity, 72: 281-283. Taylor, D. W. 1960. Distribution of the Freshwater Clam

Pisidium ultramontanum. American Journal of Science, 258A: 325-334. Taylor, D. W. 1966a. Summary of North American Blancan

Nonmarine Mollusks. Malacologia, 4: 1-172.

26 Taylor, D. W. 1966b. A Remarkable Snail Fauna from Coahuila,

Mexico. Veliger, 9: 152-228. Taylor, D. W. 1981. Freshwater Mollusks of California: A Distributional Checklist. California Fish and Game, 67:

140-163. Taylor, D. W. 1985. Evolution of Freshwater Drainages and Molluscs in Western North America. Pp. 265-321 in C.J. Smiley (ed.), Late Cenozoic History of the Pacific

Northwest, American Association for the Advancement of

Science, San Francisco, 417 pp. Taylor, D. W. 1987. Fresh-water Molluscs from New Mexico and Vicinity. New Mexico Bureau of Mines & Mineral Resources,

Bulletin, 116: 1-50. Taylor, D. W. 1988. Aspects of Freshwater Mollusc Ecological

Biogeography. Palaeogeography, Palaeoclimatology,

Palaeoecology, 62: 511-576. Taylor, D. W. and R. C. Bright. 1987. Drainage History of the Bonneville Basin. Pp. 239-256 In Cenozoic Geology of Western Utah, R. S. Kopp and R. E. Cohenour (eds.), Utah

Geological Association Publication 16, Brigham Young

University Press, Provo, 684 pp.

Thompson, F. G. 1968. The Aquatic Snails of the Family

Hydrobiidae of Peninsular Florida. University of Florida

Press, Gainesville, 268 pp.

27 • United States Department of the Interior (USDI). 1991a. Endangered and Threatened Wildlife and Plants; Final Rule to

List the and the as Endangered. Federal Register, 56: 49646-49649.

United States Department of the Interior (USDI). 1991b. Endangered and Threatened Wildlife and Plants;

Candidate Review for Listing as Endangered or Threatened Species. Federal Register, 56: 58804-58836.

Wiley, E. 0. 1981. Phylogenetics. The Theory and Practice of

Phylogenetic Systematics. John Wiley and Sons, New York,

439 pp.

28 BUDGET

Year 1 Year 2 Total

Salary:

Hershler Salaried by Smithsonian Assistant 3,000 3,000 6,000 Mulvey 3,200 3,200 6,400

12,400

Travel costs:

Airfare 1,402 386 1,788 Per diem (43 days @ $66) 2,508 330 2,838 Truck rental (38 days @ $50) 1,900 ___ 1,900 Car rental (5 days @ $25) --- 125 125 Miscellaneous 250 250

6,901

Supplies and Materials

Chemicals (1600 snails @ $3.90) 2,080 4,160 6,240 Microfuge tubes 50 50 Film and processing 250 125 375

6,665

Total Year 1 $14,640 Total Year 2 11,326

TOTAL $25,966 BUDGET EXPLANATION

Summer contracts (1 month for each year) are requested to support Mulvey's contribution to the project at a level equivalent to her current salary ($3,200 per month). Modest sums

($3K/year in contract form) also are requested for support of Hershler's laboratory studies, specifically involving measuring shells, preparation of material for scanning electron microscopy, and data entry.

Roundtrip airfares for fieldwork are to Orlando (FL), San Diego (CA), and Tucson (AZ) during Year 1. Details of the

fieldtrips, which total 38 days, are given above. Use of a 4WD truck for fieldwork is preferred, given prevalence of rough terrain in most of the collection areas. Miscellaneous expenses will cover purchase of dry ice and shipping of frozen samples back to Georgia. A short (five day) trip to Augusta (SC) during

Year 2 (RT Airfare, car rental, per diem), enabling the

investigators to work together on data analysis and manuscript

preparation, also is requested. The component of this research pertaining to establishment

of buffer and stain combinations for electrophoresis, and genetic study of species (totalling 19 populations) from the Death Valley region has been facilitated by funding ($16,219) recently

provided by the Office of the Director, National Museum of Natural History. Only an additional 40 populations (totalling

1,600 snails) require assay, with one-third of this planned for the first year and two-thirds for the second year. Costs of supplies for electrophoretic analyses are calculated at $3.90 per

30 specimen based on previous projects performed by Mulvey. The supplies requested are only those that will be consumed during the analysis (biochemical reagents, starch, film, etc.).

31

411) 1 7 8 5 The University of Georgia (803) 725-2472 Drawer E FTS 239-2472 Aiken, SC 29802 FAX 803-725-3309 Savannah River Ecology Laboratory

August 25, 1993

Mr. David R. Short Office of Sponsored Projects National Museum of Natural History Smithsonian Institution Washington, DC 20560

Dear Mr. Short: I am writing to convey the support of SREL for the research proposal, "Evolution of the North American Springsnail Tyronia". It is an important and exciting project, and we look forward to being involved in the work. It is a natural extension of the type of work that we have conducted in the laboratory. I am sure that the work when completed and published will influence the approach to similar types of problems and the thinking in this area of research. The facilities of our ecological genetics laboratory will be available for the completion of this research.

Sincerely,

jWO/fi 1^^^}1L Michael H. Smith Director

cc: Dr. Peg Mulvey File

32

An Equal Opportunity/Affirmative Action Institution Previous Scholarly Studies Funding

1988-1989. "Generic Review of Aquatic Snails of the Subfamily Littoridininae (Gastropoda: Hydrobiidae)," $42,977. Funding supported fieldwork enabling acquisition of materials for a review of the 31 genera of this predominantly Neotropical

group of fresh- and brackish-water snails. Products of this

effort were descriptions of various new taxa arising from

the study (Thompson and Hershler 1991), and a more significant companion paper reviewing the genera of this subfamily and providing speculative phylogenetic and

biogeographic hypotheses for the group (Hershler and

Thompson 1992). The latter included clarification of a large Central and South American fauna (19 described genera)

that had been little known previously and stood as a major

stumbling block to an understanding of the higher

systematics of the huge family Hydrobiidae.

1990-1993. "Biogeography of Great Basin Springsnails," $38,500, 1-year award; $40,000, 2-year award. Funding supported a

field survey of springsnails (family Hydrobiidae) throughout

the Great Basin of the western United States, which will yield material for a systematic monograph of this large, but poorly known fauna and, more importantly, a biogeographic database for study of snail distribution and evolution in relation to a historically complex drainage involving over

90 basins. Fieldwork will be completed by November 1, 1993. The first two years encompassed survey of the State of

33 Nevada, and yielded over 200 collections of snails, almost all of which represent new records. Snails were found in more than half of Nevada's 70+ basins, and many new, localized species were discovered, both suggesting that a fruitful biogeographic database will be obtained. Additional funding ($15,000) was provided in the form of a

Cooperative Agreement with the Bureau of Land Management, which will benefit greatly from the results of the study as they pertain to conservation and management of imperiled aquatic ecosystems and biota in the West.

34 CURRICULUM VITAE

Robert Hershler Department of Invertebrate Zoology, NHB STOP 118 National Museum of Natural History Smithsonian Institution Washington, D.C. 20560 phone (202) 786-2077 FAX (202) 357-2343 BITNET mmivoo7@sivm

EDUCATION B.S. 1975 SUNY at Stony Brook Biology M.S. 1980 John Hopkins University Ecology and Evolution Ph.D. 1983 Johns Hopkins University Ecology and Evolution

PROFESSIONAL EXPERIENCE 1985-Present Smithsonian Institution, Washington, D.C. Associate Curator 1984-1985. Southwest Texas State University, San Marcos, TX Research Associate 1983-1984. Australian Museum, Sydney, NSW Consulting Research Biologist

AWARDS AND CONTRACTS (since coming to Smithsonian Institution) 1987. Status survey of Hydrobiidae in Owens River drainage, $9,995. California Department of Fish and Game. 1988. Generic review of aquatic snails of the subfamily Littoridininae (Gastropoda: Hydrobiidae), $42,977. Smithsonian Institution (Scholarly Studies Program). [2 years] 1988. Status assessment of the Alabama Live-bearing snail, Tulotoma magnifica, $4,500. United States Fish and Wildlife Service [Cooperative Agreement] 1988. Status survey of Hydrobiidae in Mojave Desert and selected areas adjacent to Death Valley Hydrographic System, $16,258. California Department of Fish and Game. 1989. Survey of springsnails of Great Basin, northern California, $25,894. California Department of Fish and Game. [2 years] 1990. Status Survey of the Grapevine Creek Snail, $3,069. California Department of Parks and Recreation.

35 1990. Biogeography of Great Basin Springsnails, $38,500. Smithsonian Institution (Scholarly Studies Program).

1991. Biogeography of Great Basin Springsnails, $40,000. Smithsonian Institution (Scholarly Studies Program). [2 years]

1991. Biogeography of Great Basin Springsnails, $10,000. Bureau of Land Management. [Cooperative Agreement, 2 years]

1992. Taxonomic status of Tryonia in Long Valley, Mono County, California, $5,000. California Department of Fish and Game.

1993. California Springsnails: Taxonomic Clarification and Threatened/Endangered Status of Selected Taxa, $10,000. California Department of Fish and Game. [2 years]

PUBLICATIONS

Hershler, R. & G.M. Davis. 1980. The Morphology of Hvdrobia truncata: Relevance to Systematics of Hydrobia. Biological Bulletin 158: 195-219.

Hershler, R. 1984. Hydrobiid Snails of the Cuatro Cienegas Basin, Mexico: Systematic Relationships and Ecology of a Unique Fauna. IA: Biota of Cuatro Cienegas, Coahuila, Mexico, P. Marsh, editor, Journal of the Arizona-Nevada Academy of Sciences 19: 61-76.

Hershler, R. 1985. Systematic Revision of the Hydrobiidae (Gastropoda: Rissoacea) of the Cuatro Cienegas Basin, Coahuila, Mexico. Malacologia 26: 31-123.

Hershler, R. & G. Longley. 1986. Phreatic Hydrobiids (Gastropoda: Prosobranchia) from the Edwards (Balcones Fault Zone) Aquifer Region, South-Central Texas. Malacologia 27: 127-172.

Hershler, R. & G. Longley. 1986. Hadoceras taylori, a New Genus and Species of Phreatic Hydrobiidae (Gastropoda: Rissoacea) from South-central Texas. Proceedings of the Biological Society Of Washington 99: 121-136.

Hershler, R. & W.L. Minckley. 1986. Microgeographic Variation in the Banded Spring Snail (Hydrobiidae: Mexipyrgus) from the Cuatro Cienegas Basin, Coahuila, Mexico. Malacologia 27: 357-374.

Hershler, R. 1987. Redescription of Assiminea infima Berry 1947 from Death Valley, California. Veliger 29: 274-288.

36 Hershler, R. & F.G. Thompson. 1987. North American Hydrobiidae (Gastropoda: Rissoacea): Redescription and Systematic Relationships of Tryonia Stimpson, 1865 and Pyrgulopsis Call and Pilsbry, 1886. Nautilus 101: 25-32.

Hershler, R. & G. Longley. 1987. Phreatoceras, a New Name for Hadoceras Hershler and Longley, 1986 (Gastropoda) non Strand, 1934 (Cephalopoda). Proceedings of the Biological Society of Washington 100: 402.

Hershler, R. & G. Longley. 1987. Phreatodrobia coronae, a New Species of Cavesnail from Southwestern Texas. Nautilus 101: 133-139.

Hershler, R. & D. Sada. 1987. Springsnails (Gastropoda: Hydrobiidae) of Ash Meadows, Amargosa Basin, California- Nevada. Proceedings of the Biological Society of Washington 100: 776-843.

Hershler, R. & J. Landye. 1988. Arizona Hydrobiidae. Smithsonian Contributions to Zoology 459: 1-63.

Hershler, R. & L. Hayek. 1988. Shell Variation of Springsnail Populations in the Cuatro Cienegas Basin, Mexico: Preliminary Analysis of Limnocrene Fauna. Nautilus 102: 56- 64.

Hershler, R. & L. Hubricht. 1988. Notes on Antroselates Hubricht, 1963 and Antrobia Hubricht, 1971 (Gastropoda: Hydrobiidae). Proceedings of the Biological Society of Washington 101: 730-740.

Hershler, R. & F.G. Thompson. 1988. Notes on Morphology of Amnicola limosa (Say, 1817)(Gastropoda: Hydrobiidae) with Comments on Status of the Subfamily Amnicolinae. Malacological Review 22: 81-92.

Hershler, R. 1988. Holsingeria unthanksensis, a New Genus and Species of Aquatic Cavesnail from Eastern North America. Malacological Review 22: 93-100.

Hershler, R. 1989. Springsnails (Gastropoda: Hydrobiidae) of Owens and Amargosa River (Exclusive of Ash Meadows) Drainages, Death Valley System, California-Nevada. Proceedings of the Biological Society of Washington 102: 176-248.

Ponder, W.F., R. Hershler, & B.J. Jenkins. 1989. An Endemic Radiation of Hydrobiid Snails from Artesian Springs in Northern South Australia: Their Taxonomy, Physiology, Distribution and Anatomy. Malacologia 31: 1-140.

37 Hershler, R. & F.G. Thompson. 1990. Antrorbis breweri, a New Genus and Species of Hydrodrobiid Cavesnail from Coosa River Basin, Northeastern Alabama. Proceedings of the Biological Society of Washington 103: 197-204.

Hershler, R. & W.L. Pratt. 1990. A New Pyrgulopsis (Gastropoda: Hydrobiidae) from Southeastern California, with a Model for Historical Development of the Death Valley Hydrographic System. Proceedings of the Biological Society of Washington 103: 279-299.

Hershler, R. & J.R. Holsinger. 1990. Zoogeography of Nort American Hydrobiid Cavesnails. Stygologia 5: 5-16.

Hershler, R., J.R. Holsinger, & L. Hubricht. 1990. A Revision of the North American Freshwater Snail Genus Fontigens (Prosobranchia: Hydrobiidae). Smithsonian Contributions to Zoology 509: 1-49.

Hershler, R. 1990. Pyrgulopsis bruneauensis, a New Springsnail (Gastropoda: Hydrobiidae) from the Snake River Plain, Southern Idaho. Proceedings of the Biological Society of Washington 103: 803-814.

Hershler, R., J.M. Pierson, & R.S. Krotzer. 1990. Rediscovery of Tulotoma magnifica (Conrad)(Gastropoda: Viviparidae). Proceedings of the Biological Society of Washington 103: 815-824.

Thompson, F.G. & R. Hershler. 1991. Two New Hydrobiid Snails (Amnicolinae) from Florida and Georgia, with a Discussion of the Biogeography of Freshwater Gastropods of South Georgia Streams. Malacological Review 24: 55-72.

Thompson, F.G. & R. Hershler. 1991. New Hydrobiid Snails (Mollusca: Gastropoda: Prosobranchia: ) from North America. Proceedings of the Biological Society of Washington 104: 669-683.

Hershler, R. & F.G. Thompson. 1992. A Review of the Aquatic Gastropod Subfamily Cochliopinae (Prosobranchia: Hydrobiidae). Malacological Review, Supplement 5: 1-140.

Hershler, R. & F. Velkovrh. 1993. A New Genus of Hydrobiid Snails (Mollusca: Gastropoda: Prosobranchia: Rissooidea) from Northern South America. Proceedings of the Biological Society of Washington 106: 182-189

Kabat, A. & R. Hershler. 1993. The Prosobranch Snail Family Hydrobiidae (Gastropoda: Rissooidea): Review of Classification and Supraspecific Taxa. Smithsonian Contributions to Zoology. In Press (394 MS pp.).

38 Hershler, R. 1993. A Review of the North American Freshwater Snail genus Pyrgulopsis (Hydrobiidae). Smithsonian Contributions to Zoology (316 MS pp.). Hershler, R., T.J. Frest, E.J. Johannes, P.A. Bowler & F.G. Thompson. 1993. Two New Genera of Hydrobiid Snails (Prosobranchia: Rissooidea) from the Northwestern United States. Veliger. In Press (56 MS pp.)

INVITED LECTURES

1982. Hydrobiid snails of the Cuatro Cienegas Basin: Systematic Relationships and Ecology of a Unique Fauna. Desert Fishes Council Annual Meeting (In symposium, Biota of Cuatro Cienegas, Coahuila, Tempe, AZ. 1985. Phreatic gastropods from south-central Texas. American Malacological Union Annual Meeting (In symposium, Perspectives in Malacology), Kingston, RI.

1986. Arizona Hydrobiidae: Morphometrics and Systematics. American Malacological Union Annual Meeting (In symposium, Molluscan morphological analysis), Monterey, CA.

1989. Groundwater Hydrobiidae of North America. UNITAS MALACOLOGICA, Tenth International Malacological Congress (In workshop, Hydrobiid Phylogeny), Tubingen, F.R.G..

39 MARGARET MULVEY

617 Boardman Road Savannah River Ecology Laboratory Aiken, South Carolina 29803 Drawer E (803)648-2732 Aiken, South Carolina 29801 SS no. 041-48-1149 (803)725-2752

EDUCATION

1974 B.A. University of Connecticut, Storrs, Connecticut 1976 M.S. University of Connecticut, Storrs, Connecticut 1981 Ph.D. Rutgers University, New Brunswick, New Jersey

POSITIONS

Associate Research Ecologist: University of Georgia, Institute of Ecology and Savannah River Ecology Laboratory, 1992-present Adjunct Assistant Professor: Wake Forest University and Savannah River Ecology Laboratory, 1983-1992

USPHS Postdoctoral Fellow: University of California, San Diego, 1982-1983

TEACHING EXPERIENCE

Instructor, Rutgers University

General Ecology Part-time faculty, University of South Carolina at Aiken Man and The Environment Genetics and Society General Biology--Laboratory Part-time faculty, Limestone College, Gaffney, South Carolina

Environmental Science

Associate professor, University of Georgia General Biology

40 PROFESSIONAL AFFILIATIONS

American Genetic Association American Malacological Union Carolinas-Society of Environmental Toxicology and Chemistry Society for the Study of Evolution AWARDS Leslie Stauber Award, 1977 Leathem-Steinetz Research Award, 1979, 1980, 1981 Charles and Johanna Busch Predoctoral Fellowship, 1979-1980 RESEARCH FUNDING

NSF (Systematics) Genetic variation and systematics of Biompha- laria (Gastropoda: Planorbidae). $10,000 (Co-PI with David S. Woodruff, 1983)

NSF (Genetic Biology Program) Genetics of host-parasite compati- bility: snail resistance to a trematode. $70,000 (Co-PI with David Woodruff, 1983) DOE (subcontract to Wake Forest University) Ecological genetics of Gambusia holbrooki and Fascioloides magna populations of the Savannah River Site. approximately $35,000 annually (1984-present)

NATO Collaborative Research Grant. Population structure and adaptation by Helix aspersa to lead contamination. BF 124,000.00 (CO-PI with M.C. Newman, 1990)

PROFESSIONAL ACTIVITIES

NSF Panel, 1992, Dissertation Improvement

Nature Conservancy Workshop on Endangered Atlantic Slope Molluscs Attended: "Pathobiology of Marine Invertebrates", Marine Biological Laboratory, Woods Hole Manuscripts and research proposals reviewed for: National Science Foundation, National Institutes of Health, Evolution, Malacologia, J. Parasitology, J. Heredity

Faculty Search Committee, SREL, Stress and Wildlife Division

Organized Workshop on Fish Research at SREL

41 PRESENTATION OF RESEARCH

1993 World Congress Society for Environmnetal Toxicology and Chemistry, Lisbon, Portugal

1992 Insitiute of Ecology, University of Georgia, Athens, GA American Malacological Union, Sarasota, FL Conservation and Management of Freshwater Mussels, St. Louis, MO

1991 Biology Department, University of New Mexico, Albuquerque, NM Savannah River Ecology Laboratory, Aiken, SC

1990 Biology Department, Wake Forest University, Winston-Salem, NC Biology Department, College of Charleston, Charleston, SC, 1990

1981-1989 Society for the Study of Evolution, Asilomar, CA Society for the Study of Evolution, Bozeman, MT Southeastern Ecological Genetics Group, Beaufort Society for the Study of Evolution, Durham, NH Southeastern Ecological Genetics Group, Georgetown, SC Biology Department, University of California, San Diego, CA Biology Department, Oregon State University, Corvallis, OR International Symposium for Molluscan Genetics, New Orleans, LA American Malacological Union, Ft. Lauderdale, FL, 1981 National Shellfish Association, Williamsburg, VA, 1981 Zoology Department, Rutgers University, New Brunswick, NJ, 1981

1978-1980 New Jersey Society of Parasitologists, Edison, NJ Bureau of Biological Research, Rutgers University, New Brunswick, NJ New Jersey Academy of Sciences, Stockton State College, Pomona, NJ Bureau of Biological Research, Rutgers University, New Brunswick, NJ

PUBLICATIONS

Mulvey, M. and S.Y. Feng. 1981. Hemolymph constituents of normal and Proctoeces maculatus-infected Mytilus edulis. Comp. Biochem. Physiol. 70A:119-125.

Mulvey, M. and R.C. Vrijenhoek. 1981. Multiple paternity in the hermaphroditic snail Biomphalaria obstructa. Jour. Heredity. 72:308-312.

Mulvey, M. and R.C. Vrijenhoek. 1981. Genetic variation among laboratory strains of the planorbid snail Biomphalaria glabrata. Biochem. Genetics. 19:1169-1182.

42 Mulvey, M. and R.C. Vrijenhoek. 1982. Population structure in Biomphalaria alabrata:examination of an hypothesis for the patchy distribution of susceptibility to schistosomes. Am Jour. Trop. Med. Hyg. 31:1195-1200.

Woodruff, D.S., M. Mulvey, W.B. Saunders and M.P. Carpenter. 1983. Genetic variation in the cephalopod Nautilus balauensis. Proc. Acad. Nat. Sci. Phil. 135:147-153.

Mulvey, M. and R.C. Vrijenhoek. 1984. Genetics of Biomphalaria alabrata: linkage analysis and crossing compatibilities among laboratory strains. Malacologia. 25:513-524.

Woodruff, D.S., M. Mulvey and M. Yipp. 1985. The continued intro- duction of intermediate host snails of Schistosoma mansoni into Hong Kong. Bull. Wld. Hlth. Org. 63:621-622.

Woodruff, D.S., M. Mulvey and M. Yipp. 1985. Observations on the population genetics of Biomphalaria ptraminea in Hong Kong: a neotropical schistosome-transmitting snail recently introduced to China. Jour. heredity.76:355-360.

Mulvey, M. and D.S. Woodruff. 1985. Genetics of Biomphalaria alabrata: linkage analysis of genes for pigmentation, enzymes and resistance to infection with Schistosoma mansoni. Biochem. Genet- ics. 22:877-889.

Woodruff, D.S., L. McMeekin, M. Mulvey and M.P. Carpenter. 1986. Population genetics of Crepidula onyx: variation in a California slipper limpet recently introduced into China. Veliger. 29:53-63.

Mulvey, M., G.W. Esch, T. Goater and A. Crews. 1987. Allozyme frequency differences in Halipegus occidualis-infected and nonin- fected Helisoma anceps. Jour. Parasitology. 73:575-561.

Mulvey, M., D.S. Woodruff and M.P. Carpenter. 1988. Joint segre- gation of loci controlling enzymes and pigmentation in the snail Biomphalaria alabrata. Jour. Heredity. 79:473-476.

Mulvey, M., M.C. Newman and D.S. Woodruff. 1988. Genetic differentiation among West Indian populations of the schistosome- transmitting snail Biomphalaria alabrata. Malacology. 29:309-317.

Lydeard, C., M. Mulvey, J.M. Aho and P.K. Kennedy. 1989. Genetic variability among natural populations of the liver fluke, Fascio- loides magna, in white-tailed deer, Odocoileus virginianus. Can. Jour. Zool. 67:2021-2025.

Diamond, S.A., M.C. Newman, M. Mulvey, P. Dixon and D. Martinson. 1989. Allozyme genotype and time to death of mosquitofish, Gambu- sia affinis (Baird and Girard) during acute toxicant exposure: inorganic mercury. Environ. Toxicol. Chem. 8:613-622.

43 Newman, M.C., S.A. Diamond, M. Mulvey and P. Dixon. 1989. Allo- zyme genotype and time to death of mosquitofish, Gambusia affinis (Baird and Girard) during acute toxicant exposure: a comparison of arsenate and inorganic mercury. Aquatic Toxicol. 15:141-156. Travis, J., J.C. Trexler and M. Mulvey. 1990. Multiple paternity and its correlates in female Poecilia latipinna (Pisces, Poe- ciliidae). Copeia. 1990:722-729. Goater, T.P., M. Mulvey and G.W. Esch. 1990. Electrophoretic differentiation of two Halipegus (Trematoda: Hemiuridae) congen- ers in an amphibian population. Jour. Parasitol. 76:431-434. Bandoni, S.M., M. Mulvey, D.K. Koech and E.S. Loker. 1990. Genet- ic structure of Kenyan populations of Biomphalaria Dfeifferi (Gastropoda: Planorbidae). Jour. Molluscan Studies. 56:383-391. Mulvey, M., J.M. Aho, C. Lydeard, P.L. Leberg and M.H. Smith. 1991. Comparative population structure of the American liver fluke,Fascioloides magna, and its white-tailed deer host. Evolu- tion. 45:1628-1640.

Mulvey, M. and S.A. Diamond. 1991. Genetic factors and tolerance acquisition in populations exposed to metals and metalloids. In Metal Ecotoxicology: Concepts and Applications, eds. M.C. Newman and A.W. McIntosh. Lewis Publishers, Boca Raton, FL. p. 301-316. Diamond, S.A., M.C. Newman, M. Mulvey and S.I. Guttman. 1991. Allozyme genotype and time-to-death of mosquitofish, Gambusia holbrooki, during acute inorganic mercury exposure: a comparison of populations. Aquatic Toxicol. 21:119-134. Kramer, V.J., M.C. Newman, M. Mulvey and G.R. Ultsch. 1992. Glycolysis and Krebs cycle metabolites in mosquitofish, Gambusia holbrooki, Girard 1859, exposed to mercuric chloride: allozyme genotype effects. Environ. Toxicol. Chem. 11:357-364. Lee, C.J., M.C. Newman and M. Mulvey. 1992. Time to death of mosquitofish (Gambusia holbrooki) during acute inorganic mercury exposure: population structure effects. Arch. Environm. Contain. Toxicol. 22:284-287. Heagler, M.G. and M. Mulvey. 1992. Time effects on enzyme activity in the mosquitofish, Gambusia holbrooki. Environ. Toxicol. Chem. 11:605-607. Aho, J.M., M. Mulvey, K. Jacobson and G.W. Esch. 1992. Genetic variability and differentiation among congeneric acanthocephalans. J. Parasitol. 78:974-981. Heagler, M.G., M.C. Newman and M. Mulvey and P.M. Dixon. 1993. Allozyme genotype shifts in mosquitofish, Gambusia holbrooki, during mercury exposure: temporal stability, concentration effects and field verification. Environ. Toxicol. Chem. 12:385- 395.

44 Mulvey, M. and J.M. Aho. 1993. Traits associated with mate competition in uninfected and Fascioloides pagna-infected male white-tailed deer. Oikos 66:187-192. Mulvey, M. and S.M. Bandoni. Genetic variation in the M-line stock of Biomphalaria glabrata (Mollusca: Pulmonata). J. Helminth. Soc. Wash. (in press) Mulvey, M., G.P. Keller and G. K. Meffe. Single-and multiple- locus genotypes and life history responses of Gambusia bolbrooki under thermally ambient and stressful conditions. Evolution. (in press)

MANUSCRIPTS SUBMITTED Woodruff, D.S. and M. Mulvey. Neotropical schistosomiasis: African affinities of the snail Diomphalaria alabrata. (J. Heredity) Davis, G.M. and M. Mulvey. Species status of Mill Creek Elliptio. (SRS-NERP Report). M. Mulvey, J.M. Aho and O.E. Rhodes. Parasitism and white-tailed deer: timing and components of female reproduction. (Oikos). Bandoni, S.M., M. Mulvey and E.S. Loker. Phylogenetic analysis of eleven species of Biomphalaria Preston, 1910 (Gastropoda: Planorbidae) based on comparison of allozymes. (Biol. J. Linnean Soc.) Bandoni, S.M., M. Mulvey and E.S. Loker. Intraspecific and interspecific patterns of allozyme variation in Biomphalaria Preston, 1910 (Gastropoda: Planorbidae). (Biochemical Systematics and Ecology). MANUSCRIPTS IN PREPARATION Newman, M.C., M. Mulvey, M.G. Heagler, S. Hales and A. Chazal. Demographic shifts in mosquitofish (Gambusia holbrooki) populations exposed to low concentrations of mercuric chloride. (for Environ. Toxicol. Chem.) Mulvey, M., J.M. Aho and M.H. Smith. Genetic variation and incidence of the liver fluke, Fascioloides magna, in white-tailed deer. (for J. Wildlife Management). Keklak, M.M., M.C. Newman and M. Mulvey. Enhanced uranium tolerance of an exposed population of the Eastern mosquitofish (Gambusia holbrooki Girard 1859). (for Arch. Environ. Contam. Toxicol.)

45 STUDENTS

Susan M. Bandoni, University of New Mexico, Albuquerque, New Mexico, Ph.D. committee member

Mary M. Davis, University of South Carolina, Aiken, South Carolina, student research at SREL

Gregory Edgar, University of New Mexico, Albuquerque, New Mexico, undergraduate summer research at SREL

Jackie Fernandez, Wake Forest University, Winston-Salem, North Carolina, Ph.D. student research at SREL and WFU

Carl Freeman, Bethel College, McKenzie, Tennessee, undergraduate summer research program

Timothy Goater, Wake Forest University, Winston-Salem, North Carolina, Ph.D. student research at SREL Mary G. Heagler, Rutgers University, New Brunswick, New Jersey, Ph.D. committee member

William Scott Monks, University of Nebraska, Lincoln, Nebraska, Ph.D. student research at SREL

Sonya Piland, University of Georgia, Athens, Georgia, M.S. committee member

Dennis Stiegerwalt, Gallatin R V High School, Gallatin Missouri, Teacher Research Associate at SREL

Amy Wethington, College of Charleston, Charleston, South Carolina, M.S. committee member Marty Wolfson, Academy High School, Fort Pierce, Florida, Teacher Research Associate at SREL

46 RESEARCH AND TEACHING INTERESTS

My major research interests involve the evolutionary aspects of host-parasite relationships and population responses to environmental stress. I have worked on a variety of population level problems including the following: population structure of molluscan hosts of human schistosomiasis, multiple paternity in molluscs and fish and responses of molluscs and fish to parasitism and environmental stress. The following is a brief summary of ongoing research efforts.

1. Genetic population structure in host-parasite relationships

The American liver fluke, Fascioloides magna, and its white-tailed deer host have been studies on the Savannah River Site for five years. We are examining factors underlying population structure in host and parasite, demographic patterns of parasitism and effects of parasite burdens on host condition indices. These data provide evidence to support tt the parasite burden-mate competition hypothesis put forward to describe the impact of parasitism on host sexual selection.

2. Genetic population structure of molluscs

Molluscs exhibit a variety of reproductive patterns and life history characteristics which make them ideal for the study of factors influencing the genetic structure of natural populations. Ongoing studies focus on freshwater mussels, Elliptio sp., and factors influencing microgeographic population structure. Additionally, as this genus is defined largely on shell morphology we are using protein electrophoretic methods to define genetic differentiation within and among populations and species in this genus.

3. Population response to environmental stress

These studies focus on the mosquitofish, Gambusia holbrooki, and its response to thermal and heavy metal stress. We are particularly interested in factors influencing differential individual response to environmental perturbation. We have described size, sex and genetic factors contributing to differential response to stress. Additionally, with Dr. Michael C. Newman (SREL)and Dr. Alan Beeby (Polytechnic, London) studies are underway to assess the responses of the land snail, Helix aspersa, to lead contamination. This work combines ecology, toxicology and population genetics to examine differences in individual and population response to this heavy metal.

I enjoy teaching and have taught genetics and environmental sciences (part-time) for the University of South Carolina-Aiken and Limestone College. I have supervised research projects for graduate students and undergraduate students working at SREL and served as committee member for four graduate students.

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