A Guide for Terrestrial Gastropod Identification - University of ...

A Guide for Terrestrial Gastropod 

Identification 

Editors: Kathryn E. Perez, , James R. (Jay) Cordeiro 

Illustrator: Marla L. Coppolino

American Malacological Society 

Terrestrial Gastropod Identification Workshop 

Editors: 

Kathryn E. Perez, James R. (Jay) Cordeiro 

Illustrator: 

Marla L. Coppolino 

Southern Illinois University, Carbondale, IL 

June 29 - July 3, 2008 

1

Acknowledgements & Sponsors 

For providing financial support for this workbook and workshop we would like to thank Illinois 

Department of Natural Resources, Division of Natural Heritage, Lawrence L. Master, and 

NatureServe. 

For permission to reproduce figures and distribution of How to know the Eastern Land Snails to 

workshop participants we would like to thank John B. Burch. 

Frank E. (Andy) Anderson provided logistics and support for the entire meeting and we are most 

appreciative. 

Workbook Contributors 

John B. Burch, Mollusk Division, Museum of Zoology, University of Michigan, Ann Arbor, 

MI 48109-1079, jbburch@umich.edu . 

Marla L. Coppolino, Department of Zoology, Mailcode 6501, Southern Illinois University, 

Carbondale, IL 62901-6501, USA, http://mypage.siu.edu/mlcopp/, mlcopp@siu.edu. 

James R. (Jay) Cordeiro, Conservation Science/Zoology, NatureServe, 11 Avenue de 

Lafayette, 5th Floor, Boston, MA 02111, jay_cordeiro@natureserve.org 

Jochen Gerber, Zoology Department, Field Museum of Natural History, 1400 S. Lake Shore 

Dr, Chicago, IL 60605-2496, jgerber@fieldmuseum.org. 

Jeffrey C. Nekola, Biology Department, Castetter Hall, University of New Mexico, 

Albuquerque, NM 87131, jnekola@unm.edu, http://sev.lternet.edu/~jnekola. 

Aydin Örstan, Section of Mollusks, Carnegie Museum of Natural History, 4400 Forbes Ave, 

Pittsburgh, PA 15213-4080, pulmonate@earthlink.net. 

Megan E. Paustian, BEES Department, 2239 Bio/Psych Building, University of Maryland, 

College Park, MD 20742, mpaustia@umd.edu. 

Kathryn E. Perez, Department of Biology, Duke University, Box 90338, Durham, NC 27708, 

perezke@gmail.com, www.unc.edu/~keperez. 

Mark A. Ports, Life Sciences Department, Great Basin College, 1500 College Parkway, Elko, 

NV 89801, markp@gwmail.gbcnv.edu. 

Kevin J. Roe, Department of Natural Resources Ecology and Management, 339 Science II, Iowa 

State University, Ames, IA 50010-3221, kjroe@iastate.edu 

Barry Roth, 745 Cole Street, San Francisco, CA 94117, barry_roth@yahoo.com. 

Amy S. Van Devender, Boone, NC, asvande@hotmail.com. 

2

Table of Contents 

I. Glossary of Land Snail Terminology 

II. Global Heritage Rank criteria 

III. Shell Growth 

IV. Land Snail Collection Strategies 

V. Overview of North American Diversity 

1. Agriolimacidae 

2. Annulariidae 

3. Arionidae 

4. Bradybaenidae 

5. Bulimulidae 

6. Carychiidae 

7. Cepolidae 

8. Ceriidae 

9. Charopidae 

10. Cionellidae 

11. Discidae 

12. Haplotrematidae 

13. Helicarionidae 

14. Helicinidae 

15. Helicodiscidae 

16. Helminthoglyptidae 

17. Humboldtianidae 

18. Megomphicidae 

19. Oleacinidae 

20. Oreohelicidae 

21. Orthalicidae 

22. Philomycidae 

23. Polygyridae 

24. Pomatiopsidae 

25. Punctidae 

26. Pupillidae 

27. Sagdidae 

28. Spiraxidae 

29. Streptaxidae 

30. Strobilopsidae 

31. Succineidae 

32. Thysanophoridae 

33. Truncatellidae 

34. Urocoptidae 

35. Valloniidae 

36. Veronicellidae 

37. Vitrinidae 

38. Zonitidae 

VI. References 

VII. Species Synonymy for Burch’s (1962) “How to Know the Eastern Land Snails” 

3

Glossarry 

of lannd 

snail terminoology 

(Images: 

©20008 

M.L. CCoppolino) 

Denticcles 

of Pupillloidea 

(after Pilsbry, 19448) 

4

Anterior – directiona al term meanning 

towards the head. 

Aperturee 

– the openin ng of the snaail 

shell, fromm 

which thee 

animal exteends 

its headd 

and foot. 

Apex – thhe 

top or mo ost central wwhorls 

of the snail shell, wwhere 

the eaarliest 

growthh 

occurred. 

Apical – on the top side 

of the shhell, 

oppositee 

the base. 

Axial – rreferring 

to direction d thatt 

is parallel tto 

the colummella; 

opposite 

of spiral. 

Base – thhe 

underside or ventral suurface 

of thee 

shell, oppoosite 

the apiccal 

side. 

Basal – rrefers 

to lower 

or ventrall 

area of the shell or aperrtural 

lip. 

Beehive – a shell sha ape that lookks 

somethingg 

like a tradittional 

beehivve. 

Body whhorl 

– the out termost and largest shelll 

whorl, formmed 

most reccently 

in the snail’s growwth; 

from the aperture to 1.0 whorl baack. 

Bulimoidd 

– a higher-than-wide 

shhell 

shape, ee.g. 

of Bulimmulidae 

Callus – a thickened area of the sshell. 

Columellla 

– the cent tral axis of thhe 

whorls off 

the shell. 

Conical – a type of sh hell shape thhat 

is relativeely 

broad at its basal porrtion, 

with ann 

elongated spire 

that taperrs 

to a point. . 

Depresseed 

heliciform m – a commoon, 

wider-thaan-high 

shell 

shape. 

5

Discoidaal 

– a wider-t than-high, veery 

flattenedd, 

disc-like shhell 

shape of 

certain taxa 

(e.g., 

Helicodisscidae). 

Foot – thhe 

flat muscu ular surface oof 

the snail’ss 

body uponn 

which it craawls. 

Globose 

overall 

– a shell sha ape similar too 

heliciformm, 

but with a higher spiree 

and more roounded 

shappe 

Growth llines 

– transv verse ridges on the shell’s 

surface, fo formed durinng 

the snail’ss 

growth. 

Head – thhe 

anterior region 

of the snail’s bodyy 

that contains 

the tentaccles 

and mouuth. 

Height – the height of o the shell iss 

measured ffrom 

the apeex 

to the mosst 

basal part of the shell, , 

parallel tto 

the colume ella. 

Hirsute – describes a shell that haas 

hairs on thhe 

surface oof 

its periostrracum. 

Keel (of the shell) – a sharp edgee 

of the peripphery 

of the shell presennt 

in certain ttaxa. 

Keel (of the tail) – a ridge on thee 

tail present in certain taaxa. 

Lip or appertural 

lip – the rim of thhe 

aperture oof 

the shell, either sharpp 

or thickened 

dependingg 

upon the species. 

Lirae – raaised 

lines on o the shell’ss 

surface thaat 

run in a spiiral 

directionn. 

Malacoloogy 

– the stu udy of molluusks. 

A scienntist 

who stuudies 

molluskks 

is called a malacologist. 

Mantle – one of the features f that defines the phylum Molllusca 

as a taaxon; 

this is a tissue thatt 

covers thhe 

visceral or rgans of the animal. In ssnails, 

it is loocated 

withiin 

the shell, aand 

only thee 

edge of thhe 

mantle is observable at the apertuure. 

For sluggs, 

the mantlle 

is located dorsally behhind 

6

the head, , forming a slight s hump (in Arionidaae, 

Argiolimaacidae 

and LLimacidae 

sllugs), 

or covvers 

the lengthh 

of the body y (in Philommycidae 

slugs). 

Microscuulpture 

– des scribes any textural 

featuures 

of the shhell’s 

surface 

usually as seen with thhe 

aid of a mmicroscope. 

Operculuum 

– a chitin nous (proteinnaceous) 

or ccalcified 

platte, 

attached to the dorsall 

tail area of f 

operculatte 

snails, tha at, when the snail is withhdrawn 

into iits 

shell, covvers 

some or all of the 

aperture. 

Palatal – 

lip. 

refers to the e area of the outer portioon 

(greatest ddistance 

fromm 

columella) ) of the aperttural 

Parietal ddenticle 

or to ooth – the caalcified 

projeection 

on thee 

body whorl 

within the aperture, preesent 

in certainn 

species. 

Periostraacum 

– the th hin outer covvering 

of thee 

shell, compposed 

of chittinous 

(proteeinaceous) 

material. In certain species s it beaars 

hairs or ttriangular 

sccale-like 

projjections, 

whiile 

in others it is 

smooth. 

Pneumosstome 

– the opening o in thhe 

mantle thhat 

allows airr 

to pass throough 

and leaads 

to the maantle 

cavity wiithin, 

where gas exchangge 

occurs thrrough 

a vasccularized 

reggion 

of the tissue. 

The 

pneumosstome 

can be e seen openinng 

and closinng 

in a land snail when iit 

is active. 

Posteriorr 

– directiona al term meanning 

towardss 

the tail. 

Protoconnch 

– the sma allest, earliest-formed 

wwhorl 

or whorrls 

(developeed 

in embryoonic 

stages) at 

the centeer 

of the shel ll’s coil. 

Ribs – raaised 

ridges on o the shell’ s surface thaat 

run in an aaxial 

or transsverse 

directtion. 

Slug – a snail with a shell that is much reducced 

or absentt. 

Its mantlee 

may be possitioned 

anteriorlyy 

or may cov ver the entiree 

length of thhe 

body, deppending 

on tthe 

taxa (maiinly 

for easteern 

N. Ameriican 

slugs). 

Anterio orly-positionedd 

mantle 

Manttle 

covering lenngth 

of body 

7

Spire – thhe 

top whorl ls of the shelll 

above the last full (boddy) 

whorl 

Spiral – rrefers 

to the direction of f the coil of tthe 

whorls; oopposite 

of aaxial. 

Striae – iindented 

line es on the sheell’s 

surface; ; can be in eiither 

a spirall 

or an axial direction. 

Subcylinndrical 

– refe ers to a shell shape that iss 

higher thann 

wide, as illlustrated 

bellow. 

Striae – iimpressed 

lin nes on the shhell’s 

surface 

that run axxially. 

Succinifoorm 

– the sh hell shape of the Succineidae, 

which is higher thaan 

wide and with a very large 

aperture. Typically the t spire is relatively 

muuch 

shorter aand 

smaller tthan 

the bodyy 

whorl. 

Suture – the seam wh here the shelll’s 

whorls jooin. 

Suturess 

are describeed 

as being iimpressed 

too 

varying ddegrees, 

for example, shallowly 

or ddeeply 

impreessed, 

dependding 

upon thhe 

species. 

Tentacless 

–the sensor ry appendagges 

on the heead 

of a landd 

snail. Therre 

is an uppeer 

pair (also 

called poosterior) 

and a lower pairr 

(also calledd 

anterior; thhese 

are usuaally 

much smmaller), 

bothh 

of 

which aree 

used for ch hemical senssory 

functionn. 

The uppeer 

pair also contains 

the eeyes, 

for moost 

land snaiils 

(some sna ails are blindd, 

for example, 

Helicodisscus 

parallelus). 

In the genus Vertiggo 

only the uupper 

tentac cles bearing tthe 

eyes are present, the lower tentacles 

are abseent. 

Basommmatophoran 

sn nail species such as Caryychium, 

as wwell 

as opercculate 

land snnails, 

have oonly 

one pair of tentacles with the eyees 

at their baases. 

Umbilicuus 

– Opening g in the center 

of the basse 

of the shelll. 

It may bee 

open, with the inside oof 

the 

columella 

visible, or partially cloosed 

by part of the lowerr 

insertion off 

the aperture, 

or compleetely 

closed annd 

appearing g as a depression. 

Its widdth 

is measuured 

at its greeatest 

diameeter, 

with thee 

lower inssertion 

as an n outer point on the diammeter. 

8

Whorl – one revoluti ion of the shell’s 

coil. WWhorls 

are coounted 

from tthe 

earliest wwhorl 

outwaard 

toward thhe 

last and la argest (bodyy 

whorl). 

Width - tthe 

width of the shell is tthe 

maximumm 

measuremment 

as takenn 

across the aaxis 

perpendicular 

to the columella. c 

Globa al Rankk 

Conserrvation 

Status DDefinitioons 

James R. (Jay) CCordeiro, 

NatureSeerve 

NatureServe 

Conser rvation Stattus 

CConservation 

n status information 

– how 

rare or thrreatened 

a sppecies 

is – iss 

a crucial 

componeent 

in setting g priorities annd 

targeting conservation 

efforts. NNatureServe 

aand 

its various 

member pprograms 

ha ave developeed 

a consisteent 

method ffor 

evaluatingg 

the health and conditioon 


species. 

MMany 

things can contribuute 

to the deccline 

and ulttimate 

demisse 

of a speciees. 

The conddition 

of each species 

is ass sessed basedd 

on many crriteria, 

includding 

numberr 

of occurrennces, 

numberr 


occurrences 

with goo od viability, population ssize, 

range eextent, 

area oof 

occupancyy, 

long- and 

short-termm 

population n and/or habbitat 

trend, thhreats, 

numbber 

of protectted 

occurrennces, 

intrinsic 

vulnerability, 

environ nmental speccificity, 

and other considderations. 

CConservationn 

status rankss 

are 

assigned on a one to five scale. GGlobal 

conseervation 

stattus 

levels rannge 

from crittically 

imperriled 

(G1) to secure 

(G5). Listed separrately 

are speecies 

knownn 

to be extincct 

(GX) or thhose 

that aree 

currentlyy 

missing or known histoorically 

and mmay 

be extinnct 

(GH). Sppecies 

rankedd 

in this latteer 

category (GH) are of f highest connservation 

cooncern, 

folloowed 

by rare species classsified 

as 

criticallyy 

imperiled (G G1), imperilled 

(G2), andd 

vulnerablee 

(G3). Ofteen 

a numericc 

range rank (e.g. 

G2G4) iss 

used to ind dicate the rannge 

of uncerttainty 

in the conservation 

status of a species. In 

cases whhere 

informat tion on consservation 

stattus 

is lackingg 

or substantially 

confliccting 

informmation 

exists aboout 

status or r trends, an uunrankable 

(GGU) 

status is 

utilized. TThe 

GNR rannk 

is used foor 

species wwhere 

global l rank has noot 

yet been asssessed. 

A rrank 

qualifieer 

(Q) is usedd 

following a 

numeric conservation n status rankk 

(e.g. G1Q) is used in caases 

where taxonomic 

diistinctivenesss 


this entityy 

at the current 

level is qquestionablee. 

9

Basic Ranks: 

GX- Presumed Extinct: Not located despite intensive searches; virtually no likelihood of 

rediscovery 

GH- Possibly Extinct: Missing; known only from historical occurrences but still some hope of 

rediscovery 

G1- Critically Imperiled: At very high risk of extinction due to extreme rarity (often five or 

fewer populations), steep declines, or other factors. 

G2- Imperiled: At high risk of extinction due to very restricted range, few populations (often 20 

or fewer), steep decline, or other factors. 

G3- Vulnerable: At moderate risk of extinction due to a restricted range, relatively few 

populations (often 80 or fewer), recent and widespread declines, or other factors. 

G4- Apparently Secure: Uncommon but not rare; some cause for long-term concern (e.e. 

widespread declines). 

G5- Secure: Common; widespread and abundant. 

Variant Ranks: 

G#G#- Range Rank: A numeric range rank (e.g., G2G3) is used to indicate the range of 

uncertainty in the status of a species or community. A G2G3 rank would indicate that 

there is a roughly equal chance of G2 or G3 and other ranks are much less likely. Ranges 

cannot skip more than one rank (e.g., GU should be used rather than G1G4). 

GU- Unrankable: Currently unrankable due to lack of information or due to substantially 

conflicting information about status or trends. Whenever possible, the most likely rank is 

assigned and a question mark qualifier may be added (e.g., G2?) to express minor 

uncertainty, or a range rank (e.g., G2G3) may be used to delineate the limits (range) of 

uncertainty. 

GNR- Unranked: Global rank not yet assessed. 

Rank Qualifiers: 

Q- Questionable Taxonomy: Taxonomy distinctiveness of this entity at the current level is 

questionable; resolution of this uncertainty may result in change from a species to a 

subspecies or hybrid, or the inclusion of this taxon in another taxon, with the resulting 

taxon having a lower-priority conservation status. 

U.S. Endangered Species Act Conservation Status 

The U.S. Endangered Species Act (U.S. ESA) is the primary legislation that affords federal legal 

protections to threatened and endangered species in the United States, and is administered by the 

U.S. Fish and Wildlife Service (USFWS) (http://endangered.fws.gov/) and U.S. National Marine 

Fisheries Service (NMFS) (http://www.nmfs.noaa.gov/prot_res/overview/es.html). As defined by 

the Act, endangered refers to species that are "in danger of extinction within the foreseeable 

future throughout all or a significant portion of its range," while threatened refers to "those 

animals and plants likely to become endangered within the foreseeable future throughout all or a 

significant portion of their ranges." 

LE- Listed Endangered: 

LT- Listed Threatened: 

C- Candidate: 

10

Shell Growth 

Dan Dourson, Judy Dourson, Copperhead Environmental Consulting 

Reprinted with permission from Land Snails of the Great Smoky Mountains 

(Eastern Region) 

Mesodon Mesomphix 

Juvenile 

Sub-adult 

Adult 

Immature shells of any species are difficult, if not impossible, to identify. Determining 

the maturity of a shell can often be accomplished by examining the aperture. As shells mature, 

the shape of the aperture changes. Note the aperture of both juvenile species. The bottom seems 

to droop as if it has an invisible weight attached. This begins to round out until it finally reaches 

a more horizontal oval shape at maturity. 

Immature Mesodon species are easily confused with Mesomphix species. Mesodon and 

Triodopsis species do not form reflected lips until they reach maturity. Other species such as 

Mesomphix, Anguispira, and Ventridens do not possess reflected lips even at maturity. 

11

Land Snail Collection Strategies 

Marla L. Coppolino, Southern Illinois University Carbondale 

Despite that land snails don’t seem to share the same spotlight in the mollusk world as 

marine or even freshwater mollusks do, there are many worthwhile reasons to collect and study 

land snails. Land snails are easily accessible, ubiquitous in distribution, and they do not require 

expensive or elaborate equipment to collect. They represent an understudied group (Lydeard et 

al., 2004), and there is much knowledge to be gained from these organisms and their habitat 

associations. Land snails are a vital part of the ecosystem and take up essential nutrients from 

the detritus and soil, which are then in turn passed on to higher trophic levels, inevitably 

affecting them for better or for worse (Barker, 2004). Land snails are studied for their capacity 

as ecological indicators (Shimek, 1930), and even as indicators of the effects of pollution and 

global climate change (Graveland et al., 1994; Regoli et al., 2006). Many of these types of 

studies are in their early stages. Much remains to be learned about land snails. And so, the 

world needs more land snail collectors! 

The literature on land snail collecting suggests a wide array of methodologies. Basic 

questions should be asked before embarking on research, whether formal or informal, that 

involve the collection of land snails: 

1) Is the study exploratory in nature, that is, is the intention of the collector surveying an 

area to “see what’s there”? 

2) Are multiple areas being surveyed? 

3) Does the investigation also include comparisons of land snail populations with habitat 

and microhabitat? 

4) Is the method intended to be repeatable and quantifiable? 

Land snail collecting can be divided into two basic categories: qualitative and 

quantitative. Historically speaking, most of the collecting in North America has been qualitative. 

This means that areas are searched for snails without being under the confines of measured time 

or space. Locality data for the specimen is recorded by the collector, often along with other 

information relating to its habitat. The data from these efforts, amassed by important collectors 

of the past such as Henry Pilsbry, Frank C. Baker, and Leslie Hubricht, have offered 

contemporary collectors a very useful baseline of information on species, and their habitats and 

ranges. 

Since around the mid-20 th century, collecting efforts in North America have turned more 

towards quantifiable studies. A quantitative study implies that land snails are collected using 

some standard measurement, either by time, by volume sampled (leaf litter, soil, etc.) or by area, 

and often by some combination of these factors. Most often, snails collected in this type of study 

are also accompanied by habitat or microhabitat data, which is also measured in a quantifiable, 

repeatable way. When a collecting method is quantitative, the snail and habitat/microhabitat data 

can be used in statistical analyses. Most modern studies use quantitative methods of collection, 

from which population estimates can be obtained (Bishop, 1977). 

The basic tools for land snail collecting are quite simple. In the field, plastic vials with 

attached snap lids are a good choice for collecting snails. Most often, direct visual search is 

employed. In many areas, the greatest amount of diversity comprises “micro snails”, or 

generally speaking, snails that measure less than 5 mm at the largest shell dimension, and cannot 

be readily found in a visual search. To collect these snails most efficiently, leaf litter and usually 

12

the top 2 cm of soil are collected in a bag (either plastic or cloth), to be sorted later in the 

laboratory. The litter can be searched visually under magnification, and then passed through a 

series of sieves to pick out the smaller snails. 

Depending on the species and the region, land snails have a wide range of microhabitat 

preferences, but generally the first places to look for snails would be in moist areas: the leaf 

litter, under bark, coarse woody debris or logs, along bases of trees and rocks, and generally in 

the interface regions of the forest floor, such as the crevices between a log and the ground litter, 

and between exposed tree roots. Don’t forget to observe tree trunks, limbs and crotches between 

branches, for snails could be actively crawling there, particularly on a rainy day. In regards to 

the best time of day to search for living snails, the greatest success often occurs during the 

overnight hours, but this time isn’t necessarily practical. Humid, cool mornings and evenings 

can yield a large collection of living snails. The best seasons to collect live snails in much of 

North America are spring and fall, when they are most active, since they hibernate in the cold of 

winter and aestivate in the heat of summer. 

In both qualitative and quantitative collections, the question arises as to whether to collect 

only empty shells, only live snails, or both. Researchers have advocated different views. In 

some studies, only living specimens were counted (Boag, 1982; Sulikowska-Drozd, 2005). 

Many other studies have included both. In any case, the decision relates to the purpose of the 

study. For maximum recovery of species diversity, a combination of live and dead collected 

snails is best. A couple of exceptions apply to live-collecting: one, when the species found are 

rare or endangered, and two, when species are found that tend to exist in high abundances, but in 

extremely localized habitats. In these cases, it is preferable to limit collecting to one or a few 

individual specimens. The advantage to including empty shells in the collection is that the 

presence of even a dead animal can be useful data in a study. Another great pro to collecting 

empty shells is that, even with a broken fragment remaining, species identification can often be 

made. The disadvantages include the fact that the time the snail has been dead is largely 

unknown. Shells usually persist longer on high-pH soils and decompose quicker on low-pH 

soils. Another uncertainty exists in that often times empty shells fall downward from a higher 

area, such as a bluff, hence the recorded microhabitat locality of an empty shell could be in error. 

A quick but useful note to mention is to remind the collector to have at least some idea of 

the species he or she will encounter. More specifically, some species are carnivorous, such as 

the Haplotrema species of North America. If a live Haplotrema species is placed in the same 

collection vial as another live, non-carnivorous species, you could very well end your field day 

with a well-fed Haplotrema and the empty shell of its victim! 

It is not essential to have the ability to identify all your snails in the field. But an 

important aspect of snail collection (or of any biological collection) is to label your specimens in 

the field with locality data. These data can range from the name of the site (park, nature 

preserve, etc.), to GPS coordinates, quadrangle information, elevation and a full description of 

the habitat features. It is mainly important that the locality of the collection can be found again 

by future collectors. 

Many examples of quantitative sampling methods exist in the literature. As some basic 

examples, an area can be sectioned into plots or transects, or into quadrats that commonly 

measure one square meter and can be placed on the ground. The transect, plot or quadrats can be 

visually searched for a pre-determined length of time, or as often in the case for quadrats, the 

entire contents of it (leaf litter, coarse woody debris, etc.) are collected in a large bag and sorted 

for snails in the laboratory, rather than in the field. Transects or quadrats can be randomly 

13

placed, or samples can be obtained by a stratified random sampling method, meaning that the 

quadrat is placed in an area randomly selected from other areas that area most likely to contain 

snails. The latter school of thought follows the knowledge that snails tend to be very patchy in 

their distribution. As discussed previously, land snails are most likely to be found near bases of 

trees, rocks, under logs and various microhabitats that could otherwise be missed in a completely 

random selection of collection areas. 

Yet another method advocated by some researchers involves the use of cardboard sheets 

or masonite boards (Boag, 1982; Hawkins et al., 1998). The sheets or boards are placed at a site 

and left for a period of time to collect land snails on their undersides. As with all methods, 

various pros and cons exist with this technique. On the plus side, the sheets or boards can help 

collect species that may otherwise be found, such as some species of slugs. On the downside, 

cardboard sheets in particular need to be checked fairly often, as they are prone to warping and 

disintegrating in excessive rainfall. In general, if treating the use of sheets or boards as a 

quantitative method, the bias exists that more snails will be attracted to them than would 

normally be collected in the same two-dimensional area. 

Finally, a few words can be suggested regarding other details about land snail collecting. 

Before venturing to an area with collecting vials in hand, learn whether a permit is required to 

collect land snails there. If the area is a state park, federally-owned land (including U.S. Forest 

Service areas), or nature preserve, you will need to obtain a permit. If you happen to be 

collecting during a hunting season, it is advisable to wear bright colors, or even an orange 

hunting vest. And lastly, adhere to the basic rules of outdoor safety: it is safer to collect with 

another person, to have awareness of the animals inhabit the area (e.g. poisonous snakes), and to 

protect your exposed skin from encounters with poison ivy and biting insects. 

Enjoy land snail collecting. With well-recorded data to accompany your collected 

specimens, your work could be a valuable contribution to the knowledge, and ultimate 

preservation, of land snail species. 

14

Phylogeny and Classification of North American Terrestrial 

Gastropods 

Kevin J. Roe, Iowa State University 

After the Arthropoda, the Mollusca is considered to be the second most diverse of all 

metazoan phyla, and the gastropods represent the most diverse group of mollusks consisting of 

~80% of all mollusks (~80,000 species). It is perhaps no great surprise then that the 

classification of such a large and diverse group of organisms has continued to change over the 

years. In general, these changes have reflected our improved understanding of the evolutionary 

relationships of gastropods. The past several decades have witnessed some dramatic shifts in our 

understanding of the relationships of the major gastropod lineages, and with them the placement 

and relationships of terrestrial snails. An understanding of the changes that have taken place in 

the classification of terrestrial snails will prove to be invaluable when attempting to reconcile the 

names and associated information contained in recent publications with that found in older, but 

still very important ones. 

In order to fully appreciate the value of a taxonomic classification, it is paramount to 

realize that classifications schemes are, first and foremost, information retrieval systems. The 

information they convey is of a set of relationships (all the species in a genus should be more 

closely related to each other than to species in other genera, and likewise for genera within 

families) that is arranged hierarchically. By looking at a classification one should be able to 

understand something about the where the organism of interest fits within this hierarchy. In 

more modern classifications, there has been some movement away from the use of traditional 

rankings (class, order, suborder etc.). This fact should not cause any great concern, because what 

is most important is the relative position of taxa within the hierarchy, not the categorical rank 

associated with it. 

Traditionally, gastropods were classified based on the presence and position of their 

respiratory apparatus into three groups: Prosobranchia (gills in-front of the heart), 

Opisthobranchia (gills behind the heart), and Pulmonata (lack gills, but use a highly vascularized 

mantle for respiration). While convenient, this classification has been recognized to not reflect 

the evolutionary relationships of gastropods (a misleading information retrieval system) and has 

been abandoned in favor of classifications based on the objective analysis of both morphological 

(e.g. Salvini-Plawen and Steiner, 1996; Ponder and Lindberg, 1997; Dayrat and Tillier, 2002) 

and DNA (e.g. Tillier and Ponder, 1992; Dayrat et al. 2001) data. Current classifications place 

pulmonates and opisthobranchs together in the clade Euthyneura. This group together with 

pyramidelids and related snails form the Heterobranchia. Those interested in exploring higherlevel 

gastropod systematics should consult the publications listed in this summary in particular, 

Bieler (1992) and Ponder and Lindberg (1997). The use of ever more sophisticated methods of 

analysis and high-tech tools has resulted in a refinement of our knowledge of the relationships of 

gastropods, but, despite these improvements our understanding is far from complete and still 

very much in a state of flux. The only way to improve the current state of knowledge of these 

relationships is to continue to collect and analyze data, and just as importantly, to continue to 

train researchers to be able to locate and identify gastropods. 

15

Diversity and higher-level systematics of land snails 

Solem (1978) estimated that there are some 20,500 land pulmonates world-wide, nearly 

twice the earlier estimate provided by Boss (1971). The focus of this workshop is on the 

terrestrial snails of the continental United States. Estimates of the number of the number of land 

snail species are available for several states and regions but accurate estimates for North America 

or even the U.S are hard to come by. A back of the envelope enumeration of the species listed in 

Turgeon et al. (1998) results in slightly less than 1100 native species in the U. S. and Canada, 

with about 70 more non-native species. The increasing problem of invasive, non-native species 

emphasizes the importance of recognizing all taxa at least at a basic level, and some effort should 

be made to become familiar with these groups. The terrestrial snails were traditionally placed in 

two large groups, the Prosobranchia, which has been discarded, and the Pulmonata, which has 

been retained. The majority of the native land snails in the United States are placed within the 

latter group. In most recent classifications, the Pulmonata is further divided into three major 

lineages, the Basomatophora, Systellommatophora, and Stylommatophora based on the number 

and kind of cephalic tentacles and the location of eyes on the head. In the Basomatophora, one 

pair of contractile tentacles (they shrink, accordion-like) is present on the head and the eyes are 

located at their base. In the Systellommatophora, the head has two pairs of contractile tentacles, 

with eyes on tips of the upper pair. Lastly, the Stylommatophora also have two pairs of 

tentacles, with eyes on tips of the upper pair, but in this case the tentacles are retractile (they can 

be inverted, like the fingers of a glove). 

The Stylommatophora was traditionally divided into four subgroups, the Orthurethra, 

Mesurethra, Heterurethra, and Sigmurethra, based on the position and structure of the kidney and 

ureter (Pilsbry, 1900; Baker, 1955). Of these groups only the Orthurethra has remained 

consistently supported (e.g. Wade et al, 2001; 2006) and based on the most comprehensive 

modern analyses the stylommatophorans are divided into two groups, a smaller achatinoid clade 

and a much larger non-achatinoid clade. Traditionally, the majority of land snail species were 

placed in the Sigmurethra, and this group was further divided into two groups, the Holopoda and 

the Aulacopoda based on the morphology and placement of pedal grooves (Pilsbry, 1896; 1900). 

In recent phylogenetic reconstructions, the achatinoid clade includes only holopod species, 

whereas the non-achatinoid clade includes holopods and aulacopods. 

To a casual observer the revisions to the classifications of terrestrial snails based on 

recent phylogenetic analyses have apparently only resulted in the reshuffling of some taxa and 

the loss of several named groups to wit: most land snails are pulmonates, most pulmonates are 

stylommatophorans, and the majority of stylommatophorans are grouped together in a large nonachatinoid 

clade. What is not readily apparent is that these new hypotheses inform us about the 

way organisms evolve and how some characters that at first appear to be similar due to common 

ancestry are actually derived independently. To fully appreciate the full potential that land snails 

have to teach us about evolutionary processes requires a more detailed understanding of the 

different families of land snails. 

16

Agriolimacidae 

Megan E. Paustian, University of Maryland 

Only 4 species (Deroceras) of this family are native to the U.S. and Canada. 

Native Agriolimacids are 15-50 mm long slugs. The Agriolimacid mantle has concentric 

ridges, a pneumostome in the posterior half, and an internal shell. The tail tip is rounded with a 

short keel extending up the dorsum. Species tend to vary in the complexity or presence of the 

penial flagellum, which is often a diagnostic character. However, most individuals of Deroceras 

laeve are aphallic. 

Native Agriolimacids live in mixed-wood forests, although the habitat and food 

requirements of most species are poorly known. They may take shelter under leaf litter, 

vegetation, or coarse woody debris. D. laeve is a pervasive species that lives in marshes, moist 

fields, along riverbanks, and in gardens. This species, which can be a garden pest, consumes live 

and dead plants. 

Two species are found in the west coastal region between Alaska and California, while D. 

heterura is only found in New Mexico. D. laeve's native range is extremely broad, spanning the 

entire U.S. and the coastal provinces of Canada, as well as much of the Holarctic region 

worldwide. 

Information summarized from Pilsbry (1948), Chichester & Getz (1973), Kerney & 

Cameron (1994), Burke (1999). 

TAXON AUTHOR 

G- 

RANK DISTRIBUTION 

Deroceras hesperium Pilsbry, 1944 G2 OR, WA; Canada: BC 

Deroceras heterura Pilsbry, 1944 G1G2 NM 

AK, AL, AR, CA, DE, FL, GA, HI, IA, ID, IL, IN, KS, KY, LA, MD, 

ME, MI, MO, MS, NC, ND, NE, NJ, NM, NY, OH, OK, PA, SD, TN, 

Deroceras laeve (Muller, 1774) G5 TX, UT, VA, VT, WA, WI, WV; Canada: AB, BC, NF, NS, ON, QC 

Deroceras monentolophus Pilsbry, 1944 G4 AK, CA, OR, WA 

17

Annulariidae 

Kathryn E. Perez, Duke University 

The single U.S. species of Annulariidae is a calciphile. Hubricht (1985) states it is found 

crawling on rocks and tree trunks in wet weather. Elongate, 10-12 mm long, operculate, related 

to Cuban Chondropoma species. 

TAXON AUTHOR 

Chondropoma dentatum (Say, 1825) G2G3 FL 

G- 


Arionidae (former family) 


There are 28 species (10 genera) that are native to the western U.S and Canada. 

The family Arionidae was recently reclassified, such that the Eurasian subfamily 

Arioninae now takes the name Arionidae, and the remaining three subfamilies are raised to 

family status (below). All species are slugs. Most have a pneumostome in the anterior half of 

the mantle, a wide foot fringe, a ribbed jaw, and an epiphallus. 

The Anadenidae are 30-100 mm long. Prophysaon (taildropper slugs) are characterised 

by a tail constriction, visible as a dark line on the sole, where a portion of the tail can break off 

(autotomize) to distract predators. 

The Ariolimacidae includes some diverse slug species. Ariolimax (bananaslugs) are 

famously large slugs that are 100-260 mm long and that have a variably colorful (yellow, olive, 

or brown) and often spotted mantle. A prominent keel runs centrally along the dorsum, and the 

pneumostome is located in the posterior half of mantle (RGF, 2004). Hesperarion 

18

(westernslugs) are smaller slugs that are about 30-50 mm long. Hesperarion and Ariolimax have 

a caudal mucous pit. Magnipelta has a distinctive broad mantle that is spread over most of its 

dorsal body, and it is about 65 mm long. All of these genera have an undivided foot sole. 

The Binneyidae are 8-60 mm long slugs. The family is distinguished by a prominent, 

humped mantle and an external shell that is either tiny and plate-like or large and whorled. 

Hemphillia (jumping-slugs) are thin-bodied slugs whose solid, flexible tails enable them to flip 

away from predators. 

The Anadenids, Ariolimacids, and Binneyids tend to inhabit moist coniferous and 

deciduous forests. Some species have received special conservation status, particularly those 

that are restricted to small endemic ranges or that preferentially inhabit old-growth forests 

undergoing logging (e.g. some Hemphillia). Slugs may take shelter under bark, coarse woody 

debris, or leaf litter. Typical foods are fungus, lichens, detritus, and live plants. 

The Anadenids, Ariolimacids, and Binneyids inhabit the western U.S. and Canada westward 

from the Rocky Mountains. They span north to Alaska, south to California, and east to Montana 

and Alberta. 

Information summarized from Pilsbry (1948), Burch & Pearce (1989), COSEWIC 

(2003), Forsyth (2004), Pearce et al. (2004), Bouchet & Rocroi (2005). 

TAXON AUTHOR 

G- 


Anadenulus cockerelli (Hemphill, 1890) G1G2 CA 

Ariolimax californicus J.G. Cooper, 1872 G2 CA 

Ariolimax columbianus (Gould, 1851) G5 CA, ID, OR, WA; Canada: BC 

Ariolimax dolichophallus Mead, 1943 G2 CA 

Binneya notabilis J.G. Cooper, 1863 G1 CA 

Gliabates oregonius Webb, 1959 G1 OR 

Hemphillia burringtoni Pilsbry, 1948 G1G2 WA 

Hemphillia camelus Pilsbry and Vanatta, 1897 G4 ID, MT, WA; Canada: AB, BC 

Hemphillia danielsi Vanatta, 1914 G2G3 MT 

Hemphillia dromedarius Branson, 1972 G3G4 WA; Canada: BC 

Hemphillia glandulosa 

Bland and W.G. Binney, 

1872 G3G4 OR, WA; Canada: BC 

Hemphillia malonei Pilsbry, 1917 G3 OR, WA; Canada: BC 

Hemphillia pantherina Branson, 1975 G1 WA 

Hesperarion hemphilli (W.G. Binney, 1875) G2 CA 

Hesperarion mariae Branson, 1991 G2 OR 

Hesperarion niger (J.G. Cooper, 1872) G2 CA 

Hesperarion plumbeus Roth, 2004 G1G3 CA 

Kootenaia burkei 

Leonard, Chichester, Baugh, 

and Wilke, 2003 G2 ID, MT 

Magnipelta mycophaga Pilsbry, 1953 G3 ID, MT, WA; Canada: BC 

Prophysaon andersoni (J.G. Cooper, 1872) G5 AK, CA, ID, MT, OR, WA; Canada: BC 

Prophysaon boreale Pilsbry, 1948 G1G3 AK 

Prophysaon coeruleum Cockerell, 1890 G3G4 CA, ID, OR, WA; Canada: BC 

Prophysaon dubium Cockerell, 1890 G4 CA, ID, OR, WA 

Prophysaon fasciatum Cockerell, 1890 G1G3 CA 

Prophysaon foliolatum (Gould, 1851) G4G5 OR, WA; Canada: 

Prophysaon humile Cockerell, 1890 G3 ID, MT 

Prophysaon obscurum Cockerell, 1890 G1G3 WA 

Prophysaon vanattae Pilsbry, 1948 G4 CA, OR, WA; Canada: BC 

Udosarx lyrata Webb, 1959 G2 ID, MT 

Zacoleus idahoensis Pilsbry, 1903 G3G4 ID, MT, WA 

19

Bradybaenidae 

Mark A. Ports, Great Basin College 

Taxonomic note: In this workbook we are including the family Monadeniidae and genus 

Monadenia in Bradybaenidae following Roth & Sadeghian (2003) and Roth (1997). 

This family is primarily distributed from south-central California, on the west side of the 

Sierra Nevada and into the Cascade Mountains of southwestern Oregon. Monadenia fidelis, a 

common species, is found in wet, coastal forests of California, Oregon, Washington, British 

Columbia, and Alaska. There is only one genus, Monadenia in this entire region with 37 records 

of species and subspecies. 

The species and subspecies of this genus range in size from 24 mm to 1 cm. They are 

typically blackish brown, with a sub-peripheral, black band. Species of Helminthoglypta have a 

supra-peripheral band. Monadenia shells may be depressed or conical in shape. The periostracum 

may be smooth, have globular bumps, or bristles. The shell is usually very thin and fragile. The 

aperature is oval and the number of whorls is 5 to 6. The most effective means to separate these 

species is by shell size, coloration, and dissection of genitalia. 

Most of the Monadenia have a life span of 10 - 15 years, are slow growing, and may not 

reach maturity for 8 -10 years. In dry habitats they may aestivate for up to 8 years. All of these 

species are hermaphroditic, transfer sperm with a dart apparatus, and lay eggs in loose soil, litter, 

or beneath downed, rotten trees. Monadenia are found in old growth forests along the coast, dry 

coniferous and deciduous forests of the Sierra Nevada and Cascade mountains, along riparian 

zones with deciduous trees especially maple, and in wet, mountain meadows. These species 

depend on talus slopes with a complete canopy of forest which keeps their habitats cool and 

moist. Aestivation and hibernation in talus slopes require a litter composed of deciduous leaves 

and detritus. During the wet months, colonies of Monadenia will leave the talus slopes and 

forage into the surrounding habitats to feed on green vegetation, fungi, feces, and dried fruit. 

Ten of the species and eleven subspecies of Monadenia are considered critically 

imperiled or sensitive according to survey work of the Northwest Forest Plan and the Sierra 

Nevada - Cascade management plan. The species of this genus are impacted by lumber removal 

and cattle grazing which can lead to a dryer and hotter microhabitat. Colony fragmentation, as in 

all the western land snails, can lead to extinction as individuals die off with little immigration 

from colonies along riparian zones. Draining of wet meadows for agriculture and urban use, 

disturbance of talus slopes, and ground fires will also impact these colonies. The status of all 

western land snails in different habitats are important indicators of the general ecosystem health. 

20

Information summarized from: NatureServe, Roth & Sadeghian (2006), Pilsbry (1939), 

Northwest Forest Plan (2002), Sierra Nevada-Cascade Mountain Management Plan (1997), Roth 

(2002). 

TAXON AUTHOR 

G- 


Monadenia callipeplus S.S. Berry, 1940 G1G2 CA 

Monadenia chaceana S.S. Berry, 1940 G2 CA, OR 

Monadenia churchi 

Hanna and A.G. Smith, 

1933 G2 CA 

Monadenia circumcarinata (Stearns, 1879) G1 CA 

Monadenia cristulata S.S. Berry, 1940 G1G2 CA 

Monadenia fidelis (J.E. Gray, 1834) G4G5 AK, CA, OR, WA; Canada: BC 

Monadenia infumata (Gould, 1855) G2G3 CA 

Monadenia marmarotis S.S Berry, 1940 G1 CA 

Monadenia mormonum (Pfeiffer, 1857) G2 CA 

Monadenia scottiana S.S. Berry, 1940 G1G2 CA 

Monadenia setosa Talmadge, 1952 G2 CA 

Monadenia troglodytes Hanna and A.G.Smith, 1933 G1G2 CA 

Monadenia tuolumneana S.S. Berry, 1955 G1 CA 

Monadenia yosemitensis (Lowe, 1916) G1 CA 

Bulimulidae 


The Bulimulidae are a family of hundreds of species, mostly found in South America. 

Shells are bulimoid in form and typically white or brown background color with grey or brown 

streaks or stripes. The largest species reach ~40 mm in length. 

The genus Drymaeus is restricted in the U.S. to Florida where these snails live on 

trees, bushes, or herbaceous vegetation. The species in this genus are semi-arboreal and are 

typically found estivating or crawling on vegetation (Hubricht, 1985). 

Rabdotus is one of the characteristic snails of the arid and semi-arid regions of Texas and 

adjacent Mexico (Fullington & Pratt, 1973). The species of Rabdotus occupy a range of habitats 

from coastal plain scrubland to Chihuahuan desert. These snails are colonial and can reach high 

densities locally and are often found crawling or estivating on tall grass, shrubs, mesquite brush, 

on human build structures, or under rocks and other cover. They are typically active during 

times of high humidity, at night, or after rain. In parts of their range after rainfall they have been 

known to be active in high enough concentrations that slicks caused by smashed snails have 

21

caused cars to run off roads. Rabdotus are of interest to a wide range of fields as they have 

signal specific habitat or environmental types and shells are found in large numbers at 

archaelogical sites presumably they formed a food source for various cultures. There is a great 

deal of taxonomic confusion surrounding the species of Rabdotus in the U.S. and a thorough 

modern treatment is necessary. 

TAXON AUTHOR 

G- 


Drymaeus dominicus (Reeve, 1850) G2G3 FL 

Drymaeus dormani (W.G. Binney, 1857) G2G3 FL 

Drymaeus multilineatus (Say, 1825) G5 FL 

Naesiotus christenseni 

(W.B. Miller and Reeder, 

1984) G1 AZ 

Naesiotus nigromontanus (Dall, 1897) G5 

Rabdotus alternatus (Say, 1830) G5 TX 

Rabdotus dealbatus (Say, 1830) G5 AL, AR, IL, KS, KY, LA, MO, MS, NM, OK, TN, TX, WI 

Rabdotus durangoanus (von Martens, 1893) G3G5 NM, TX 

Rabdotus mooreanus (Pfeiffer, 1868) G5 AL, MS, OK, TX 

Rabdotus pilsbryi (Ferriss, 1925) G5 TX 

Carychiidae 

Jochen Gerber, Field Museum of Natural History 

Shells of the only carychiid genus found in North America, Carychium, are minute, 

elongate, transparent or white, 1.2-2.5 mm high. They have expanded and often thickened 

peristomes. A characteristic of the group is a columellar fold whose shape is species-specific. It 

can be seen through the shell wall in transparent shells or can be studied by carefully breaking a 

“window” into the body whorl. Snails in the genus Carychium are found in very moist to wet 

habitats, such as floodplains, swamps, and moist woods under rotting logs or in leaf litter. 

Carychium stygium is only found in caves. 

TAXON AUTHOR 

G- 


Carychium clappi Hubricht, 1959 G5 AL, GA, IN, KY, MD, NC, PA, SC, TN, VA, WV 

CO, DE, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, NC, 

NE, NJ, NM, NY, OH, OK, PA, SC, SD, TN, TX, VA, VT, WI, WV; 

Carychium exiguum (Say, 1822) G5 Canada: AB, NB, NS, ON, QC, NF 

22

Carychium exile I. Lea, 1842 G5 

AL, AR, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, 

MS, NC, ND, NJ, NY, OH, OK, PA, SC, SD, TN, VA, VT, WI, WV; 

Canada: AB, NS, ON, QC 

Carychium mexicanum Pilsbry, 1891 G5 AL, FL, GA, LA, MS, OK, SC, TN, TX 

Carychium nannodes G.H. Clapp, 1905 G5 

AL, GA, IN, KY, MD, MI, MS, NC, NY, OH, PA, TN, VA, WV; 

Canada: ON 

Carychium occidentale Pilsbry, 1891 G3G4 CA, ID, OR, WA; Canada: BC 

Carychium riparium Hubricht, 1978 G2G3 IN, KY; Canada: ON 

Carychium stygium Call, 1897 G3G4 KY, TN 

Cepolidae 


This family occurs in the Greater Antilles and in the Bahamas. Only one species, 

Hemitrochus varians, also lives in North America (South Florida). Its medium-sized (ca 15 mm 

in diameter) shell is globose-conic, perforate, the aperture not expanded. The relatively smooth 

shell is white with varying arrangements of dark spiral bands. Aperture pinkish within. 

Hemitrochus varians lives on shrubs and trees. 

TAXON AUTHOR 

G- 


Hemitrochus varians (Menke, 1829) G2G4 FL 

23

Ceriidae 


Cerion is a large genus (~600 spp) endemic to the Caribbean with one U.S. 

representative. These snails typically found attached to grass stems, trees, and shrumbs near the 

beach, but above the tide and spray lines. They are large (up to 28 mm length), elongate, many 

whorled, with opaque-white ribbed shells, sometimes streaked with gray or brown. Cerion tend 

to be inactive by day, estivating attached to vegetation and active at night and in rainy weather. 

Several Caribbean Cerion species have been introduced into Florida. 

TAXON AUTHOR 

G- 


Cerion incanum (A. Binney, 1851) G1 FL, LA 

Charopidae 


Charopids are helicoid snails with minute, ribbed shells ranging from 1-2.6 mm in height 

and 2-6 mm width (Pilsbry, 1946). In the U.S. they are limited in distribution to Western states at 

high elevation. Other members of this family extend throughout South America and they are 

particularly diverse in South Africa, New Zealand and Australia (Bequaert & Miller, 1973). 

24

Radiodiscus millecostatus range extends north into NM and AZ, but is mostly found in Mexico 

and south to Costa Rica (Bequaert & Miller, 1973). Metcalf & Smartt (1997) report finding this 

species in leaf litter in Aspen forest in mountains in New Mexico at 6,800 -10,800 ft. elevation. 

R. abietum is about twice as large as R. millecostatus (Pilsbry, 1946) and inhabitats coniferous 

forests dominated by fir, tends to be found near streams (Pilsbry, 1946). 

TAXON AUTHOR 

G- 


Radiodiscus abietum H.B. Baker, 1930 G4 ID, MT, OR, WA 

Radiodiscus millecostatus Pilsbry and Ferriss, 1906 G3 AZ, NM 

Cionellidae 


Roth & Sadeghian (2003) state Cochlicopa lubrica (formerly Cionella lubrica) is 

Holarctic in distribution, in North America ranging from northern Alaska to Chihuahua and 

Nuevo Leon Mexico, habitats include near sea level in boreal forest and cold-temperate areas to 

higher elevations in the western and mountain states. Forsyth (2004) describes C. lubrica as 4.4- 

6.5 mm high, glossy, spindle shaped, with a thickened apertural lip. Forsyth (2004) describes the 

natural history of C. lubrica as occurring mostly in disturbed habitats such as roadsides and 

gardens where they eat living and dead plant material (summarized from Forsyth 2004). Pilsbry 

(1946) stated this species lives among damp leaves in densly shaded places, under wood, or in 

chinks of stone walls. 

TAXON AUTHOR 

G- 


AK, CA, CO, IA, ID, IL, IN, KS, MA, ME, MI, MN, MO, NC, NE, 

NH, NJ, NM, NY, OH, PA, RI, SD, TN, UT, VA, VT, WA, WI, WV; 

Cochlicopa lubrica (Muller, 1774) G5 Canada: AB, BC, NB, NS, ON, QC, NF 

CT, DE, IA, IL, IN, KS, MA, ME, MI, MN, MO, NJ, NY, OH, PA, RI, 

Cochlicopa lubricella (Porro, 1838) G5 SD, TX, VA, VT, WI, WV; Canada: AB, NS, ON 

AL, AR, CA, GA, IA, IN, KY, MA, MD, ME, MI, MN, MO, MS, NC, 

Cochlicopa morseana (Doherty, 1878) G5 NY, OH, PA, SC, TN, VA, VT, WI, WV; Canada: NS, ON 

Cochlicopa nitens (Gallenstein, 1848) G4 IL, MA, ME, MI, MN, NY, OH, PA, VT, WI; Canada: ON 

25

Discidae Thiele, 1931 (Patulidae Tryon, 1866) 

Aydin Örstan, Carnegie Museum of Natural History 

Taxonomic Note: Species in this family are discussed under the name Discidae Thiele, 

1931 following Bouchet & Rocroi (2005: 66) who consider Patulidae Tryon, 1866 a synonym of 

Discidae. 

The genus Anguispira is endemic to North America. Anguispira are woodland snails. At 

least two species, A. alternata and A. fergusoni, characteristically climb trees in warm and wet 

weather, especially at night, to feed on fungi and rotting wood. Both A. alternata and A. 

fergusoni (and perhaps other species also) become dormant in the winter and laboratory 

populations of A. alternata are known to require exposure to low temperatures prior to 

reproduction. Increased winter temperatures due to global warming may, therefore, threaten 

especially the more southern populations of A. alternata. The known Anguispira species are 

conchologically variable, especially in spire height and peripheral angulation of the body whorl. 

In some cases, these variations may reflect cryptic species lumped under currently accepted taxa. 

The genus is in need of a taxonomic revision. 

Discus is a Holarctic genus. Most North American Discus species live in forests; D. 

whitneyi inhabits wet meadows and marshy places. The European D. rotundatus has been 

recorded in parks and disturbed areas in northeast and northwest U.S. and Canada. 

TAXON AUTHOR 

G- 

RANK LISTED DISTRIBUTION 

Anguispira alabama (G.H. Clapp, 1920) G2 AL, TN 

Anguispira alternata (Say, 1816) G5 

AL, AR, CT, DE, GA, IA, IL, IN, KS, KY, LA, MA, MD, 

ME, MI, MN, MO, MS, NC, NE, NH, NJ, NY, OH, OK, PA, 

RI, SD, TN, VA, VT, WI, WV; Canada: NB, NS, ON, QC 

Anguispira cumberlandiana (I. Lea, 1840) G3 AL, TN 

Anguispira fergusoni (Bland, 1861) G4 DE, MD, NC, NJ, NY, PA, SC, VA 

Anguispira jessica Kutchka, 1938 G3G4 AL, NC, TN, VA 

Anguispira knoxensis (Pilsbry, 1899) G1G2 TN 

Anguispira kochi (Pfeiffer, 1821) G5 

ID, IL, IN, KY, MI, MO, MT, OH, OR, PA, TN, WA, WI, 

WV; Canada: BC, ON 

Anguispira macneilli Walker, 1928 G2 TX 

Anguispira mordax (Shuttleworth, 1852) G4 AL, KY, NC, TN, VA, WV 

Anguispira nimapuna H.B. Baker, 1932 G1 ID 

Anguispira picta (G.H. Clapp, 1920) G1 LT TN 

Anguispira rugoderma Hubricht, 1938 G2 KY 

AL, AR, FL, GA, IL, KY, LA, MO, MS, NC, OK, SC, TN, 

Anguispira strongyloides (Pfeiffer, 1854) G5 

TX, VA 

Discus brunsoni S.S. Berry, 1955 G1 MT 

26

Discus bryanti (Harper, 1881) G3 AL, KY, NC, TN, VA 

CT, DE, IA, IN, KY, MA, MD, ME, MI, MN, MS, NH, NJ, 

NY, PA, RI, SD, TN, VA, VT, WI, WV; Canada: MB, NS, 

Discus catskillensis (Pilsbry, 1896) G5 

ON 

Discus clappi (Pilsbry, 1924) G1 AL, TN 

Discus macclintocki (F.C Baker, 1928) G1 LE IA, IL, MO 

Discus marmorensis H.B. Baker, 1932 G1G2 IA, ID 

Discus nigrimontanus (Pilsbry, 1924) G4 AL, AR, KY, MO, NC, TN, VA 

AL, FL, GA, IA, IL, IN, KY, LA, MD, MI, MO, MS, NC, 

Discus patulus (Deshayes, 1830) G5 

NY, OH, PA, SC, TN, VA, WI, WV; Canada: ON 

Discus selenitoides (Pilsbry, 1890) G1 CA 

AZ, CO, IA, IL, KS, MO, MT, NE, NM, OR, SD, UT, WY; 

Discus shimekii (Pilsbry, 1890) G5 

Canada: AB, BC, YT 

AK, AZ, AR, CA, CT, DE, IA, ID, IL, IN, KS, KY, MA, 

MD, ME, MI, MN, MO, MT, NC, ND, NE, NJ, NM, NY, 

OH, OK, PA, SD, TX, UT, VA, VT, WA, WI, WV; Canada: 

Discus whitneyi (Newcomb, 1864) G5 

AB, BC, NB, NS, QC, LB, NF 

Speleodiscoides spirellum A.G. Smith, 1957 G1 CA 

Haplotrematidae 


These snails have medium-sized to large, depresseed to almost planispiral, openly 

umbilicated shells, mostly light-colored shells. The peristome is not or very narrowly expanded 

and usually blunt, but not particularly thickened, and it lacks folds or teeth. 

The Family is restricted to the Americas and the West Indies. In North America, only 

two species live east of the Rocky Mountains, one, Haplotrema concavum, with a very wide 

distribution. The bulk of the North American species is found in the states and provinces 

bordering the Pacific. 

Haplotrematids live usually in forest habitats in leaf litter and under logs and stones. 

They are known to be omnivorous: they eat other snails, earth worms, etc., as well as plant 

material. 

TAXON AUTHOR 

G- 


Ancotrema hybridum (Ancey, 1888) G5 CA, ID, OR, WA; Canada: BC 

Ancotrema sportella (Gould, 1846) G4 AK, CA, ID, OR, WA; Canada: BC 

Ancotrema voyanum (Newcomb, 1865) G1G2 CA, OR 

Ancotrema zopherum Roth, 1990 G1 CA 

Haplotrema alameda Pilsbry, 1930 G1G2 CA 

27

Haplotrema caelatum (Mazyck, 1886) G1 CA 

Haplotrema catalinense (Hemphill, 1890) G1 CA 

AL, AR, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, 

MO, MS, NC, NH, NJ, NY, OH, OK, PA, SC, TN, TX, VA, VT, WI, 

Haplotrema concavum (Say, 1821) G5 WV; Canada: ON, QC 

Haplotrema costatum A.G. Smith, 1957 G1 CA 

Haplotrema duranti (Newcomb, 1864) G2G3 CA 

Haplotrema guadalupense Pilsbry, 1927 G2G4 CA 

Haplotrema keepi (Hemphill, 1890) G1 CA 

Haplotrema kendeighi Webb, 1951 G2 NC, TN 

Haplotrema minimum (Ancey, 1888) G1G2 CA 

Haplotrema mokelumnense Roth, 1990 G1 CA 

Haplotrema transfuga (Hemphill, 1892) G1G2 CA 

Haplotrema vancouverense (I. Lea, 1839) G5 AK, CA, ID, MT, OR, WA; Canada: BC 

Helicarionidae 


These three genera were previously part of the Zonitiidae. Guppya are low spired, small 

(3.5 mm wide), helicoid snails and Euconulus are higher spired bee-hive shaped snails. Hubricht 

(1985) states that most Euconulus species are found in moist leaf litter on wooded hillsies and in 

ravines. Forsyth (2004) describes E. fulvus as being common and widespread, preferring logs 

and debris, under grass and leaf litter at dry and moist sites at all elevations. E. dentatus is also 

found in leaf litter but in dryer situations than the other species. Dryachloa dauca is found in 

lawns and on roadsides (Hubricht, 1985) and Guppya species are very small from

Euconulus polygyratus (Pilsbry, 1899) G5 

DE, IL, IN, KS, MD, MA, ME, MI, MO, NY, PA, WI, WV; Canada: 

AB, MB, ON, SK 

Euconulus praticola (Reinhardt, 1883) G5 Canada: AB, BC 

Euconulus trochulus (Reinhardt, 1883) G5 AL, AR, FL, GA, IL, IN, KY, LA, MO, MS, NC, OK, SC, TN, TX, VA 

Guppya gundlachi (Pfeiffer, 1839) G3 FL, TX 

Guppya miamiensis Pilsbry, 1903 G3Q FL, TX 

AL, AR, FL, GA, IA, IL, IN, KY, LA, MD, MI, MO, MS, NC, NJ, NY, 

Guppya sterkii (Dall, 1888) G5Q OH, OK, PA, SC, TN, VA, WI, WV; Canada: ON 

Helicinidae 


This family of small operculate snails is largely tropical with only a few species in the 

U.S. Helicinids tend to occur in large colonies in moist shaded areas, often in disturbed areas. 

These species tend to climb and occur on sides of buildings, on grass, shrubs, and in trees. 

Fullington & Pratt (1974) state that Oligyra orbiculata estivates arboreally in the summer and 

hibernate in the winter in soil at the base of shrubs and rocks. Hendersonia occulata is relatively 

uncommon and lives on well shaded, humid, and leafy slopes in limestone terrain (Pilsbry, 

1946). From the more widely distributed fossil distribution it appears this species was much 

more widespread in the past and now is restricted to a more limited distribution. 

TAXON AUTHOR 

G- 


Helicina clappi Pilsbry, 1909 G2G3 FL 

Hendersonia occulta (Say, 1831) G4 

IA, IL, IN, KS, KY, MD, MI, MN, MO, MS, NC, NE, OH, PA, TN, 

VA, WI, WV; Canada: ON 

Lucidella tantilla Pilsbry, 1902 G4 FL, MD 

Oligyra orbiculata Say, 1818 G5 AR, FL, GA, KY, LA, MO, MS, OK, TX, AL, TN 

29

Helicodiscidae 

Barry Roth, San Francisco, CA 

The Helicodiscidae are distributed from North America to northern South America and in 

Western Europe and the Indo-Pacific region. The shell is small to minute, with few, flatly coiled, 

slowly expanding whorls, and broadly open umbilicus; sculpture, when present, consists of spiral 

ridges or striations. One or more sets of paired lamellae are present in body whorl of some 

species. The kidney is elongate, triangular, and reaches the hindgut. The central tooth of the 

radula is three-cusped and reduced in size, lateral teeth are three to five three-cusped, and 

marginals are short and broad. An epiphallus is present. The ovotestis is elongate, unbranched, 

and served by a long, straight hermaphroditic duct. 

Habitats of helicodiscids include rock piles, leaf litter, and the undersides of rocks on 

wooded hillsides. Many species are calciphiles, found in limestone rubble and/or caves. Some 

have a tendency to burrow in soil. A few are characteristic of open grassy places, such as 

roadsides, meadows, old fields, along railroads, and (Helicodiscus parallelus) in vacant city lots. 

In its native range in the eastern USA, Lucilla singleyana (formerly Helicodiscus singleyanus) is 

also found in greenhouses; in California, where it is introduced, it is found on roots and bulbs in 

gardens. 

Information summarized from Pilsbry (1948), Hubricht (1985), Solem (1985), Falkner et 

al. (2002). 

TAXON AUTHOR 

G- 


Helicodiscus aldrichianus (G.H. Clapp, 1907) G3 AL, TN 

Helicodiscus barri Hubricht, 1962 G3G4 AL, GA, TN 

Helicodiscus bonamicus Hubricht, 1978 G1 NC, TN 

Helicodiscus diadema Grimm, 1967 G1 VA 

Helicodiscus eigenmanni Pilsbry, 1900 G5Q AR, AZ, KS, NM, TX, UT 

Helicodiscus enneodon Hubricht, 1967 G3G4Q TN, VA 

Helicodiscus fimbriatus Wetherby, 1881 G4 AL, GA, KY, NC, SC, TN, WV 

Helicodiscus hadenoecus Hubricht, 1962 G3 AL, KY, TN, VA 

Helicodiscus hexodon Hubricht, 1966 G1 TN 

Helicodiscus inermis H.B. Baker, 1929 G4 

AL, FL, GA, IL, IN, LA, MD, MO, MS, NC, NE, NJ, OH, 

OK, TN, TX, VA, WI, WV 

Helicodiscus lirellus Hubricht, 1975 G1 VA 

Helicodiscus multidens Hubricht, 1962 G3 TN, VA 

AL, AR, FL, GA, IL, IN, KS, KY, LA, MD, MO, MS, NC, 

Helicodiscus notius Hubricht, 1962 G5Q 

OK, PA, SC, TN, TX, VA, WV 

30

Helicodiscus nummus (Vanatta, 1899) G1G2 AR, KY, OK, TX 

AL, AR, CT, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, 

MD, ME, MI, MN, MO, MS, NC, NE, NH, NJ, NY, OH, 

OK, PA, RI, SC, TN, TX, VA, VT, WI, WV; Canada: NB, 

Helicodiscus parallelus (Say, 1817) G5 

NS, ON, QC, NF 

Helicodiscus punctatellus Morrison, 1942 G1 KY, TN 

Helicodiscus salmonaceus Hemphill, 1890 G2 ID, WA 

Helicodiscus saludensis (Morrison, 1937) G1 NC, SC 

CT, IA, IL, IN, KY, MA, MD, ME, MI, MN, NH, NY, PA, 

Helicodiscus shimeki Hubricht, 1962 G4G5 

TX, VA, VT, WI, WV; Canada: ON 

AL, AR, CA, DE, IL, IN, KS, KY, LA, MD, MI, MO, NE, 

NJ, NM, NY, OH, OK, PA, TN, TX, VA, WI, WY; Canada: 

Helicodiscus singleyanus (Pilsbry, 1889) G5 

ON 

Helicodiscus tridens (Morrison, 1935) G2 OK, TX 

Helicodiscus triodus Hubricht, 1958 G2 NC, VA, WV 

Polygyriscus virginianus (P.R. Burch, 1947) G1 LE VA 

Helminthoglyptidae 


The family Helminthoglyptidae is a diverse and widespread group of land snails from 

California, Oregon, Arizona, New Mexico, and west Texas. This family is composed of eleven 

genera with approximately 83 species of Helminthoglypta, 13 species of Sonorelix, 9 species of 

Micrarionta, 1 species of Chamaeariontales, 1 species of Herpeterous, 8 species of Xerarionta, 

2 species of Cahuillus, 7 species of Eremarionta, 1 species of Noyo, 4 species of Rothelix, and 

72 species of Sonorella. 

The Helminthoglyptidae vary in diameter from 5 mm to 4 cm, and range from a dark 

brown color to a pearly white color. The shells are generally depressed-helicoid in shape, thin in 

texture, with a thin, reflexed peristome, normally without aperatural teeth. Most have a single 

dark supra-peripheral band while some of the Mohave Desert species may not have any band. In 

all of these genera (especially the Sonorella) the best way to identify them is the dissection of 

their genitalia or the study of their phylogeny. 

This family occupies habitat types that include wet temperate forest in Oregon and 

northern California, dry coniferous and deciduous forests of the west side of the Sierra Nevada, 

coastal mountains and canyons of conifers and oak, chaparral brush of coastal California, desert 

valleys and mountains of California, Arizona, Texas and New Mexico. Most species of this 

family are associated with rocky or talus slopes and ledges of dolomite, granite, or shale. 

Existing in large to small colonies most species are active in the morning and evening or after 

31

precipatation during the day. Those species (Helminthoglypta hertlieni), that occupy the moist 

temperate forests will feed on fungi, green herbaceous plants, and underground roots. Desert 

genera will also feed on fungi and green forbs and feces. 

All of the Helminthoglyptidae are hermaphroditic and some of the snails in this family 

lay eggs. Some species (Helminthoglypta) reproduce throughout the fall and deposit eggs in litter 

or in talus slopes. In northern and central California Helminthoglypta may lay two broods which 

will hatch in the late summer and following spring. In the desert genera, Sonorelix, Sonorella, 

and Eremarionta, these species may lay eggs twice in wet summers or once every three years 

during droughts. Some of the Sonorella are viviparous, the eggs hatching in the uterus where the 

young grow for a time before leaving the adult snail, while other species of this genus will lay 40 

eggs at a time. The desert species may live up to six years, spending most of the dry summer and 

fall in aestivation. 

There are several species and subspecies of Helminthoglypta, Sonorella, Xerarionta, and 

Cahuillus that are considered critically imperiled due to a limited distribution (on mountain tops 

or along a riparian zone), which results in fragmented colonies with little immigration. 

Impacts on these genera also include disruption of talus and foraging habitats by cattle 

grazing, loss of water in riparian zones to agriculture, loss of aspen forests, human recreation, 

invasive plants, and hot ground fires. The status of all western land snails in different habitats are 

important indicators of the general ecosystem health. 

Information is summarized from Pilsbry (1939), NatureServe, Bequaert & Miller (1973), 

Roth & Sadeghian (2006), Northwest Forest Plan (2002), Natural Heritage Research, Sierra 

Nevada and Cascade Mountains Management Plan (1997). 

TAXON AUTHOR 

G- 


Cahuillus greggi (W.B. Miller, 1981) G1G2 CA 

Cahuillus indioensis (Yates, 1890) G2G3 CA 

Chamaearionta aquaealbae (S.S. Berry, 1922) G1 CA 

Eremarionta brunnea (Willett, 1935) G1 CA 

Eremarionta immaculata (Willett, 1937) G1 CA 

Eremarionta millepalmarum (S.S. Berry, 1930) G1 CA 

Eremarionta morongoana (S.S Berry, 1929) G1G3 CA 

Eremarionta newcombi (Pilsbry and Ferriss, 1923) GHQ AZ 

Eremarionta orocopia (Willett, 1939) G1 CA 

Eremarionta rowelli (Newcomb, 1865) G3G4 AZ, CA 

Eremariontoides argus (Edson, 1912) G2 CA 

Helminthoglypta allyniana (S.S. Berry, 1920) G2 CA 

Helminthoglypta allynsmithi Pilsbry, 1939 G1 CA 

Helminthoglypta arrosa (W.G. Binney, 1858) G2G3 CA 

Helminthoglypta avus (Bartsch, 1916) G1 CA 

Helminthoglypta ayresiana (Newcomb, 1861) G1G2 CA 

Helminthoglypta benitoensis Lowe, 1930 G2G4 CA 

Helminthoglypta berryi Hanna, 1929 G1 CA 

Helminthoglypta californiensis (I. Lea, 1838) G1G2 CA 

Helminthoglypta callistoderma Pilsbry, 1917 G1 CA 

Helminthoglypta carpenteri (Newcomb, 1861) G2 CA 

Helminthoglypta caruthersi Willett, 1934 G1 CA 

Helminthoglypta coelata (Bartsch, 1916) G1 CA 

Helminthoglypta concolor Roth and Hochberg, 1988 G1G3 CA 

Helminthoglypta contracostae (Pilsbry, 1895) G1G2 CA 

Helminthoglypta crotalina S.S. Berry, 1928 G1 CA 

32

Helminthoglypta cuyama 


1937 

(W.G. Binney and Bland, 

G3 CA 

Helminthoglypta cypreophila 1869) G5 CA 

Helminthoglypta diabloensis (J.G. Cooper, 1869) G2 CA 

Helminthoglypta dupetithouarsii (Deshayes, 1840) 

Gregg and W.B. Miller, 

G2G3 CA 

Helminthoglypta edwardsi 

1976 G1G3Q CA 

Helminthoglypta euomphalodes S.S Berry, 1938 G1 CA 

Helminthoglypta exarata (Pfeiffer, 1857) G2 CA 

Helminthoglypta expansilabris (Pilsbry, 1898) G2 CA 

Helminthoglypta fairbanksi 

Reeder and W.B. Miller, 

1986 G1 CA 

Helminthoglypta ferrissi Pilsbry, 1924 G1 CA 

Helminthoglypta fieldi Pilsbry, 1930 G1 CA 

Helminthoglypta fisheri (Bartsch, 1904) G1 CA 

Helminthoglypta fontiphila Gregg, 1931 G1 CA 

Helminthoglypta graniticola S.S Berry, 1926 G1 CA 

Helminthoglypta greggi Willett, 1931 


G1 CA 

Helminthoglypta hertleini 

1937 G1 CA, OR 

Helminthoglypta inglesi S.S. Berry, 1938 G1 CA 

Helminthoglypta isabella S.S Berry, 1938 G1 CA 

Helminthoglypta jaegeri S.S Berry, 1928 G1 CA 

Helminthoglypta liodoma S.S Berry, 1938 G1 CA 

Helminthoglypta mailliardi Pilsbry, 1927 G3 CA, OR 

Helminthoglypta micrometalleoides W.B. Miller, 1970 G1 CA 

Helminthoglypta milleri Reeder, 1986 G1 CA 

Helminthoglypta mohaveana S.S Berry, 1927 

Reeder and W.B. Miller, 

G1 CA 

Helminthoglypta montezuma 

1986 G1 CA 

Helminthoglypta morroensis (Hemphill, 1911) G2G3 CA 

Helminthoglypta napaea S.S. Berry, 1938 G1 CA 

Helminthoglypta nickliniana (I. Lea, 1838) G3 CA 

Helminthoglypta orina S.S. Berry, 1938 G1 CA 

Helminthoglypta petricola (S.S Berry, 1916) G1 CA 

Helminthoglypta phlyctaena (Bartsch, 1916) G1G2 CA 

Helminthoglypta piutensis Willett, 1938 G1 CA 

Helminthoglypta proles (Hemphill, 1892) G1 CA 

Helminthoglypta reediana Willett, 1932 G1 CA 

Helminthoglypta salviae Roth, 1987 G2 CA 

Helminthoglypta sanctaecrucis Pilsbry, 1927 G1 CA 

Helminthoglypta sequoicola (J.G. Cooper, 1866) 


G2 CA 

Helminthoglypta similans 

1937 G1 CA 

Helminthoglypta sonoma Pilsbry, 1937 G1 CA 

Helminthoglypta stageri Willett, 1938 G1 CA 

Helminthoglypta stiversiana (J.G. Cooper, 1876) G1G2 CA 

Helminthoglypta talmadgei Roth, 1988 G1G3 CA 

Helminthoglypta taylori Reeder and Roth, 1988 G1 CA 

Helminthoglypta tejonis S.S. Berry, 1930 G1 CA 

Helminthoglypta thermimontis S.S. Berry, 1953 G1 CA 

Helminthoglypta traskii (Newcomb, 1861) G1G2 CA 

Helminthoglypta tudiculata (A. Binney, 1843) G2G3 CA 

Helminthoglypta tularensis (Hemphill, 1892) G1 CA 

Helminthoglypta umbilicata (Pilsbry, 1898) G2 CA 

Helminthoglypta uvasana Roth and Hochberg, 1992 G1G2 CA 

33

Helminthoglypta vasquezi Roth and Hochberg, 1992 G1 CA 

Helminthoglypta venturensis (Bartsch, 1916) G1Q CA 

Helminthoglypta walkeriana (Hemphill, 1911) 


G2 LE CA 

Helminthoglypta waltoni 

1976 G1G3Q CA 

Helminthoglypta willetti (S.S. Berry, 1920) G1 CA 

Herpeteros angelus (Gregg, 1949) G1G2 CA 

Maricopella allynsmithi 


1969 G1 AZ 

Micrarionta beatula Cockerell, 1929 G1 CA 

Micrarionta facta (Newcomb, 1864) G1G2 CA 

Micrarionta feralis (Hemphill, 1901) G1 CA 

Micrarionta gabbi (Newcomb, 1864) G1 CA 

Micrarionta opuntia Roth, 1975 G1 CA 

Micrarionta rufocincta (Newcomb, 1864) G1 CA 

Mohavelix micrometalleus (S.S. Berry, 1930) G1 CA 

Myotophallus rooseveltianus (S.S Berry, 1917) G2 AZ 

Noyo intersessa (Roth, 1987) G2 CA 

Rothelix cuyamacensis (Pilsbry, 1895) G1 CA 

Rothelix lowei (Bartsch, 1918) 

(Reeder and W.B. Miller, 

G1 CA 

Rothelix rhodophila 

1987) 

(Reeder and W.B. Miller, 

G1 CA 

Rothelix warnerfontis 

1988) G1 CA 

Sonorelix avawatzica (S.S. Berry, 1930) G1G2 CA 

Sonorelix baileyi (Bartsch, 1904) G1 CA 

Sonorelix borregoensis (S.S. Berry, 1929) G1 CA 

Sonorelix harperi (Bryant, 1900) G1 CA 

Sonorelix melanopylon (S.S. Berry, 1930) G1 CA 

Sonorelix rixfordi (Pilsbry, 1919) G1 CA 

Sonorella ambigua Pilsbry and Ferriss, 1915 G5 AZ 

Sonorella anchana S.S. Berry, 1948 G1 AZ 

Sonorella animasensis Pilsbry, 1939 G1 NM 

Sonorella apache Pilsbry and Ferriss, 1915 G1 AZ 

Sonorella ashmuni Bartsch, 1904 G2 AZ 

Sonorella baboquivariensis Pilsbry and Ferriss, 1915 G5 AZ 

Sonorella bagnarai W.B. Miller, 1967 G1 AZ 

Sonorella bartschi Pilsbry and Ferriss, 1915 G1 AZ 

Sonorella bequaerti W.B. Miller, 1967 G2 AZ 

Sonorella bicipitis Pilsbry and Ferriss, 1910 G3G4 AZ 

Sonorella binneyi Pilsbry and Ferriss, 1910 G1 AZ 

Sonorella bowiensis Pilsbry, 1905 G1 AZ, CA 

Sonorella bradshaveana W.B. Miller, 1984 G1 AZ 

Sonorella caerulifluminis Pilsbry and Ferriss, 1919 

Fairbanks and Beeder, 

G1G2 AZ 

Sonorella christenseni 

1980 G1 AZ 

Sonorella clappi Pilsbry and Ferriss, 1915 G1 AZ 

Sonorella coloradoensis (Stearns, 1890) G5 AZ 

Sonorella coltoniana Pilsbry, 1939 G1 AZ 

Sonorella compar Pilsbry, 1919 G1 AZ 

Sonorella dalli Bartsch, 1904 G1 AZ 

Sonorella danielsi Pilsbry and Ferriss, 1910 G3 AZ 

Sonorella delicata Pilsbry and Ferriss, 1919 G1 AZ 

Sonorella dragoonensis Pilsbry and Ferriss, 1915 G1 AZ 

Sonorella eremita Pilsbry and Ferriss, 1915 G1 AZ 

Sonorella ferrissi Pilsbry, 1915 G1 AZ 

34

Sonorella franciscana Pilsbry and Ferriss, 1919 G2 AZ 

Sonorella galiurensis Pilsbry and Ferriss, 1919 G2 AZ 

Sonorella grahamensis Pilsbry and Ferriss, 1919 G1 AZ 

Sonorella granulatissima Pilsbry, 1905 G3G4 AZ 

Sonorella hachitana (Dall, 1896) G2 NM 

Sonorella huachucana Pilsbry, 1905 

Gilbertson and Metcalf, 

G4G5 AZ 

Sonorella hueconensis 

2005 


G1G2 TX 

Sonorella imitator 

1974 G2 AZ 

Sonorella imperatrix Pilsbry, 1939 G1 AZ 

Sonorella imperialis Pilsbry and Ferriss, 1923 G1 AZ 

Sonorella insignis Pilsbry and Ferriss, 1919 

Fairbanks and Reeder, 

G1 AZ 

Sonorella macrophallus 

1980 G1 AZ 

Sonorella magdalenensis (Stearns, 1890) G2G3 AZ 

Sonorella meadi W.B. Miller, 1966 G1 AZ 

Sonorella metcalfi W.B. Miller, 1976 G2 NM, TX 

Sonorella micra Pilsbry and Ferriss, 1910 G1G2 AZ 

Sonorella micromphala Pilsbry, 1939 

Christensen and Reeder, 

G1 AZ 

Sonorella milleri 

1981 G2Q AZ 

Sonorella mustang Pilsbry and Ferriss, 1919 G3 AZ 

Sonorella neglecta Gregg, 1951 G1G2 AZ 

Sonorella odorata Pilsbry and Ferriss, 1919 G2 AZ 

Sonorella optata Pilsbry and Ferriss, 1910 G2 AZ 

Sonorella orientis Pilsbry, 1936 G3 NM 

Sonorella papagorum Pilsbry and Ferriss, 1915 G1 AZ 

Sonorella parva Pilsbry, 1905 

Gilbertson and Radke, 

G4 AZ 

Sonorella pedregosensis 

2006 G1G2 AZ 

Sonorella reederi W.B. Miller, 1984 G1 AZ 

Sonorella rinconensis Pilsbry and Ferriss, 1910 G2 AZ 

Sonorella rosemontensis Pislbry, 1939 G3 AZ 

Sonorella russelli W.B. Miller, 1984 G1 AZ 

Sonorella sabinoensis Pilsbry and Ferriss, 1919 G4 AZ 

Sonorella santaritana Pilsbry and Ferriss, 1915 G3G4 AZ 

Sonorella simmonsi W.B. Miller, 1966 G2G3 AZ 

Sonorella sitiens Pilsbry and Ferriss, 1915 G4 AZ 

Sonorella superstitionis Pilsbry, 1939 G3 AZ 

Sonorella todseni W.B. Miller, 1976 G1 NM 

Sonorella tortillita Pilsbry and Ferriss, 1919 G3 AZ 

Sonorella tryoniana Pilsbry and Ferriss, 1923 G1 AZ 

Sonorella vespertina Pilsbry and Ferriss, 1915 G1 AZ 

Sonorella virilis Pilsbry, 1905 G2 AZ 

Sonorella walkeri Pilsbry and Ferriss, 1915 G5 AZ 

Sonorella waltoni W.B. Miller, 1968 G1 AZ 

Sonorella xanthenes Pilsbry and Ferriss, 1923 G2 AZ 

Xerarionta intercisa (W.G. Binney, 1857) G1 CA 

Xerarionta kellettii (Forbes, 1850) G1 CA 

Xerarionta redimita (W.G. Binney, 1858) G1G2 CA 

Xerarionta stearnsiana (Gabb, 1868) G2 CA 

Xerarionta tryoni (Newcomb, 1864) G1 CA 

35

Humboldtianidae 


Humboldtiana comprises ~50 species most of which are endemic to Mexico. Ten species 

in the group are found in the Trans-Pecos mountainous region in West Texas and the Guadalupe 

mountains of New Mexico. The rest of the species are found in mountainous areas south to the 

Trans-Volcanic belt in Mexico D.F. Populations of Humboldtiana occur in isolated mountainous 

habitat and individuals of different species are not known to co-occur. Their low vagility and 

dispersal potential has resulted in high levels of endemism and very restricted distributions 

(Mejía & Zúñiga, 2007). The majority of Humboldtiana species are known only from dry shell 

material and many species remain to be described making designations of relationships within 

the group tentative (Thompson & Brewer, 2000). Also included in the family Humboldtianidae is 

the genus Bunnya, three slug-like species of snail with reduced shells. 

These snails are typically found in high elevation habitat, 10,000-13,000 feet above sea 

level in pine forests, oak forests, pine-oak forests and xerophytic shrubland (Mejía & Zúñiga, 

2007). Humboldtiana have been observed eating lichen and ripe cactus tuna (KEP, pers obs.). 

TAXON AUTHOR 

G- 


Humboldtiana agavophila Pratt, 1971 G1 TX 

Humboldtiana cheatumi Pilsbry, 1935 G2 TX 

Humboldtiana chisosensis Pilsbry, 1927 G1 TX 

Humboldtiana edithae Parodiz, 1954 G1 TX 

Humboldtiana ferrissiana Pilsbry, 1928 G2 TX 

Humboldtiana fullingtoni Cheatum, 1972 G1 TX 

Humboldtiana hoegiana (Pilsbry, 1939) G3 TX 

Humboldtiana palmeri Clench, 1930 G2 TX 

Humboldtiana texana Pilsbry, 1927 G2 TX 

Humboldtiana ultima Pilsbry, 1927 G2 NM, TX 

36

Megomphicidae 


Megomphicidae consists of about eight living species, all occurring in western North 

America between Idaho and western Montana, USA, and northern Baja California, Mexico. 

Most of the species have restricted ranges, some of them being known from only a few localities. 

Fossil species, the earliest dating from the late Cretaceous Period, extend the historic range to 

Alberta, Canada, and eastern Wyoming, USA. The possibility that the localized species are 

"long-branch endemics" (i.e., members of a little-ramified clade that is the sister-group of a much 

more branching clade) lends them special interest for conservation purposes and phylogenetic 

analysis. 

The shell is medium sized to large, many-whorled, discoidal, and conspicuously 

umbilicate, with the lip of the aperture simple or thickened by a ridge, but not reflected. The 

periphery is rounded or compressed. Species of Polygyroidea and Polygyrella develop apertural 

barriers when adult; Polygyrella has one or two radial series of lamellae within the body whorl. 

The reproductive system includes an accessory sac arising from the free oviduct near the 

insertion of the spermathecal duct. 

Species of Glyptostoma are found on rocky hillsides under plant debris, in rock piles, 

wood rat nests, and spaces beneath logs, stumps, and boulders. Ammonitella inhabits talus 

around limestone ledges and leaf litter under shrubs or trees; it also occurs in caves. Polygyrella 

and Polygyroidea are typically found in shaded rockslides. Megomphix occurs in and under 

rotting logs and in caves and rock crevices. 

Information summarized from Pilsbry (1939; 1946), Smith (1957). 

TAXON AUTHOR 

G- 


Ammonitella yatesii J.G. Cooper, 1869 G1 CA 

Glyptostoma gabrielense Pilsbry, 1938 G2 CA 

Glyptostoma newberryanum (W.G. Binney, 1858) G2 CA 

Megomphix californicus A.G. Smith, 1960 G1G2 CA 

Megomphix hemphilli (W.G. Binney, 1879) G3 OR, WA 

Megomphix lutarius H.B. Baker, 1932 G1 OR, WA 

Polygyrella polygyrella 

(Bland and J.G. Cooper, 

1861) G3 ID, MT, OR, WA 

Polygyroidea harfordiana (J.G. Cooper, 1870) G1 CA 

37

Oleacinidae 


A single species of this family is found in the U.S. in Florida; the family is more widely 

distributed in the Greater Antilles. This small (6-8 mm) elongate, ribbed snail was previously 

called Varicella gracillima (Pfeiffer, 1839). Hubricht (1985) states that this snail is a calciphile 

and tends to be found in leaf litter, under rocks or trash, usually in hammocks or under sea grape 

plants above beaches. This species also climbs up the trunks of trees in wet weather. 

TAXON AUTHOR 

G- 


Melaniella gracillima (Pfeiffer, 1839) G4 FL 

Oreohelicidae 


There are 96 species, several subspecies, and two genera of the family Oreohelicidae. 

Approximately 50 species and subspecies are considered critically imperiled or sensitive. 

Species of the genus Radiocentrum are found in southern Arizona, southwestern New Mexico, 

and south into northwestern Chihuahua. The genus Oreohelix is centered in the mountain states, 

south to western Chihuahua, north to British Columbia and Alberta, and east to the Black Hills of 

South Dakota. This includes the states of Colorado, Wyoming, Utah, Nevada, Montana, and 

Idaho. 

The genus Oreohelix range in size from 9 mm to 2.5 cm in diameter. They typically have 

one to two brown, supra-peripheral bands circling the whorls, some with indistinct bands, to no 

bands at all. Most species have different degrees of color varying from brown to blackish red. 

38

The various species have 4 to 6 tubular or carinate whorls and a typically depressed shell, 

sometimes discoidal or pyramidal in some species. The aperature is rounded to angular with no 

teeth, the peristome blunt or sharp, and the lip of the aperature is not reflected. 

Species of the genus Oreohelix are viviparous, the eggs hatching in the uterus where the 

young grow for a time before leaving the adult snail. In the genus Radiocentrum the adults are 

oviparous, the eggs leaving the uterus before hatching. The genus Radiocentrum has species 

located in isolated mountain ranges of Arizona and New Mexico, California, and west Texas. 

The greatest diversity and abundance of species in the genus Oreohelix, including many 

subspecies, are found in two regions that include the Wasatch Mountains of Utah and the 

northwest river basins of Idaho. Many of the subspecies are taxonomically difficult to classify 

because of the great variation in shell morphology within a single canyon or even a single 

colony. Undescribed species and subspecies are still being found today using genetic techniques 

and the dissection of the genitalia. 

The one habitat requirement of both genera is the presence of limestone and dolomite 

talus slopes, although some colonies are found in granite and shale. They exist in a wide variety 

of habitat sites that have vegetation varying from dry sagebrush, aspen forests, and coniferous 

forests to riparian zones that have a brush understory and an overstory of deciduous trees. 

Colonies may occupy an area ranging from 15 meters on dry slopes up to several miles in once 

glaciated canyons. They can be found in elevations of 600 ft. to 10,000 ft. These snails are 

typically active in the spring and fall, feeding on fungi, aspen and willow leaves, green forbs, and 

feces. They are most active in the early morning and evening and after rain showers. 

Most colonies of Oreohelix and Radiocentrum exist in limestone talus on north facing 

slopes. During the winter they hibernate 5 to 8 ft. down into the talus slopes with minimal litter. 

During the summer or in stressful conditions they aestivate 3 to 15 cm inside talus slopes and in 

litter next to streams. During these times they cover their aperature with a thin wall of mucus and 

calcium. 

Smaller species live up to 3 years while larger species may live up to 10 years in 

undisturbed habitat. The isolated mountains of Utah, Montana, southern Arizona, New Mexico, 

and eastern Nevada have colonies of the Oreohelicidae that are endemic and in some cases, 

undescribed. Populations of Oreohelicids throughout their range are impacted through 

overgrazing by cattle, drought, and invasive weeds such as cheat grass, loss of aspen forest, 

colony fragmentation, and ground fires. The status of all western land snails in different habitats 

are important indicators of the general ecosystem health. 

Information summarized from Pilsbry (1939), Bequaert & Miller (1973), Frest (1994), 

Metcalf and Smartt (1997), Weaver et al. (2007), Ports (2004). 

TAXON AUTHOR 

G- 


Oreohelix alpina (Elrod, 1901) G1 MT 

Oreohelix amariradix Pilsbry, 1934 G1G2 MT 

Oreohelix anchana Gregg, 1953 G2 AZ 

Oreohelix barbata Pilsbry, 1905 G1 AZ, NM 

Oreohelix californica S.S. Berry, 1931 G1 CA 

Oreohelix carinifera Pilsbry, 1912 G1 MT, WY; Canada: MB 

Oreohelix concentrata (Dall, 1896) G2 AZ 

Oreohelix confragosa Metcalf, 1974 G1 NM 

Oreohelix elrodi (Pilsbry, 1900) 

J. Henderson and Daniels, 

G1 MT 

Oreohelix eurekensis 

1916 G1 UT 

39

Oreohelix florida Pilsbry, 1939 


GX NM 

Oreohelix grahamensis 

1974 G2 AZ 

Oreohelix hammeri Fairbanks, 1984 G1 ID, OR 

Oreohelix handi Pilsbry and Ferriss, 1918 G1 CA, NV 

Oreohelix haydeni (Gabb, 1869) G2G3 CO, ID, NM, UT 

Oreohelix hemphilli (Newcomb, 1869) G1G3 ID, NV 

Oreohelix hendersoni Pilsbry, 1912 G1 CO 

Oreohelix houghi W.B. Marshall, 1929 G1 AZ, NM 

Oreohelix howardi Jones, 1944 G1 UT 

Oreohelix idahoensis (Newcomb, 1866) G1G2 ID 

Oreohelix intersum (Hemphill, 1890) G1 ID 

Oreohelix jaegeri S.S. Berry, 1931 G1 NV 

Oreohelix jugalis (Hemphill, 1890) G1G2 ID 

Oreohelix junii Pilsbry, 1934 G2 WA 

Oreohelix litoralis Crews and Metcalf, 1982 G1 NM 

Oreohelix loisae Ports, 2004 G1G3 NV 

Oreohelix magdalenae Pilsbry, 1939 G1 NM 

Oreohelix metcalfei Cockerell, 1905 G2 NM 

Oreohelix neomexicana H. A. Pilsbry, 1905 G3 NM, TX 

Oreohelix nevadensis S.S. Berry, 1932 G1 NV 

Oreohelix parawanensis Gregg, 1941 G1 UT 

Oreohelix peripherica (Ancey, 1881) G2 ID, OR, UT 

Oreohelix pilsbryi Ferriss, 1917 G1 NM, WY 

Oreohelix pygmaea Pilsbry, 1913 G1 WY 

Oreohelix strigosa (Gould, 1846) G5 AZ, ID, MT, NM, NV, OR, SD, UT, WA, WY; Canada: AB, BC 

Oreohelix subrudis (Reeve, 1854) G5 AZ, ID, MT, NM, UT, WA; Canada: AB, BC, SK 

Oreohelix swopei Pilsbry and Ferriss, 1917 

J. Henderson and Daniels, 

G1 NM, WY 

Oreohelix tenuistriata 

1916 GH ID 

Oreohelix variabilis Henderson, 1929 G2Q OR 

Oreohelix vortex S.S. Berry, 1932 G1G2 ID 

Oreohelix waltoni Solem, 1975 G1G2 ID 

Oreohelix yavapai Pilsbry, 1905 G5 AZ, MT, UT 

Radiocentrum avalonense (Hemphill, 1905) G1 CA 

Radiocentrum chiricahuana (Pilsbry, 1905) G2 AZ 

Radiocentrum clappi (Ferriss, 1904) G2 AZ 

Radiocentrum ferrissi (Pilsbry, 1915) G1 NM, TX 

Radiocentrum hachetanum (Pilsbry, 1915) G1 NM 

40

Orthalicidae 


Snails of the family Orthalicidae are some of the most commonly encountered and visible 

snails in Florida. Species in the genus Liguus are restricted to the Greater Antilles with one 

highly morphologically, but not genetically variable species (Hillis et al. 1991) present in 

Florida. Shells of Liguus fasciatus are bulimoid in shape, range from 40-72 in length, and have 

incredibly colored and patterned shells, with colors including pink, orange, yellow, green, blue, 

brown, and black. These snails seem to prefer limestone-rich areas and are found on smoothbarked 

trees including Tamarisk, Poisonwood, Black Ironwood, and Pigeonplum. 

Orthalicus species also have colorfully striped shells and their conspicuous abundance 

and colorful stripes have led to a great deal of attention from collectors and taxonomists. 

Orthalicus reses reses is believed to have been extirpated from its native habitat by the invasive 

fire ant Solenopsis invicta (Forys et al. 2001). 

TAXON AUTHOR 

Liguus fasciatus (Muller, 1774) G3 FL 

Orthalicus floridensis Pilsbry, 1899 G3 FL 

G- 


Orthalicus reses (Say, 1830) G2 FL 

41

Philomycidae 


There are 21 species (3 genera) that inhabit the eastern U.S. and Canada. All species of 

Philomycidae (mantleslugs) are slugs. Pallifera are small slugs that are 7-30 mm long, while the 

medium-large Philomycus and Megapallifera are 50-100 mm long. The Philomycidae are 

characterised by a mantle that covers the entire dorsal body, and these slugs lack an internal shell 

and divisions on the foot sole. Philomycus alone employs a calcified dart in mating. Philomycid 

mantles are usually marked with longitudinal rows of gray, brown, or black spots, stripes, or 

chevrons, which form patterns diagnostic to species. Some Pallifera species have solid color 

mantles. 

Philomycids occupy both deciduous and coniferous forest, although they favor old, moist 

deciduous forest (particularly beechwood and basswood). Some species prefer rocky cliffsides 

or upland pine forest. Some species have received special conservation status, particularly those 

that are restricted to small endemic ranges. Slugs are found in moist locations beneath loose 

bark, under dead logs, in tree crevices, and in leaf litter. They are best observed during rainy 

weather and at night when they move and forage on tree trunks (Philomycus, Megapallifera) or 

the ground (Pallifera). As implied by the family name, most species feed on fungus and lichen. 

Megapallifera consume algae that grow on tree trunks and other surfaces. 

This family is restricted to the eastern half of the U.S. and Canada. Species range north 

to Ontario and Nova Scotia, south to Florida and Louisiana, and as far west as Iowa and 

Oklahoma. Pallifera pilsbryi is found in Arizona. 

Information summarized from Pilsbry (1948), Burch (1962), Chichester & Getz (1973), 

Hubricht (1985), Fairbanks (1990). 

TAXON AUTHOR 

G- 


AL, AR, DE, GA, IA, IL, IN, KY, LA, MD, MO, MS, NC, NY, OH, 

Megapallifera mutabilis (Hubricht, 1951) G5 PA, SC, TN, TX, VA, WI, WV; Canada: ON 

Megapallifera ragsdalei (Webb, 1950) G3 AR, IL, KY, MO, OK 

Megapallifera wetherbyi (W.G. Binney, 1874) G2G3 KY, TN, VA 

IA, IL, IN, KY, MA, MD, ME, MI, NC, NY, OH, PA, VA, WI, WV; 

Pallifera dorsalis (A. Binney, 1842) G5 Canada: NS, ON 

AL, FL, GA, IL, IN, KY, MA, MD, MI, MO, MS, NC, OH, SC, TN, 

Pallifera fosteri F.C. Baker, 1939 G5 VA, WI, WV; Canada: ON 

Pallifera hemphilli (W.G. Binney, 1885) G4 IN, MI, NC, TN, VA 

Pallifera marmorea Pilsbry, 1948 G3 AR, IL, KY, LA, MO, OK 

Pallifera megaphallica Grimm, 1961 G5 

Pallifera ohioensis (Sterki, 1908) G5 ME, OH 

42

Pallifera pilsbryi C.D. Miles and Mead, 1960 G2 AZ 

Pallifera secreta Cockerell, 1900 G4 IN, KY, MD, NC, PA, TN, VA, WV 

Pallifera tournescalis Branson, 1968 G1 OK 

Pallifera varia Hubricht, 1953 G2G4 KY, VA 

Philomycus batchi Branson, 1968 G1 KY, OK 

Philomycus bisdodus Branson, 1968 G1 KY, OK 

AL, AR, FL, GA, IA, IL, IN, KS, KY, LA, MD, ME, MI, MO, MS, 

Philomycus carolinianus (Bosc, 1802) G5 NC, NJ, NY, OH, OK, PA, SC, TN, TX, VA, WI, WV; Canada: ON 

DE, GA, KY, MD, ME, NC, NH, NJ, NY, PA, TN, VA, WV; Canada: 

Philomycus flexuolaris Rafinesque, 1820 G5 NS, ON, QC 

Philomycus sellatus Hubricht, 1972 G2G3 AL, TN 

AL, CT, GA, KY, LA, MD, MS, NC, NY, OH, PA, TN, VA, WV; 

Philomycus togatus (Gould, 1841) G5 Canada: ON 

Philomycus venustus Hubricht, 1953 G4 KY, NC, SC, TN, VA, WV 

Philomycus virginicus Hubricht, 1953 G3 KY, NC, TN, VA, WV 

Polygyridae 


There are ~ 230 named extant species in 23 genera in the family Polgyridae in the U.S. and 

Canada, another ~ 40 species occur only in Mexico. 

Polygyrids range in size from 4 - 44 mm. They range in shape from globose to strongly 

keeled or flattened. Apertures have a reflected lip and many have 1-3 apertural teeth. The 

majority of polygyrid shells are a brownish color without color bands. A few banded exceptions 

are: Triodopsis multilineata, Stenotrama fraternum fasciatum, Mesodon elevatus, sAllogona 

profunda, Praticolella spp., Daedalochila texasiana, and D. scintilla. 

Polygyrids can be found occupying deep leaf litter, on top of ground cover, urban areas, 

under rocks and decaying logs and other objects that retain moisture (such as roadside 

cardboard). A few are almost arboreal as Praticolella and Daedalochila species often occupy 

grass, shrubs, and small trees in wet weather. Polygyrids are most typically active at night or 

whenever moisture in the atmosphere is most prevalent, such as after rain. During drought or 

cold months of the year, some species burrow deep in soil or humus. It is assumed that most 

polygyrids feed on mycelia or fruiting bodies of fungi, but they can grow on plant material such 

as lettuce, carrots, and tomatoes when held in the laboratory. 

43

This family is widespread in North America, occupying all U.S. states except Wyoming, 

Colorado, Utah, and Nevada. It also occurs in eastern Canada and southern Alaska. The genus 

Ashmunella occupies the semi-arid mountain ranges of Texas, New Mexico, and Arizona. 

Information summarized from Pilsbry (1941), Cheatum & Fullington (1971), Hubricht (1985). 

TAXON AUTHOR 

G- 


Allogona lombardii A.G. Smith, 1943 G1 ID 

Allogona profunda (Say, 1821) G5 

AL, AR, IA, IL, IN, KS, KY, LA, MD, MI, MN, MO, MS, 

NC, NE, NY, OH, PA, TN, VA, WI, WV; Canada: ON 

Allogona ptychophora (A.D. Brown, 1870) G5 ID, MT, OR, WA; Canada: BC 

Allogona townsendiana (I.Lea, 1838) G3G4 OR, WA; Canada: BC 

Appalachina chilhoweensis (J. Lewis, 1870) G4 KY, NC, TN 

Appalachina sayana (Pilsbry, 1906) G5 

KY, MA, MD, ME, MI, NH, NC, NY, PA, RI, TN, VA, VT, 

WV; Canada: ON, NB, NS, QC 

Ashmunella amblya Pilsbry, 1940 G3 NM, TX 

Ashmunella angulata Pilsbry, 1905 G2G3 AZ 

Ashmunella animasensis Vagvolgyi, 1974 G1 NM 

Ashmunella ashmuni (Dall, 1897) G1 NM 

Ashmunella auriculata Vagvolgyi, 1974 

Clench and W.B Miller, 

G2 NM 

Ashmunella bequaerti 

1966 G1 TX 

Ashmunella binneyi Pilsbry and Ferriss, 1917 G1 NM 

Ashmunella carlsbadensis Pilsbry, 1932 G1 NM, TX 

Ashmunella chiricahuana (Dall, 1896) G1G2 AZ 

Ashmunella cockerelli Pilsbry and Ferriss, 1917 G2 NM 

Ashmunella danielsi Pilsbry and Ferriss, 1915 G1 NM 

Ashmunella edithae Pilsbry and Cheatum, 1951 G1 TX 

Ashmunella esuritor Pilsbry, 1915 G1G2 AZ 

Ashmunella ferrissi Pilsbry, 1905 G1 AZ 

Ashmunella harrisi Metcalf and Smartt, 1977 G1 NM 

Ashmunella hebardi Pilsbry and Vanatta, 1923 G1 NM 

Ashmunella kochii G.H. Clapp, 1908 G1 NM 

Ashmunella lenticula Gregg, 1953 G1 AZ 

Ashmunella lepiderma Pilsbry and Ferriss, 1910 G1G2 AZ 

Ashmunella levettei (Bland, 1881) G1G2 AZ, NM 

Ashmunella macromphala Vagvolgyi, 1974 G1 NM 

Ashmunella mearnsii (Dall, 1895) G2 NM 

Ashmunella mendax Pilsbry and Ferriss, 1917 G1 NM 

Ashmunella mogollonensis Pilsbry, 1905 G1 AZ, NM 

Ashmunella mudgei Cheatham, 1971 G1 TX 

Ashmunella organensis Pilsbry, 1936 G2 NM 

Ashmunella pasonis (Drake, 1951) G2G3 NM, TX 

Ashmunella pilsbryana Ferriss, 1914 G1 AZ 

Ashmunella proxima Pilsbry, 1905 G2G3 AZ 

Ashmunella pseudodonta (Dall, 1897) G1 NM 

Ashmunella rhyssa (Dall, 1897) G1G2 NM 

Ashmunella rileyensis Metcalf and Hurley, 1971 G1 NM 

Ashmunella salinasensis Vagvolgyi, 1974 G1 NM 

Ashmunella sprouli 

R.W. and K.E. Fullington, 

1978 G1G3 TX 

Ashmunella tetrodon Pilsbry and Ferriss, 1915 G3 NM 

Ashmunella thompsoniana (Ancey, 1887) G2G3 NM 

Ashmunella todseni Metalf and Smartt, 1977 G1 NM 

Ashmunella varicifera (Ancey, 1901) G2G3 AZ 

44

Ashmunella walkeri Ferriss, 1904 G1 NM 

Cryptomastix devia (Gould, 1846) G3 OR, WA; Canada: BC 

Cryptomastix germana (Gould, 1851) G4 OR, WA; Canada: BC 

Cryptomastix harfordiana (W.G. Binney, 1886) G3G4 ID, OR, WA 

Cryptomastix hendersoni (Pilsbry, 1928) G1G2 ID, OR, WA 

Cryptomastix magnidentata (Pilsbry, 1940) 

(Bland and J.G. Cooper, 

G1 ID 

Cryptomastix mullani 

1861) G4 ID, MT, OR, WA; Canada: BC 

Cryptomastix populi (Vanatta, 1924) G2 ID, OR, WA 

Cryptomastix sanburni (W.G. Binney, 1886) G1 ID, MT 

Daedalochila auriculata Say, 1818 G3 FL, LA 

Daedalochila auriformis (Bland, 1859) G4 AL, FL, GA, LA, MS, TN, TX 

Daedalochila avara (Say, 1818) G3 FL 

Daedalochila bisontes Coles and Walsh, 2006 G2 AR 

Daedalochila chisosensis (Pilsbry, 1936) G2G3 TX 

Daedalochila delecta (Hubricht, 1976) G2G3 FL 

Daedalochila dorfeuilliana (I. Lea, 1838) G4G5 AR, IL, KS, LA, MO, OK, TN, TX 

Daedalochila fatigiata (Say, 1829) G3 AL, IN, KY, TN 

Daedalochila hausmani (Jackson, 1948) G2 FL 

Daedalochila hippocrepis (Pfeiffer, 1848) G1 TX 

Daedalochila jacksoni (Bland, 1866) G3 AR, KS, MO, OK 

Daedalochila leporina (Gould, 1848) G4G5 AL, AR, IL, IN, KY, LA, MO, MS, OK, TN, TX 

Daedalochila peninsulae (Pilsbry, 1940) G2 FL 

Daedalochila peregrina (Rehder, 1932) G2 AR 

Daedalochila plicata (Say, 1821) G4 AL, GA, IN, KY, TN, VA 

Daedalochila polita (Pilsbry and Hinkley, 1907) G3 TX 

Daedalochila postelliana (Bland, 1859) 

(Pilsbry and Hubricht, 

G3 GA, LA, NC, SC 

Daedalochila scintilla 

1962) G1 TX 

Daedalochila subclausa (Pilsbry, 1899) G3 AL, FL, GA 

Daedalochila triodontoides (Bland, 1861) G3 LA, MS, TX 

Daedalochila troostiana (I. Lea, 1839) G4 AL, KY, TN 

Daedalochila uvulifera (Shuttleworth, 1852) G3 FL 

Euchemotrema fasciatum (Pilsbry, 1940) G3 AL, NC, TN 

Euchemotrema fraternum (Say, 1824) G5 

Euchemotrema hubrichti (Pilsbry, 1940) G1 IL, MI, WI 

AL, AR, CT, DE, GA, IA, IL, IN, KY, LA, MA, MD, ME, 

MI, MN, MO, MS, NC, NH, NJ, NY, OH, OK, PA, RI, SC, 

TN, VA, VT, WI, WV; Canada: NB, NS, ON 

Euchemotrema leai (A. Binney, 1841) G5 

AL, AR, GA, IA, IL, IN, KS, KY, LA, ME, MI, MN, MO, 

MS, NE, NY, OH, OK, PA, SD, TN, TX, VA, WI; Canada: 

NB, ON, QC 

Euchemotrema wichitorum (Branson, 1972) G2 OK 

Fumonelix archeri (Pilsbry, 1940) G1 NC, TN 

Fumonelix christyi (Bland, 1860) G3 GA, LA, NC, SC, TN, VA 

Fumonelix jonesiana (Archer, 1938) G1 NC, TN 

Fumonelix orestes (Hubricht, 1975) G1 NC 

Fumonelix wetherbyi (Bland, 1873) G2G3 KY, TN 

Fumonelix wheatleyi (Bland, 1860) G4 GA, NC,TN, VA 

Hochbergellus hirsutus Roth and W.B. Miller, 1992 G1 OR 

Inflectarius approximans (G.H. Clapp, 1905) G2 AL 

Inflectarius downieanus (Bland, 1861) G3 AL, KY, NC, TN 

Inflectarius edentatus (Sampson, 1889) G2G3 AR, MO, OK 

Inflectarius ferrissi (Pilsbry, 1897) G2 NC, TN 

AL, AR, FL, GA, IL, IN, KS, KY, LA, MI, MO, MS, NC, 

Inflectarius inflectus (Say, 1821) G5 

OH, OK, TX, VA, WV, TN; Canada: ON 

Inflectarius kalmianus (Hubricht, 1965) G3 KY, NC, TN, VA 

45

Inflectarius magazinensis (Pilsbry and Ferriss, 1907) G1 LT AR 

Inflectarius rugeli (Shuttleworth, 1852) G5 AL, GA, IN, KY, NC, SC, TN, VA, WV 

Inflectarius smithi (G.H. Clapp, 1905) G2 AL, TN 

Inflectarius subpalliatus (Pilsbry, 1893) G2 NC, SC, TN 

Inflectarius verus (Hubricht, 1954) G1 NC, SC 

Linisa tamaulipasensis (I. Lea, 1857) G3 TX 

Linisa texasiana (Moricand, 1833) G3G4 AL, AR, KS, LA, MO, MS, NM, OK, TX 

Lobosculum pustula (Ferussac, 1832) G3G4 AL, FL, GA, SC 

Lobosculum pustuloides (Bland, 1858) G3G4 AL, FL, GA, KY, MS, NC, SC, TN, VA 

Mesodon altivagus (Pilsbry, 1900) G2G3 NC, TN 

Mesodon andrewsae W.G. Binney, 1879 G3 NC, TN, VA, WV 

AL, AR, GA, IA, IL, IN, KS, KY, LA, MD, MI, MN, MO, 

MS, NC, NY, OH, OK, PA, TN, TX, VA, WI, WV; Canada: 

Mesodon clausus (Say, 1821) G5 

ON 

AL, AR, IL, IN, KY, MI, MO, MS, NC, NY, OH, OK, TN, 

Mesodon elevatus (Say, 1821) G5 

VA; Canada: ON 

Mesodon mitchellianus (I. Lea, 1839) G4 IN, KY, MI, NC, NY, OH, PA, TN, VA, WV 

Mesodon normalis (Pilsbry, 1900) G5 AL, GA, KY, NC, SC, TN, VA 

Mesodon sanus (Clench and Archer, 1933) G3 AL, LA, TN 

AL, AR, CT, DE, FL, GA, IA, IL, IN, KS, KY, LA, MD, 

ME, MI, MN, MO, MS, NC, NJ, NY, OH, OK, PA, SC, TN, 

Mesodon thyroidus (Say, 1816) G5 

TX, VA, WI, WV; Canada: ON 

Mesodon trossulus Hubricht, 1966 G1 AL 

AL, AR, GA, IA, IL, IN, KY, LA, MD, MI, MO, MS, NC, 

Mesodon zaletus (A. Binney, 1837) G5 

NY, OH, OK, PA, TN, TX, VA, WI, WV; Canada: ON, QC 

Millerelix deltoidea (Simpson, 1889) G2 AR, MO, OK 

Millerelix gracilis Hubricht, 1961 G2G3 TX 

Millerelix lithica (Hubricht, 1961) G3 AR, OK 

Millerelix mooreana (W.G. Binney, 1858) G3 TX 

Millerelix simpsoni (Pilsbry and Ferriss, 1907) G2 AR, OK 

CT, DE, IL, IN, KY, LA, MA, MD, ME, MI, MS, NC, NH, 

NJ, NY, OH, PA, RI, SC, TN, VA, VT, WI, WV; Canada: 

Neohelix albolabris (Say, 1816) G5 

NB, QC 

Neohelix alleni (Sampson, 1883) G5 AL, AR, IA, IL, KS, LA, MI, MN, MO, MS, OK, TN 

CT, KY, MA, MD, ME, NC, NH, NY, OH, PA, VT, WV, 

Neohelix dentifera (A. Binney, 1837) G5 

VA; Canada: QC 

Neohelix divesta (Gould, 1848) G3G4 AR, KS, LA, MO, OK, TX 

Neohelix lioderma (Pilsbry, 1902) G1G2 OK 

Neohelix major (A. Binney, 1837) G4G5 AL, DE, GA, MD, MS, NC, NJ, SC, TN, VA 

Neohelix solemi Emberton, 1988 G4 MD, NC, NJ, NY 

Patera appressa (Say, 1821) G4 IL, KY, MD, NC, OH, SC, VA, WV, AL, IN, TN 

Patera binneyana (Pilsbry, 1899) G2G3 AR, OK, TX 

Patera clarki (I. Lea, 1858) G3 GA, KY, NC, SC, TN 

Patera clenchi (Rehder, 1932) G1 AR 

Patera indianorum (Pilsbry, 1899) G2G3 AR, OK 

Patera kiowaensis (Simpson, 1888) G2G3 AR, OK 

Patera laevior (Pilsbry, 1940) G4 AL, GA, IN, KY, MD, MS, NC, OH, TN, VA 

Patera leatherwoodi Pratt, 1971 G1 TX 

Patera panselenus (Hubricht, 1976) G2 KY, VA, WV 

Patera pennsylvanica (Green, 1827) G4 IL, IN, KY, MI, MO, OH, PA, WV, TN; Canada: ON 

Patera perigrapta Pilsbry, 1894 G5 AL, AR, GA, KY, LA, MO, MS, NC, SC, TN 

Patera roemeri (Pfeiffer, 1848) G3G4 OK, TX 

Patera sargentiana 

(C.W. Johnson and Pilsbry, 

1892) G2 AL 

Polygyra cereolus (Muhlfeld, 1816) G4 AL, FL, GA, HI, KY, LA, MS, NC, SC, TX 

Polygyra septemvolva Say, 1818 G5 AL, FL, GA, LA, MI, MS, NC, NM, SC, TX 

Praticolella bakeri Vanatta, 1915 G2G3 FL 

46

Praticolella berlandieriana (Moricand, 1833) G2G3 TX 

Praticolella candida Hubricht, 1983 G2 TX 

Praticolella griseola (Pfeiffer, 1841) G3 TX 

Praticolella jejuna (Say, 1821) G3 FL, SC 

Praticolella lawae (J. Lewis, 1874) G3 AL, GA, MS, NC, TN 

Praticolella mobiliana (I. Lea, 1841) G3 AL, FL, GA, MS 

Praticolella pachyloma (Menke, 1847) G3G4 TX 

Praticolella taeniata Pilsbry, 1940 G3G4 TX 

Praticolella trimatris Hubricht, 1983 G2 TX 

Stenotrema altispira (Pilsbry, 1894) G3 NC, TN, VA 

Stenotrema angellum Hubricht, 1958 G4 IN, KY, TN 

Stenotrema barbatum (G.H. Clapp, 1904) G5 

AL, CT, DE, GA, IA, IL, IN, KS, KY, LA, MA, MD, MI, 

MN, MO, MS, NC, NJ, NY, OH, PA, SC, TN, VA, WI, 

WV; Canada: ON 

Stenotrema barbigerum (Redfield, 1856) G3G4 AL, GA, KY, NC, SC, TN 

Stenotrema blandianum (Pilsbry, 1903) G2 AR, MO 

Stenotrema brevipila (G.H. Clapp, 1907) G2 AL, GA 

Stenotrema burringtoni Grimm, 1971 G5Q CT, NJ, NY, VA, WV 

Stenotrema calvescens Hubricht, 1961 G3 AL, TN 

Stenotrema cohuttense (G.H. Clapp, 1914) G2 GA, TN 

Stenotrema deceptum (G.H. Clapp, 1905) G3G4 AL, TN 

Stenotrema depilatum (Pilsbry, 1895) G2 NC, TN 

Stenotrema edgarianum (I. Lea, 1841) G2G3 TN 

Stenotrema edvardsi (Bland, 1856) G4G5 GA, KY, NC, PA, TN, VA, WV 

Stenotrema exodon (Pilsbry, 1900) G2 AL, GA, TN 

Stenotrema florida Pilsbry, 1940 G3 AL, FL, GA 

AL, CT, DE, IN, KS, KY, MA, MD, MI, MS, NC, NJ, NY, 

Stenotrema hirsutum (Say, 1817) G5 

OH, PA, TN, VA, WI, WV, IL; Canada: NS, ON 

Stenotrema labrosum (Bland, 1862) G3G4 AR, IA, LA, MO, OK 

Stenotrema magnafumosum (Pilsbry, 1900) G4 AL, GA, NC, SC, TN 

Stenotrema maxillatum (Gould, 1848) G3 AL, GA 

Stenotrema morosum Hubricht, 1978 GH TN 

Stenotrema pilsbryi (Ferriss, 1900) G2 AR, OK 

Stenotrema pilula (Pilsbry, 1900) G3G4 GA, NC, SC, TN, VA 

Stenotrema simile Grimm, 1971 G2 MD, WV 

Stenotrema spinosum (I. Lea, 1830) G4 AL, GA, MS, TN, VA 

AL, AR, GA, IL, IN, KS, KY, LA, MD, MO, MS, NC, OH, 

Stenotrema stenotrema (Pfeiffer, 1842) G5 

OK, SC, TN, TX, VA, WV 

Stenotrema unciferum (Pilsbry, 1900) G2 AR, OK 

Stenotrema waldense Archer, 1938 G2 TN 

Trilobopsis loricata (Gould, 1846) G2G3 CA, OR 

Trilobopsis penitens (Hanna and Rixford, 1923) G1 CA 

Trilobopsis roperi (Pilsbry, 1889) G1 CA 

Trilobopsis tehamana (Pilsbry, 1928) G1 CA 

Trilobopsis trachypepla (S.S. Berry, 1933) G1 CA 

Triodopsis alabamensis (Pilsbry, 1902) G4 AL, GA, SC, TN, VA 

Triodopsis anteridon Pilsbry, 1940 G3 KY, TN, VA, WV 

Triodopsis burchi Hubricht, 1950 G3 NC, VA, WV 

Triodopsis claibornensis Lutz, 1950 G2 KY, TN 

Triodopsis complanata (Pilsbry, 1898) G2 KY, TN 

Triodopsis cragini Call, 1886 G4 AR, KS, LA, MO, OK, TX 

Triodopsis discoidea (Pilsbry, 1904) G3 IL, IN, KY, MO, OH 

Triodopsis fallax (Say, 1825) G5 DE, MD, NC, NJ, PA, SC, TN, VA, WV 

Triodopsis fraudulenta (Pilsbry, 1894) G4 IL, KY, MD, PA, TN, VA, WV 

Triodopsis fulciden Hubricht, 1952 G1G2 NC 

Triodopsis henriettae (Mazyck, 1877) G3 TX 

47

Triodopsis hopetonensis (Shuttleworth, 1852) G4G5 AL, FL, GA, KY, LA, MD, MO, MS, NC, NJ, SC, TN, VA 

Triodopsis juxtidens (Pilsbry, 1894) G5 GA, KY, MD, ME, NC, NJ, NY, PA, SC, VA, WV 

Triodopsis messana Hubricht, 1952 G4 NC, SC, VA 

Triodopsis neglecta (Pilsbry, 1899) G3 AR, KS, MO, OK 

Triodopsis obsoleta (Pilsbry, 1894) G4 MD, NC, VA 

Triodopsis palustris Hubricht, 1958 G3 FL, GA, SC 

Triodopsis pendula Hubricht, 1952 G3 NC, VA 

Triodopsis picea Hubricht, 1958 G3 MD, PA, VA, WV 

Triodopsis platysayoides (Brooks, 1933) 

Brooks and McMillan, 

G1 LT WV 

Triodopsis rugosa 

1940 G1 TN, VA, WV 

Triodopsis soelneri (J.B. Henderson, 1907) G2 NC 

Triodopsis tennesseensis (Walker and Pilsbry, 1902) G4 AL, GA, IN, KY, NC, TN, VA, WV 

AL, CT, DE, GA, IL, IN, KY, MA, MD, MI, MS, NC, NH, 

NJ, NY, OH, PA, SC, TN, VA, VT, WI, WV; ; Canada: ON, 

Triodopsis tridentata (Say, 1816) G5 

QC 

Triodopsis vannostrandi (Bland, 1875) G4 AL, FL, GA, SC 

AL, GA, IL, IN, KY, MI, MS, NC, NY, OH, PA, TN, VA, 

Triodopsis vulgata Pilsbry, 1940 G5 

WI; Canada: ON 

Triodopsis vultuosa (Gould, 1848) G3G4 AR, LA, TX 

Vespericola armiger (Ancey, 1881) G1 CA 

Vespericola columbianus (I. Lea, 1838) 

(Pilsbry and Henderson, 

G5 AK, OR, WA; Canada: BC 

Vespericola depressa 

1936) G2Q OR, WA 

Vespericola embertoni Roth and Miller, 2000 G2G3 CA 

Vespericola eritrichius (S.S. Berry, 1939) G1 CA 

Vespericola euthales (Berry, 1939) G3 CA, OR 

Vespericola haplus (S.S Berry, 1933) G1 CA 

Vespericola karokorum Talmadge, 1962 G2G3 CA 

Vespericola klamathicus Roth and W.B. Miller, 1995 G2 CA 

Vespericola marinensis Roth and W.B. Miller, 1993 G2G3 CA 

Vespericola megasoma (Pilsbry, 1928) G3 CA, OR 

Vespericola ohlone Roth, 2003 GX CA 

Vespericola orius (S.S. Berry, 1933) G1G2 CA 

Vespericola pilosus (J. Henderson, 1928) G2G3 AK, CA, OR, WA 

Vespericola pinicola (S.S Berry, 1916) G1 CA 

Vespericola pressleyi Roth, 1985 G1 CA 

Vespericola rhodophila Roth and Miller, 2000 

Cordero and W.B. Miller, 

G1G3 CA 

Vespericola rothi 

1995 G1 CA 

Vespericola sasquatch Roth and Miller, 2000 

Cordero and W.B. Miller, 

G1G3 CA 

Vespericola scotti 

1995 G1 CA 

Vespericola shasta (S.S Berry, 1921) G1 CA 

Vespericola sierranus (S.S. Berry, 1921) G2 CA, OR 

Webbhelix chadwicki (Ferriss, 1907) G1Q KS, NE 

Webbhelix multilineata (Say, 1821) G5 

AR, IA, IL, IN, KS, KY, MD, MI, MN, MO, NE, NY, OH, 

PA, WI, WV, TN; Canada: ON 

Xolotrema caroliniense (I. Lea, 1834) G4 AL, AR, GA, LA, MS, NC, SC, TN 

AR, KY, MA, MD, MO, MS, NC, NJ, NY, OH, PA, VT, 

Xolotrema denotatum (Ferussac, 1821) G5 

WV, AL, IN, MI, TN, VA; Canada: ON 

AL, AR, DE, GA, IA, IL, IN, KY, LA, MD, MO, NJ, OH, 

Xolotrema fosteri (F.C. Baker, 1921) G5 

TN, TX, WI 

Xolotrema obstrictum (Say, 1821) G4 AL, IL, IN, KY, LA, MS, TN 

Xolotrema occidentale (Pilsbry and Ferriss, 1907) G1 AR 

48

Punctidae 

Aydin Örstan, Carnegie Museum of Natural History 

Punctum is a Holarctic genus. The North American Punctum species are among our 

smallest land snails. In fact, P. smithi, whose adult shell diameters barely reach ~1.2 mm, is one 

of the smallest land snails in the world. Punctum are primarily woodland snails that are 

widespread throughout North America. The range of P. conspectum [P. conspectum is a junior 

synonym of Paralaoma servilis] extends to Alaska. Although Punctum specimens may, on 

occasion, be abundant in litter samples, because of their diminutive sizes, live Punctum are 

difficult to observe and study in the wild. Consequently, virtually nothing is known about the 

natural histories of individual species. The European P. pygmaeum is known to be able to 

reproduce without mating in captivity. The anatomies of most North American Punctum species 

have also not been studied. 

Paralaoma servilis (also known as P caputspinulae) is known from a few disjunct 

locations in western North America. It is probably a non-native species. 

TAXON AUTHOR 

G- 


Paralaoma servilis (Shuttleworth, 1852) G5 CA, ID, NM, WA; Canada: BC 

Punctum blandianum Pilsbry, 1900 G4 AL, KY, NC, TN, VA 

Punctum californicum Pilsbry, 1898 G5 AK, AZ, CA, CO, MT, NM, OR, SD, WA; Canada: BC 

Punctum hannai Roth, 1985 G1G3 CA 

AL, AR, CA, CO, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, 

ME, MI, MN, MO, MS, NC, NE, NH, NJ, NM, NY, OH, OK, OR, 

PA, SC, SD, TN, TX, UT, VA, VT, WI, WV; Canada: AB, NB, NS, 

Punctum minutissimum (I. Lea, 1841) G5 ON, QC, NF 

Punctum randolphi (Dall, 1895) G4 AK, CA, ID, OR, WA; Canada: BC 

Punctum smithi Morrison, 1935 G4 AL, IL, IN, KY, MD, NC, TN, VA, WV 

Punctum vitreum (H.B. Baker, 1930) G5 

AL, AR, DE, IA, IL, IN, KY, LA, MD, MN, MO, MS, NC, NJ, OK, 

PA, TN, TX, VA, WI, WV 

“Zonites” diegoensis Hemphill, 1892 G1Q CA 

49

Pupillidae 

Jeffrey C. Nekola, University of New Mexico 

Nine genera (Bothriopupa, Chaenaxis, Columella, Gastrocopta, Pupilla, Pupisoma, 

Pupoides, Sterkia and Vertigo) and ~125 species are currently thought to constitute this family in 

Canada and the continental United States, with an unknown number of additional taxa residing in 

Mexico. All have shells taller than wide (approaching equal in Pupisoma), with maximum 

dimensions ranging from 1.2 mm (Vertigo hebardi) - 5.0 mm (Pupoides albilabris). Shell color 

typically ranges from yellow-brown-brick red, while in Chaenaxis and Gastrocopta this often 

tends to be white or horn-yellow. Aperture margins vary considerably, ranging from simple to 

reflected to thickened. Apertural lamellae range from 0-9 or more, with their number and 

arrangement being vital for species-level taxonomy. Pupillids occur in almost all habitat types, 

ranging from arctic tundra and semi-tropical grasslands to forests, peatlands, and bluffs. While 

individuals tend to be most common in leaf litter accumulations, they may also be found in wet 

turf, vertical rock outcrops (e.g. Vertigo meramecensis, Gastrocopta corticaria) or mossy tree 

trunks (e.g. Vertigo rowelli). Based on observations of lab-reared populations, most Pupillids 

appear to be generalist detritivores that feed on a variety of fungal hyphae and/or algal mats. All 

species are also capable of uniparental reproduction, allowing for the founding of new 

populations by the successful establishment of only a single individual. 

The global biodiversity center for Gastrocopta, Pupilla, and Vertigo is in North America 

which supports at least 2/3 of all known global taxa. While found in every U.S. state and 

Canadian province, the Pupillidae become particularly abundant and diverse with increasing 

latitude, constituting more than 90% of taxa and individuals in taiga and tundra situations. Many 

of these have yet to be scientifically described. This family is also abundant and diverse in some 

lower latitude landscapes, including the Upper Midwest, the Southern Appalachians, ‘sky island’ 

mountains of the Southwest, the central Rockies, and the fog-belt of the California coast. 

Information summarized from Pilsbry (1948), Hubricht (1985). 

TAXON AUTHOR 

G- 


Bothriopupa variolosa (Gould, 1848) G1 FL 

Chaenaxis tuba (Pilsbry and Ferriss, 1906) G4 AZ 

Columella columella (Martens, 1830) G5 

Columella edentula (Draparnaud, 1805) G5 

Columella simplex (Gould, 1840) G5Q 

AK, AZ, CO, IA, IL, ID, IN, KS, KY, MI, MO, MS, MT, NE, NM, 

OH, SE, TX, UT, WA, WI, WY; Canada: AB, BC, ON 

AK, AL, AR, CT, GA, IA, ID, IL, IN, KY, MA, MD, ME, MI, MO, 

MS, MT, NJ, NY, OH, OK, OR, PA, TN, WA, WI, WV, WY; 

Canada: AB, BC, LB, MB, NB, NF, NS, ON, QC, YT 

AL, AR, CT, GA, IA, IL, IN, KY, MA, MD, ME, MI, MN, MO, MS, 

NC, NM, NY, OH, OK, PA, SD, TN, TX, VA, WI, WV; Canada: ON 

50

Gastrocopta abbreviata (Sterki, 1909) G4 AL, IA, IL, KS, LA, MN, MO, MS, ND, NE, OK, SD, TX, WI 

AL, AR, DE, FL, GA, IA, IL, IN, KS, KY, LA, MD, ME, MI, MN, 

MO, MS, NC, ND, NE, NJ, NM, NY, OH, OK, PA, SC, SD, TN, TX, 

Gastrocopta armifera (Say, 1821) G5 VA, VT, WI, WV; Canada: QC 

Gastrocopta ashmuni (Sterki, 1898) G4G5 AZ, NM, TX, UT 

Gastrocopta carnegiei (Sterki, 1916) G1G3Q OH 

Gastrocopta clappi (Sterki, 1909) G4G5 AL, KY, TN, VA 

Gastrocopta cochisensis (Pilsbry and Ferriss, 1910) G3G4 AZ, NM 

AL, AR, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, 

MN, MO, MS, NC, ND, NE, NJ, NM, NY, OH, OK, PA, SC, SD, TN, 

Gastrocopta contracta (Say, 1822) G5 TX, VA, VT, WI, WV; Canada: ON, QC 

AL, CT, FL, GA, IA, IL, IN, KS, KY, LA, MD, ME, MI, MN, MO, 

MS, NC, NE, NJ, NY, OH, OK, PA, TN, TX, VA, VT, WI, WV; 

Gastrocopta corticaria (Say, 1816) G5 Canada: NB, ON, QC 

Gastrocopta cristata (Pilsbry and Vanatta, 1900) G5 AZ, CO, DE, KS, LA, MD, MO, NJ, NM, OK, TX, VA 

Gastrocopta dalliana (Sterki, 1898) G2G4 AZ, NM, TX 

AR, IA, IL, IN, KS, KY, MI, MN, MO, NC, ND, NE, NM, NY, OH, 

Gastrocopta holzingeri (Sterki, 1889) G5 OK, SD, TN, TX, VA, WI, WV; Canada: BC, ON, QC 

AL, CA, FL, GA, KS, LA, MD, MO, MS, NC, NE, NJ, NM, OK, SD, 

Gastrocopta pellucida (Pfeiffer, 1841) G5 TX, UT, VA, WI 

AL, AR, CA, DE, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, 

MI, MN, MO, MS, NC, NE, NJ, NY, OH, OK, PA, SC, SD, TN, TX, 

Gastrocopta pentodon (Say, 1822) G5 VA, VT, WI, WV; Canada: AB, NS, ON, QC 

Gastrocopta pilsbryana (Sterki, 1890) G4G5 AZ, CO, NM, TX, UT 

AL, AR, FL, GA, IA, IL, IN, KS, KY, LA, MD, MO, MS, NC, NE, 

Gastrocopta procera (Gould, 1840) G5 NM, NY, OH, OK, SC, SD, TN, TX, VA, WI 

Gastrocopta prototypus (Pilsbry, 1899) G1 AZ, NM 

Gastrocopta quadridens Pilsbry, 1916 G2G3 AZ, NM, UT 

Gastrocopta riograndensis (Pilsbry, 1916) GH TX 

Gastrocopta riparia Pilsbry, 1916 G3G5 AL, FL, GA, LA, MS, TX 

Gastrocopta rogersensis Nekola and Coles, 2001 G3G4 AR, IA, IL, MO, WI 

Gastrocopta ruidosensis (Cockerell, 1899) G1 KS, NE, NM, OK, TX 

Gastrocopta rupicola (Say, 1821) G3G4 AL, FL, GA, LA, MS, NC, SC, TN, TX 

Gastrocopta servilis (Gould, 1843) G3G4 FL, HI 

IA, IL, IN, KS, KY, MI, MN, MO, ND, NY, OH, SD, WI; Canada: 

Gastrocopta similis (Sterki, 1909) G5 AB, ON 

Gastrocopta sterkiana Pilsbry, 1917 G2G3Q AR, OK, TX 

AL, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, 

MS, NC, NE, NJ, NY, OH, OK, PA, SC, SD, TN, TX, VA, WI, WV; 

Gastrocopta tappaniana (C.B. Adams, 1842) G5 Canada: AB, ON 

Nearctula rowellii (Rowell, 1861) G3 CA 

Pupilla blandi E.S. Morse, 1865 G4 KS, MO, ND, NE, NM, SD, TX, UT 

Pupilla hebes (Ancey, 1881) G5 AK, CA, ID, MT, NM, SD, TX, UT, WA, WY; Canada: AB, BC 

IA, IL, KS, MA, MD, ME, MI, MN, MO, ND, NE, NJ, NM, NY, OH, 

Pupilla muscorum (Linnaeus, 1758) G5 OK, SD, TX, UT, VA, VT, WI, WV; Canada: AB, NF, NS, ON, QC 

Pupilla sonorana (Sterki, 1899) G4G5 NM, TX 

Pupilla syngenes (Pilsbry, 1890) G4 NM, TX, UT 

Pupisoma dioscoricola (C.B. Adams, 1845) G3 AL, FL, GA, LA, MS, SC, TX 

Pupisoma macneilli (G.H. Clapp, 1918) G3 AL, FL, GA, LA, MS, SC, TX 

Pupisoma minus Pilsbry, 1920 G2Q FL 

AL, AR, CA, DE, FL, GA, IA, IL, IN, KS, KY, LA, MD, ME, MI, 

MO, MS, NC, NE, NJ, NM, NY, OH, OK, PA, RI, SC, SD, TN, TX, 

Pupoides albilabris (C.B. Adams, 1841) G5 UT, VA, VT, WI, WV; Canada: ON, QC 

Pupoides hordaceus (Gabb, 1866) G4 AZ, CO, KS, NM, OK, TX, UT, WY 

Pupoides inornatus Vanatta, 1915 G2 KS, NE, NM, OK, SD, TX 

Pupoides modicus (Gould, 1848) G3 FL, GA, LA 

Sterkia clementina (Sterki, 1890) G2G3 CA 

Sterkia eryiesii (Pilsbry, 1899) G1 FL 

Sterkia hemphilli (Sterki, 1890) G2 CA 

51

Vertigo alabamensis G.H. Clapp, 1915 G3 AL, NC, SC 

Vertigo allyniana S.S Berry, 1919 G1 CA 

Vertigo andrusiana (Pilsbry, 1899) G2G3 CA, OR, WA; Canada: BC 

Vertigo arthuri von Martens, 1882 G3Q AK, MN, ND, SD, WY; Canada: AB, BC, MB, ON, YT 

Vertigo berryi Pilsbry, 1919 G1 AZ, CA 

Vertigo binneyana Sterki, 1890 G1 IA, KS, MT, NM, WI; Canada: AB, BC, MB, ON 

Vertigo bollesiana (E.S. Morse, 1865) G4 

IA, KY, MA, MD, ME, MI, MN, NC, NH, NY, OH, PA, TN, VA, 

VT, WI, WV; Canada: NS, ON, QC 

Vertigo brierensis (Leonard, 1972) G1 IA, IL, MN, WI; Canada: MB 

Vertigo clappi Brooks and Hunt, 1936 G1G2 KY, TN, VA, WV 

Vertigo columbiana Pilsbry and Vanatta, 1900 G5 AK, OR, UT, WA; Canada: BC 

Vertigo concinnula Cockerell, 1897 G4G5 AZ, CO, ID, KS, NE, NM, UT, WA, WY 

Vertigo conecuhensis G.H. Clapp, 1915 G2 AL 

Vertigo cristata (Sterki in Pilsbry, 1919) G5 MA, ME, MI, MN, NY, WI, WV; Canada: AB, BC, ON 

Vertigo dalliana Sterki, 1890 G1 CA, OR 

AZ, DE, IA, IL, IN, KS, MA, ME, MI, MN, MO, MT, ND, NE, NM, 

Vertigo elatior Sterki, 1894 G5 NY, OH, SD, TX, UT, VA, WI, WY; Canada: AB, BC, ON, NF 

AL, IA, IL, IN, KS, KY, LA, MD, ME, MI, MN, MO, NC, NH, NJ, 

NM, NY, OH, PA, TN, TX, UT, VA, WI, WV; Canada: AB, BC, NB, 

Vertigo gouldi (A. Binney, 1843) G5 NF, NS, ON, QC 

Vertigo hannai Pilsbry, 1919 G1 IL, KS; Canada: ON 

Vertigo hebardi Vanatta, 1912 G1 FL 

Vertigo hinkleyi Pilsbry, 1921 G3 AZ, NM 

Vertigo hubrichti Pilsbry, 1934 G3 IA, IL, IN, KY, MI, MN, MO, MS, NE, SD, WI; Canada: MB 

Vertigo idahoensis Pilsbry, 1934 G1G2 ID 

Vertigo malleata Coles and Nekola, 2007 G5 AL, FL, GA, MA, ME, NC, NJ, SC 

Vertigo meramecensis Van Devender, 1979 G2G3 IA, IL, LA, MN, MO; Canada: MB 

AL, AR, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, 

MO, MS, NC, NE, NJ, NM, NY, OH, OK, PA, SC, SD, TN, TX, VA, 

Vertigo milium (Gould, 1840) G5 VT, WI, WV; Canada: ON, QC 

AK, CA, IA, ID, IL, IN, KS, KY, ME, MI, MN, ND, NE, NM, SD, 

Vertigo modesta (Say, 1824) G5 TX, UT, WA, WI; Canada: AB, BC, LB, NF, NS, ON 

IL, IN, MA, ME, MI, MN, NJ, NY, OH, WI; Canada: AB, MB, ON, 

Vertigo morsei Sterki, 1894 G3 QC 

Vertigo nylanderi Sterki, 1909 G3G4 ME, MI, MN, WI; Canada: MB, NS, ON 

Vertigo occidentalis Sterki, 1907 G1Q CA 

Vertigo occulta Leonard, 1972 G2 IA, IL, MN, WI; Canada: MB 

Vertigo oralis Sterki, 1898 G5 AL, FL, GA, LA, MD, MS, NC, SC, TX, VA 

Vertigo oscariana Sterki, 1890 G4 AL, AR, FL, GA, KY, LA, MD, MO, MS, NC, TN, TX, VA, WV 

AK, AL, AR, CA, CT, DE, FL, GA, IA, ID, IL, IN, KS, KY, LA, 

MA, MD, ME, MI, MN, MO, MS, NC, ND, NE, NJ, NM, NY, OH, 

OK, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV,; Canada: 

Vertigo ovata Say, 1822 G5 AB, BC, LB, NS, ON, QC 

IN, ME, MI, MN, NY, SD, VT, WI, WY; Canada: AB, NF, NS, ON, 

Vertigo paradoxa Sterki, 1900 G4G5Q QC 

Vertigo parvula Sterki, 1890 G3 KY, NC, OH, TN, VA 

Vertigo perryi Sterki, 1905 G3G4 CT, MA, ME, NH, RI 

DE, IA, IN, MA, MD, ME, MI, MN, MO, NJ, NY, OH, PA, TN, VA, 

Vertigo pygmaea (Draparnaud, 1801) G5 WI, WV; Canada: NS, ON 

Vertigo rugosula Sterki, 1890 G4 AL, AR, FL, KY, LA, MS, OK, SC, TN, TX 

Vertigo sterkii Pilsbry, 1919 G2 CA 

Vertigo teskeyae Hubricht, 1961 G5 AL, DE, FL, GA, LA, MD, MS, NC, SC, TN, TX, VA 

AR, DE, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MO, MS, NE, 

Vertigo tridentata Wolf, 1870 G5 NJ, NY, OH, OK, PA, SD, TN, TX, VA, WI, WV; Canada: ON 

IA, IL, IN, KY, MA, MD, ME, MI, MO, MS, NC, NY, OH, OK, PA, 

Vertigo ventricosa (E.S. Morse, 1865) G5 TN, VA, VT, WI, WV; Canada: NB, NS, ON, QC 

Vertigo wheeleri Pilsbry, 1928 G1 AL 

52

Sagdidae 


This family is distributed in Central America and the West Indies. A single species, 

Lacteoluna selenina, is found in North America (S. Florida). It has a small (ca. 5 mm), 

depressed, umbilicate, dull-white shell. The whorls are “shouldered”. The outer lip of the 

aperture remaines straight and thin even in fully grown specimens. 

They live in areas with some tree or shrub cover under rocks and plant debris. 

TAXON AUTHOR 

G- 


Lacteoluna selenina (Gould, 1848) G2 FL 

Spiraxidae 


There are only 4 species of the carnivorous snails in this family in the U.S. and Canada, 

but several hundred species of several genera in Mexico, the Caribbean, Central and South 

America. Euglandina rosea is a large (up to 76 mm length, up to 30 mm width), common snail 

of the southeastern US and has been introduced to many places worldwide, often intentionally, 

into CA as well as numerous Pacific Islands. The introductions of E. rosea rate as one of the 

worst described cases of a failed attempt at biological pest control resulting in heavy predation 

pressure on native land snails of those areas (Cowie, 2001; Lydeard, et al. 2004). In Florida they 

can be found foraging for terrestrial snails on the ground, or in wet weather, climbing on the 

stems and leaves of small trees, ~1 m off the ground (Davis, Perez & Bennett, 2004). This 

53

species has even been observed foraging underwater for aquatic snail in Hawaii (Kinzie, 1992). 

This species has been observed in a variety of habitats, in woods dominated by salt cedar (Davis, 

Perez & Bennett, 2004), roadsides, edges of marsh, and disturbed urban areas (Hubricht, 1985). 

Euglandina singleyana (placed in the genus Glandina by Schileyko) is a large snail (up to 

51 mm length) distributed across Central Texas ranging from Galveston County to Val Verde 

County with a few specimens collected in Mexico near the Texas border (Perez & Strenth, 

2003). In the eastern part of its range this snail is found under rocks and logs in wooded stream 

valleys, in the western part of the range it is found under fallen Yucca and under rocks in desert 

shrub habitat dominated by Lechugilla (Fullington & Pratt, 1974). These snails are active on 

humid days, early morning, or after rain. They can be observed following the trails of prey snails 

and often leave small piles of empty shells of their favorite prey near rocky hiding places (KEP 

pers obs., Humboldtiana spp.). E. texasiana is a calciphile, found in much wetter places where it 

stays damp. This species can also be found crawling on buildings in urban areas. The range of 

this species is the Rio Grande Valley of Texas south throughTamaulipas and San Luis Potosi. E. 

texasiana is smaller than the other two U.S. Euglandina species reaching maximum of ~34 mm 

length (Pilsbry, 1946). 

Pseudosubulina cheatumi is the northernmost respresentative of a largely tropical group. 

This species is restricted to leaf litter and stabilized talus in bottoms of canyons as well as in the 

Evergreen Zone in the Chisos Mountains, Big Bend National Park, Texas (Fullington & Pratt, 

1974). 

TAXON AUTHOR 

G- 


Euglandina rosea (Ferussac, 1818) G5 AL, CA, FL, GA, HI, LA, MS, NC, SC, TX 

Euglandina singleyana (W.G. Binney, 1892) G3 TX 

Euglandina texasiana (Pfeiffer, 1857) G1G2 TX 

Pseudosubulina cheatumi Pilsbry, 1950 G1 TX 

Strobilopsidae 


These snails are very small with uniformly brownish shells. Shells are usually lowconical 

or beehive-shaped with a narrow umbilicus, and usually sculptured with conspicuous 

transverse ribs. One North American species (Strobilops hubbardi) is depressed helicoid with a 

wide umbilicus; its shell surface has weakly developed, dense riblets. The aperture is dilated in 

all species and more or less thickened. As a family characteristic the strobilopsids possess a 

number of lamellae inside the last whorl. Number, shape and arrangement of these lamellae are 

54

species-specific. The family is currently distributed in East and Southeast Asia, the Philippines, 

the eastern half of North America, and Central America. 

The snails are found in moderately moist forests in leaf litter and under dead wood and 

bark. 

TAXON AUTHOR 

G- 


AL, AR, DE, FL, GA, HI, IA, IL, IN, KY, LA, MA, MD, MI, MO, 

MS, NC, NJ, NY, OH, OK, PA, SC, TN, TX, VA, WI, WV; Canada: 

Strobilops aeneus Pilsbry, 1926 G5 NS 

IL, IN, KY, MA, ME, MI, MN, MO, NJ, NY, OH, PA, RI, TN, WI; 

Strobilops affinis Pilsbry, 1893 G4G5 Canada: MB, ON, QC 

Strobilops hubbardi A.D. Brown, 1861 G3G4 AL, FL, GA, TX 

AL, AR, CT, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, 

MO, MS, NC, NE, NH, NJ, NY, OH, OK, PA, SC, SD, TN, TX, VA, 

Strobilops labyrinthicus (Say, 1817) G5 VT, WI, WV; Canada: NB, NS, ON, QC 

AL, AR, DE, FL, GA, KS, LA, MD, MS, NC, NJ, OK, PA, SC, TN, 

Strobilops texasianus Pilsbry and Ferriss, 1906 G5 TX, VA 

Succineidae 

John B. Burch, University of Michigan 

The Succineidae are a land snail family nearly worldwide in distribution, whose shells are 

characteristically thin and amber-colored, the latter aspect the source of their common name, 

“amber shells.” The succineids are distinct enough from other stylommatophoran snails to be 

placed in their own separate suborder, Heterurethra Pilsbry 1900. Pertinent among their unusual 

characteristics are their heterurethrus kidney and ureter, and their elasmognathous jaw. Another 

unusual aspect of the Succineidae is the range of their chromosome numbers---the widest range 

known for any land snail family. Further, the lowest chromosome number known for the 

Mollusca is found in the Succineidae. 

Four genera of Succineidae are recognized in North America (north of Mexico), 

Succinea, Oxyloma, Novosuccinea and Catinella. With some exceptions, the three genera can be 

recognized in many regions by characteristics of their shells. Succinea and Oxyloma have shells 

with large body whorls and short spires, whereas Catinella has a proportionately longer spire, 

approaching that of the aperture in length. Succinea generally has a more oval shell, while the 

shell of Oxyloma is more elongate and narrow. However, the most important characters for 

assigning North American species to genera are aspects of genital anatomy; these are more 

important than shell characters. 

55

The taxonomy of the North American Succineidae is in need of studies using modern 

techniques to determine taxon validity and relationships. 

TAXON AUTHOR 

G- 


Catinella aprica Hubricht, 1968 G2 AL, MS; Canada: ON 

Catinella baldwini (Ancey, 1889) GNR HI 

Catinella exile (Leonard, 1972) G2 DE, IL, IN, ME, MI, MN, WI; Canada: ON 

Catinella explanata (Gould, 1852) GNR HI 

Catinella gabbi (Tryon, 1866) G1G2 CA, ID, WA 

Catinella gelida (F.C. Baker, 1927) G1 IA, IL, IN, KY, MI, MO, MS, OH, SD, WI 

Catinella hubrichti Grimm, 1960 G3 DE, MD, NC, SC, VA 

AL, AR, DE, FL, GA, KY, LA, MD, MO, MS, NC, OK, 

Catinella oklahomarum (Webb, 1953) G5 

PA, SC, TN, VA, WV 

Catinella parallela Franzen, 1979 G3 IL, IN, WI 

Catinella paropsis Cooke, 1921 GNR HI 

Catinella pinicola Grimm, 1960 G5 MD 

Catinella protracta Franzen, 1983 G2Q MI 

Catinella pugilator Hubricht, 1961 G1G2 AL, GA, NC, SC 

Catinella rehderi (Pilsbry, 1948) G3 CA, ID, MT, WA 

Catinella rotundata (Gould, 1846) G1G3 HI 

Catinella rubida Pease, 1870 GNR HI 

Catinella stretchiana Bland, 1865 G3 CA, SD, UT, WY 

Catinella texana Hubricht, 1961 G1Q LA, TX 

Catinella tuberculata Cooke, 1921 GNR HI 

Catinella vagans (Pilsbry, 1900) G3Q KS, NJ 

AL, AR, CA, FL, GA, IA, ID, IN, KS, KY, LA, MA, MD, 

ME, MI, MN, MO, MS, NC, ND, NH, NJ, NM, NY, OH, 

OK, PA, SC, TX, UT, VT, WA, WI, WV, IL, TN, VA; 

Catinella vermeta (Say, 1829) G5 

Canada: AB, BC, NB, NF, NS, ON, QC 

Catinella waccamawensis Franzen, 1981 G1Q NC 

Catinella wandae (Webb, 1953) G2 AR, IA, KS, OK, WY 

Novisuccinea chittenangoensis (Pilsbry, 1908) G1 LT AR, IA, IL, MO, NY, VA 

AR, CA, CT, DE, GA, IA, IN, KS, KY, LA, MA, MD, ME, 

MI, MN, MO, MS, NC, ND, NE, NH, NJ, NY, OH, OR, PA, 

RI, SC, SD, VT, WI, WV, IL, TN, VA; Canada: NB, NF, 

Novisuccinea ovalis (Say, 1817) G5 

NS, ON, QC 

Oxyloma chasmodes Pilsbry, 1948 G1G3 CA 

Oxyloma decampi (Tryon, 1866) G5Q KY, NJ, WY 

Oxyloma deprimidum Franzen, 1973 G2 IL 

Oxyloma effusum (Pfeiffer, 1853) G3 DE, FL, MD, NC, NJ, NY, VA; Canada: ON 

Oxyloma groenlandicum (Moller, 1842) G3G4 NY; Canada: BC, NS, QC, NF, YT 

Oxyloma hawkinsi (Baird, 1863) G3G4 AK, ID, UT, WA; Canada: AB, BC, MB 

Oxyloma haydeni (W.G. Binney, 1858) G2G3 AZ, NE, UT, WA, WY; Canada: AB, MB, NT 

Oxyloma kanabense Pilsbry, 1948 G1 LE AZ, UT; Canada: AB 

Oxyloma missoula Hubricht, 1982 G2G4 

Oxyloma nuttallianum (I. Lea, 1841) G2G4 AK, CA, ID, MT, OR, UT, WA; Canada: BC 

Oxyloma peoriense (Wolf, 1894) G4G5 IL, IN, MI, NY, OH; Canada: NF, ON 

CA, CT, IA, IL, IN, KS, KY, MA, ME, MI, MN, ND, NE, 

NH, NJ, NM, NY, OH, PA, SD, TN, UT, VA, VT, WI; 

Oxyloma retusum (I. Lea, 1834) G5 

Canada: AB, NS, QC 

Oxyloma salleanum (Pfeiffer, 1849) G3 AR, IL, LA, MO, MS, TN, TX, WI 

Oxyloma sillimani (Bland, 1865) G2 CA, NV, UT 

Oxyloma subeffusum Pilsbry, 1948 G3 MD, NJ, PA, VA 

Oxyloma verrilli (Bland, 1865) G1G2 Canada: NF, QC 

Succinea bakeri Hubricht, 1963 GH IL, KY, MS, WI 

Succinea barberi (W. B. Marshall, 1926) G2 FL 

56

Succinea californica 

P. Fischer and Crosse, 

1878 G1G2 CA 

Succinea campestris Say, 1817 G4 FL, GA, LA, MD, ME, NC, NJ, SC, VA 

Succinea floridana Pilsbry, 1905 G2G3 FL 

AL, AR, FL, IA, IL, KS, KY, LA, MO, NC, NE, NM, OK, 

Succinea forsheyi I.Lea, 1864 G4 

TN, TX, WI 

Succinea greerii Tryon, 1866 G3 AL, LA, MS, OK, TN, TX 

AL, AR, AZ, FL, KS, KY, LA, MO, MS, NM, OK, TN, TX, 

Succinea grosvenori I. Lea, 1864 G5 

UT, WY; Canada: AB, MB, NT, ON, SK 

AL, AR, DE, FL, GA, IN, KS, MD, ME, MO, NC, ND, NE, 

Succinea indiana Pilsbry, 1905 G5 

NJ, NY, OK, SC, SD; Canada: ON 

Succinea luteola Gould, 1848 G4 AR, AZ, CA, FL, LA, MS, NM, SD, TX 

Succinea oregonensis I. Lea, 1841 G2G4 CA, ID, OR, UT, WA; Canada: AB, BC 

Succinea paralia Hubricht, 1983 G2 AL, FL, TX 

Succinea pennsylvanica Pilsbry, 1848 G1G2 NY, PA 

Succinea rusticana Gould, 1846 G2G3 AK, CA, ID, OR, UT, WA; Canada: BC 

Succinea solastra Hubricht, 1961 G2G3 TX 

Succinea strigata Pfeiffer, 1855 G4 AK; Canada: BC, SK, YT 

Succinea unicolor Tryon, 1866 G3G4 AL, AR, FL, GA, LA, MI, MS, NC, SC, TX 

Succinea urbana Hubricht, 1961 G2G3 AL, MS 

Succinea vaginacontorta C.B. Lee, 1951 G2G3Q KS, NM, TX 

Succinea wilsonii I. Lea, 1864 G4 DE, GA, MD, ME, NC, NJ, NY, SC, VA; Canada: NS, PE 

Thysanophoridae 


Hojeda inaguensis is found in moist leaf litter in hammocks in Florida. Its range also 

includes the Bahamas. The Thysanophora species are small, helicoid to flattened snails typically 

found under logs, dead palm fronds in woods, and rocks, but are also found in open scrublands. 

T. plagioptycha is usually found in wet places (Hubricht, 1985). Individuals of Microphysula 

cookei prefer wet montane coniferous forest, usually in wet places under vegetated rocks 

(Forsyth, 2004). M. ingersolii prefers subalpine meadows, spruce forests, and Trembling Aspen 

groves (Forsyth, 2004). 

TAXON AUTHOR 

G- 


Hojeda inaguensis (Weinland, 1880) GH FL 

Microphysula cookei (Pilsbry, 1922) G4 AK, AZ, WA; Canada: BC 

Microphysula ingersolli (Bland, 1875) G5 AZ, ID, MT, NM, OR, TX, UT, WA, WY; Canada: AB, BC 

Thysanophora hornii (Gabb, 1866) G5 AZ, NM, TX 

Thysanophora plagioptycha (Shuttleworth, 1854) G5 FL, TX 

57

Urocoptidae 


Urocoptidae is a large family related to the Bulimulidae and Achatinidae. The shells of snails in 

this family tend to be elongate with a tapering spire and an expanded apertural lip. The two 

genera (and species) found in Florida are similarly elongate, but Cochlodinella poeyana is 

smaller with wider whorls and typically has a decollate spire in contrast with Microceramus 

pontificus which is somewhat larger with a wider shell. The shape of the aperture is also very 

different. 

The majority of U.S. Urocoptids are in the genus Holospira. These are snails of the 

limestone terrain of the arid southwestern U.S. Holospira and related genera found in Mexico are 

very species-rich. Important characteristics for identification of Holospira species are the length 

and width of the shell, the number of whorls, the smoothness or ribbing of the shell, and the 

shape (angulate, ovate, or oblong) and folding of the aperture. For final identification the shell 

must be opened, usually by cutting or grinding away the penultimate whorl to look at the central 

axis of the shell. Lamellae are associated with the central axis of the shell, especially in the final 

whorl. These lamellae are variously developed and shaped in different species. 

Species of Holospira tend to live in isolated large colonies and be very restricted in 

geographic location, for example, found only in single mountain ranges. One exception to this is 

Metastoma roemeri which is found throughout central Texas to the Rio Grande Valley of New 

Mexico. It is most abundant around New Braunfels, Texas. Other Holospira species are found 

hanging attached to or under limestone ledges, a few species in West Texas are associated with 

Selaginella (Resurrection plant). They can also be found along canyon walls, under stones, dead 

Yucca stems, dead stems of Sotol and in limestone talus (Metcalf & Smartt, 1997). Specific 

localities and habitat discussions of each New Mexico Holospira species can be found in Land 

Snails of New Mexico (Metcalf & Smartt, 1997). 

TAXON AUTHOR 

G- 


Cochlodinella poeyana (d'Orbigny, 1841) G1G2 FL 

Coelostemma pyrgonasta F. G. Thompson, 1989 

Gilbertson and Worthington, 

G1 NM 

Holospira animasensis 

2003 G1G2 NM 

Holospira arizonensis Stearns, 1890 G2 AZ 

Holospira campestris Pilsbry and Ferriss, 1915 G3Q AZ 

Holospira chiricahuana Pilsbry, 1905 G2G3 AZ 

Holospira cionella Pilsbry, 1905 G3Q AZ 

Holospira cockerelli Dall, 1897 G1 NM 

Holospira crossei Dall, 1895 G2 NM 

58

Holospira danielsi Pilsbry and Ferriss, 1915 G3G4 AZ, TX 

Holospira ferrissi Pilsbry, 1905 G2 AZ 

Holospira goldfussi (Menke, 1847) G2G3 TX 

Holospira hamiltoni Dall, 1897 G1 TX 

Holospira mesolia Pilsbry, 1912 G1 TX 

Holospira metcalfi F.G. Thompson, 1974 G1 NM 

Holospira millestriata Pilsbry and Ferriss, 1915 G1G2Q AZ 

Holospira montivaga Pilsbry, 1946 G2 AZ, NM, TX 

Holospira oritis Pilsbry and Cheatum, 1951 G1 TX 

Holospira pasonis Dall, 1895 G1 TX 

Holospira pityis Pilsbry and Cheatum, 1951 G1 TX 

Holospira riograndensis Pilsbry, 1946 G1 TX 

Holospira sherbrookei Gilbertson, 1989 G1 AZ 

Holospira tantalus Bartsch, 1906 G1G2 AZ 

Holospira whetstonensis Pilsbry and Ferriss, 1923 G1G2 AZ, NM 

Holospira yucatanensis Bartsch, 1906 G1 TX 

Metastoma roemeri (Pfeiffer, 1848) G4 NM, TX 

Microceramus pontificus (Gould, 1848) G2G3 FL 

Microceramus texanus (Pilsbry, 1898) G2 TX 

Valloniidae 


The valloniids are minute snails (ca. 1.5-4 mm in diameter) of uniformly white, greyish 

or brown shell color. Their shells are mostly depressed-helicoid, but some species are more 

globular. Many species have an ornamentation of transverse ribs and sometimes spiral lines. 

The aperture is either simple (Planogyra, Zoogenetes) or expanded (Vallonia). In some Vallonia 

species there is a threshold-like callus within the aperture, however, all valloniids lack apertural 

teeth or lamellae. The umbilicus is wide to very wide, with the exception of the tall-helicoid 

Zoogenetes which has a merely perforate shell. 

The distribution of the family is holarctic. In North America they have a mainly northern 

distribution, with some Vallonia species extending further southward in the Western mountain 

ranges and into Northern Mexico. Some Vallonia species (V. costata, V. excentrica, V. pulchella) 

have been accidentally introduced in many areas outside their natural range. 

Vallonia species live usually in open habitats, such as meadows, lawns, and on exposed 

rock outcrops, occasionally extending into more open forest vegetation, but they avoid dense 

woods. Moisture requirements differ by species. The two Planogyra species live in leaf litter in 

moist forests and swampy areas. Zoogenetes is a snail of boreal forests of NE North America 

and of high-mountain forest habitats in the Rockies. 

59

TAXON AUTHOR 

G- 


IL, MA, ME, MI, MN, NH, NY, OH, VT, WI; Canada: NF, NS, ON, 

Planogyra asteriscus (E.S. Morse, 1857) G4 QC 

Planogyra clappi (Pilsbry, 1898) G3G4 BC, AK, CA, ID, OR, WA 

CA, CO, ID, MA, ME, MI, NM, OR, UT, WA, WY; Canada: AB, BC, 

Vallonia albula Sterki, 1893 G4Q MB, NF, ON, QC 

CA, DE, IA, IL, IN, KY, MA, MD, ME, MI, MN, NC, NE, NJ, NY, 

Vallonia costata (Muller, 1774) G5 OH, PA, SD, VA, WI, WV; Canada: ON, QC 

AK, AZ, CA, CO, IA, ID, MT, ND, NM, OR, SD, TX, UT, VT, WA, 

Vallonia cyclophorella Sterki, 1892 G5 WI, WY; Canada: AB, BC, MB, ON, SK 

CA, GA, IL, IN, KY, MA, MD, ME, MI, MO, NC, NJ, NY, OH, PA, 

Vallonia excentrica Sterki, 1893 G5 RI, TN, VA, VT, WI, WV; Canada: AB, BC, NF, NS, ON, QC 

AZ, CA, CO, IA, ID, IL, IN, KS, KY, MA, ME, MN, MO, MT, ND, 

NE, NM, NY, OK, RI, SD, TX, UT, WI, WY; Canada: AB, BC, MB, 

Vallonia gracilicosta Reinhardt, 1883 G5Q NF, NU, ON, QC 

IA, IL, KS, KY, MI, MN, MO, NE, NM, NY, OH, OK, SD, TN, TX, 

Vallonia parvula Sterki, 1893 G4 VA, WI; Canada: ON 

AL, AR, IA, IL, KY, MD, MN, MO, MS, NC, ND, NE, NJ, NM, SD, 

Vallonia perspectiva Sterki, 1893 G4G5 TN, TX, UT, VA, WI, WV; Canada: AB 

CA, CT, DE, IA, IL, IN, KY, MA, MD, ME, MI, MN, MO, NC, NE, 

NJ, NY, OH, PA, SD, TX, UT, VA, WA, WI, WV; Canada: AB, BC, 

Vallonia pulchella (Muller, 1774) G5 NB, NF, NS, ON, QC 

Vallonia terraenovae Gerber, 1996 G1 Canada: NF 

AK, MA, ME, MI, MN, NH, RI, WI; Canada: AB, BC, NB, NF, NS, 

Zoogenetes harpa (Say, 1824) G5 ON, QC 

Veronicellidae 


Only one species of Veronicellidae, Leidyula floridana, is native to the U.S. 

In contrast to the majority of slugs, L. floridana belongs to the order Systellommatophora 

instead of Stylommatophora. L. floridana is easily distinguished from most slugs by the 

presence of contractile instead of invaginable tentacles, the absence of a pneumostome, an anus 

at the end of the foot (due to detorsion of the gut), and a wide mantle covering the entire dorsum. 

This species is 50-70 mm long. 

L. floridana occupies any habitats with sufficient shelter, including woods, roadsides, and 

gardens. 

L. floridana is native to southern Florida and was introduced to Louisiana. Several nonnative 

Veronicellids overlap in range with L. floridana. 

Information summarized from Burch (1962), Hubricht (1985). 

60

TAXON AUTHOR 

G- 


Leidyula floridana (Leidy, 1851) G2G4 FL, LA 

Vitrinidae 


Vitrinids are snails or semislugs with the shell small to medium sized, very thin, glassy, 

transparent, helicoid to ear shaped, with few, rapidly expanding whorls. The body whorl is 

capacious, the aperture large, strongly oblique, and broader than high. The umbilicus is narrow 

or absent. In most cases the animal is unable to retract completely within its shell. One or more 

lobes of the mantle sometimes lap onto the shell. The radular teeth have long cusps; with two or 

more cusps present on marginal teeth. The foot is narrow, with a tripartite sole. There is no 

caudal mucus pit. The anterior cephalic tentacles are inconspicuous. 

Vitrinidae is a Holarctic clade, also extending to central Africa and some Atlantic islands. Only 

the semislug genus Vitrina occurs in North America. 

In Europe and western North America, vitrinids are found in a variety of moderately 

humid places, including woods and grassland. In the southern part of their range, such as the 

Sierra Nevada, California, and the southern Rocky Mountains, they occupy higher elevations and 

are found in habitats such as moist montane meadows, stream banks in moss and grasses, on 

bases of plants such as corn lily (Veratrum californicum), under pine bark, logs, and sticks on 

ground; in aspen thickets, and under willows at edges of meadows. 

Information summarized from Pilsbry (1946), Forcart (1955), Hubricht (1985). 

TAXON AUTHOR 

G- 


MA, MD, ME, MI, MN, NH, NY, PA, RI, VT, WI; Canada: NB, NF, 

Vitrina angelicae Beck, 1837 G5 NS, ON, QC 

Vitrina pellucida (Muller, 1774) G5 AK, CA, CO, ID, MT, NM, OR, SD, TX, UT; Canada: AB, BC, QC 

Vitrina pellucida alaskana Dall, 1905 G5T5 AK, AZ, CA, NM, OR, WA; Canada: BC 

61

Zonitidae 

Amy S. Van Devender, Boone, NC 

Taxonomic note: The genera traditionally included in the family Zonitidae have been reclassified 

(Bouchet & Rocroi, 2005). Following this new arrangement, the genera occuring in the 

U.S. and Canada can be assigned to several families as follows: Gastrodontidae: Gastrodonta, 

Striatura, Ventridens, Zonitoides; Oxychilidae: Glyphyalinia, Mesomphix, *Nesovitrea*, 

Ortizius, Oxychilus, Paravitrea, Pilsbryna, Vitrinizonites; Pristilomatidae: Hawaiia, 

Ogaridiscus, Pristiloma, Vitrea. 

The Zonitidae are considered to be a difficult group to identify. They are found in nearly 

every place in the world but probably originated and had their initial radiation of species in the 

Appalachain Mountains of the Eastern U. S. At present the family contains about 147 species 

assorted among 14 genera. It is interesting to note that Hubricht described or changed our 

understanding of over 1/5 of the species in this family. 

The family consists of moderate to minute snails which have shells with 3 to 9 whorls 

that are usually shiny in a dazzling array of subtle colors ranging from yellow to green, copper to 

pink. Their shells tend to be wider than high and may have a variety of incised lines or patterns 

of micro dots as ornamentation. Younger shells especially may have teeth in various patterns. 

The lip of the shell is sharp and thin; and though it may fold back on itself at the umbilical 

insertion, it is never reflected. Like its allies in the Aulacopoda (the slugs and discids) these 

animals have noticeable pedal grooves lying well above the sole of the foot. 

Most of these species seem to love leaf litter of mature woods and the deep coves of the 

mountains often harbor a dozen species. Little is known about how they divvy up the 

microhabitats but prying apart wet layers of compacted leaves usually turns up live animals. 

Some species especially in the Paravitrea and Mesomphix are found in talus. Some like Hawaiia 

minuscula and Zonitoides arboreus (even Ventridens demissus) act like tramp snails turning up 

in all kinds of strange places. 

TAXON AUTHOR 

G- 


Gastrodonta fonticula Wurtz, 1948 G3G4 KY, VA, WV, TN 

Gastrodonta interna (Say, 1822) G5 AL, GA, IN, KY, MS, NC, OH, PA, SC, TN, VA, WV 

Glyphyalinia carolinensis (Cockerell, 1890) G4 AL, KY, NC, TN, VA 

Glyphyalinia clingmani (Dall, 1890) G1 NC 

Glyphyalinia cryptomphala (G.H. Clapp, 1915) G5 AL, GA, KY, MO, TN, VA 

Glyphyalinia cumberlandiana (G.H. Clapp, 1919) G4 AL, KY, MD, NC, TN, VA, WV 

Glyphyalinia floridana (Morrison, 1937) GH FL 

62

Glyphyalinia indentata (Say, 1823) G5 

AL, AR, FL, GA, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MO, 

MS, NC, NJ, NM, NY, OH, OK, PA, SC, TN, TX, UT, VA, VT, WI, 

WV; Canada: NS, ON, QC 

Glyphyalinia junaluskana (Clench and Banks, 1932) G2 GA, NC, TN 

Glyphyalinia latebricola Hubricht, 1968 G1G2 AL, IN 

Glyphyalinia lewisiana (G.H. Clapp, 1908) G4 AL, AR, IN, KY, LA, MO, MS, NC, TN, VA, WV 

AL, AR, FL, GA, LA, MD, MS, NC, NJ, OH, OK, SC, TN, VA; 

Glyphyalinia luticola Hubricht, 1966 G4 Canada: ON 

Glyphyalinia ocoae Hubricht, 1978 G1 NC, TN 

Glyphyalinia pecki Hubricht, 1966 G1G2 AL 

Glyphyalinia pentadelphia (Pilsbry, 1900) G2G3 GA, NC, TN 

Glyphyalinia picea Hubricht, 1976 G3 DE, MD, VA, WV 

Glyphyalinia praecox (H.B. Baker, 1930) G4 AL, GA, KY, LA, MS, NC, SC, TN, VA, WV 

Glyphyalinia raderi (Dall, 1898) G2 KY, MD, PA, VA, WV 

CT, DE, GA, IL, KY, MA, MD, ME, MI, NC, NH, NJ, NY, OH, PA, 

Glyphyalinia rhoadsi (Pilsbry, 1899) G5 SC, TN, VA, VT, WI, WV; Canada: ON 

Glyphyalinia rimula Hubricht, 1968 G3 IN, KY, TN, WV 

Glyphyalinia roemeri (Pilsbry and Ferriss, 1906) G3 LA, TX 

Glyphyalinia sculptilis (Bland, 1858) G4 AL, GA, KY, LA, MS, NC, SC, TN, VA 

AL, AR, FL, GA, IL, IN, KY, MD, MI, MO, MS, NC, NJ, OK, SC, 

Glyphyalinia solida (H.B. Baker, 1930) G5 TN, VA, WV 

Glyphyalinia specus Hubricht, 1965 G4 AL, GA, KY, TN, WV 

Glyphyalinia umbilicata (Cockerell, 1893) G5 FL, GA, LA, NC, SC, TX, UT 

Glyphyalinia virginica (Morrison, 1937) G3 KY, VA 

AL, AR, CT, DE, FL, GA, IL, IN, KY, LA, MA, MD, MI, MO, MS, 

Glyphyalinia wheatleyi (Bland, 1883) G5 NC, NJ, NY, OH, OK, PA, TN, VA, WI, WV; Canada: ON 

Godwinia caperata (Gould, 1846) GNR HI 

Godwinia haupuensis Cooke, 1921 GNR HI 

Godwinia newcombi (Reeve, 1854) GNR HI 

AR, FL, IL, IN, KY, LA, MD, MI, MO, MS, NC, NJ, NY, OH, OK, 

Hawaiia alachuana (Dall, 1885) G4G5Q PA, SC, TN, TX, VA, WV; Canada: ON 

AL, AR, CA, DE, FL, GA, HI, IA, ID, IL, IN, KS, KY, LA, MA, 

MD, ME, MI, MN, MO, MS, MT, NC, NE, NJ, NM, NY, OH, OK, 

PA, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV; Canada: AB, NF, 

Hawaiia minuscula (A. Binney, 1841) 

(Cockerell and Pilsbry, 

G5 NS, ON, QC 

Hawaiia neomexicana 

1900) G2 UT, NM 

Mesomphix andrewsae (Pilsbry, 1895) G3N4 NC, TN 

Mesomphix anurus Hubricht, 1962 G4N5 AL, GA, KY, MS, NC, TN 

Mesomphix capnodes (W.G. Binney, 1857) G5 AL, AR, GA, KS, KY, LA, MO, MS, NC, OK, TN, VA, WV 

CT, DE, IL, IN, KS, KY, MD, MI, NC, NJ, NY, OH, OK, PA, TN, 

Mesomphix cupreus (Rafinesque, 1831) G5 VA, VT, WV; Canada: ON 

Mesomphix friabilis (W.G Binney, 1857) G5 AL, AR, IL, IN, KY, LA, MO, MS, OH, OK, TN, TX 

Mesomphix globosus (MacMillan, 1940) G5 AL, AR, FL, GA, IL, IN, KY, LA, MS, SC, TN, TX 

IN, KY, MA, MD, NJ, NY, OH, PA, TN, VA, VT, WV; Canada: ON, 

Mesomphix inornatus (Say, 1821) G5 QC 

Mesomphix latior (Pilsbry, 1900) G3G4 AL, GA, KY, NC, TN 

Mesomphix perfragilis (Wetherby, 1894) G1 TN 

Mesomphix perlaevis (Pilsbry, 1900) G4G5 GA, IL, KY, MD, NC, OH, PA, TN, VA, WV 

Mesomphix pilsbryi (G.H. Clapp, 1904) G4 AL, FL, GA, MS, NC, SC 

Mesomphix rugeli (W.G. Binney, 1879) G4 GA, KY, NC, TN, VA 

Mesomphix subplanus (A. Binney, 1842) G3G4 GA, NC, TN, VA 

Mesomphix vulgatus H.B. Baker, 1933 G4 IN, KY, LA, OH, TN, WI 

CA, CO, IA, ID, IN, KY, MA, ME, MI, MN, MT, ND, NE, NH, NY, 

OH, OR, PA, SD, UT, WA, WI, WY; Canada: AB, BC, MB, NS, ON, 

Nesovitrea binneyana (E.S. Morse, 1864) G5 QC 

Nesovitrea dalliana (Pilsbry and Simpson, 1889) G3G4 FL, GA, SC 

IL, AK, AR, AZ, CT, DE, IA, ID, IN, KS, KY, LA, MA, MD, ME, 

MI, MN, MO, NE, NH, NJ, NM, NY, OH, OK, PA, SD, TN, TX, UT, 

Nesovitrea electrina (Gould, 1841) G5 VA, VT, WA, WI, WV; Canada: AB, BC, LB, NB, NF, NS, ON, QC 

63

Nesovitrea hawaiiensis (Ancey, 1904) GNR HI 

Nesovitrea molokaiensis (Sykes, 1897) GNR HI 

Nesovitrea pauxilla (Gould, 1852) GNR HI 

Nesovitrea suzannae Pratt, 1978 G1 TX 

Ogaridiscus subrupicola (Dall, 1877) G1 ID, OR, UT 

Paravitrea alethia Hubricht, 1978 G1 TN, TX 

Paravitrea amicalola Hubricht, 1976 G1 GA 

Paravitrea andrewsae (W.G Binney, 1879) G2 KY, NC, TN 

Paravitrea aulacogyra (Pilsbry and Ferriss, 1906) GHQ AR 

Paravitrea bellona Hubricht, 1978 G1 NC, WV 

Paravitrea bidens Hubricht, 1963 G1 AL 

Paravitrea blarina Hubricht, 1963 G3 KY, TN, VA 

Paravitrea calcicola H.B. Baker, 1931 G1 TN, VA 

Paravitrea capsella (Gould, 1851) G4 AL, GA, IL, IN, KS, KY, NC, OH, PA, TN, VA, WI, WV 

Paravitrea ceres Hubricht, 1978 G1 WV 

Paravitrea clappi (Pilsbry, 1898) G2G3 NC, TN 

Paravitrea conecuhensis (G.H. Clapp, 1917) G3 AL, FL, LA, MS, TX 

Paravitrea dentilla Hubricht, 1978 G1 VA 

Paravitrea diana Hubricht, 1983 G1 GA 

Paravitrea grimmi Hubricht, 1968 G1G3Q VA 

Paravitrea hera Hubricht, 1983 G1 VA 

Paravitrea lacteodens (Pilsbry, 1903) G1 NC, PA 

Paravitrea lamellidens (Pilsbry, 1898) G2 ME, NC, TN 

Paravitrea lapilla Hubricht, 1965 G2 KY, TN 

Paravitrea metallacta Hubricht, 1963 G3 TN 

Paravitrea mira Hubricht, 1975 G2 KY, VA 

AL, AR, CT, IN, KY, LA, MA, MD, ME, MI, MS, NC, NH, NY, OH, 

Paravitrea multidentata (A. Binney, 1840) G5 PA, TN, VA, VT, WI, WV; Canada: NB, NS, ON 

Paravitrea petrophila (Bland, 1883) G4 AL, AR, KY, OK, TN, WV 

Paravitrea pilsbryana (G.H. Clapp, 1919) G2 AL, TN 

Paravitrea placentula (Shuttleworth, 1852) G3 GA, KY, NC, TN, VA 

Paravitrea pontis H.B. Baker, 1931 G3 KY, MD, VA, WV 

Paravitrea reesei Morrison, 1937 G3 KY, NC, TN, VA, WV 

Paravitrea septadens Hubricht, 1978 G1 KY, VA 

Paravitrea seradens Hubricht, 1972 G3 KY, VA, WV 

Paravitrea significans (Bland, 1866) G3 AR, IL, KS, LA, MO, MS, OK, TN, WI 

Paravitrea simpsoni (Pilsbry, 1889) G4 AR, KS, MO, OK, TX 

Paravitrea subtilis Hubricht, 1978 G2 KY, TN, VA 

Paravitrea tantilla Hubricht, 1963 G3 AL, KY, TN 

Paravitrea ternaria Hubricht, 1978 G1G2 NC, TN 

Paravitrea tiara Hubricht, 1978 G1G2 AL 

Paravitrea toma Hubricht, 1975 G1 AL 

Paravitrea tridens Pilsbry, 1946 G2 TN, VA 

Paravitrea umbilicaris (Ancey, 1887) G2 AL, GA, NC, TN 

Paravitrea variabilis H.B. Baker, 1929 G2G3 AL, TN 

Paravitrea varidens Hubricht, 1978 G1G2 NC, TN 

Pilsbryna aurea H.B. Baker, 1929 G1 TN 

Pilsbryna castanea H.B. Baker, 1931 G2 TN 

Pilsbryna nodopalma Slapcinsky and Coles, 2004 G1G2 NC, TN 

Pilsbryna quadrilamellata Slapcinsky and Coles, 2004 G1 TN 

Pilsbryna vanattai (Walker and Pilsbry, 1902) G2G3 NC, TN 

Pristiloma arcticum (Lehnert, 1884) G3G4 AK, ID, OR, WA; Canada: BC, YT 

Pristiloma cavator Roth, 1998 G1G2 CA 

Pristiloma chersinella (Dall, 1886) G3G4 CA, ID, MT, NV, OR, WA; Canada: BC 

Pristiloma gabrielinum (S.S. Berry, 1924) G1G2 CA 

64

Pristiloma idahoense Pilsbry, 1902 G2G3 ID, OR, WA 

Pristiloma johnsoni (Dall, 1895) G2G3 OR, WA; Canada: BC 

Pristiloma juniperum A.G. Smith, 1957 G1G2 CA 

Pristiloma lansingi (Bland, 1875) G5 CA, OR, WA; Canada: BC 

Pristiloma nicholsoni H.B. Baker, 1930 G1G2 CA 

Pristiloma orotis (S.S. Berry, 1930) G1G2 CA 

Pristiloma pilsbryi Vanatta, 1899 G1 OR, WA 

Pristiloma shepardae (Hemphill, 1892) G1 CA 

Pristiloma stearnsi (Bland, 1875) G4G5 AK, CA, OR, WA; Canada: BC 

Pristiloma wascoense (Hemphill, 1911) G2 ID, MT, OR, WA 

Striatura exigua (Stimpson, 1850) G5 

CT, IA, KY, LA, MA, MD, ME, MI, MN, MS, NC, NH, NJ, NY, OH, 

OR, PA, TN, VA, VT, WI, WV; Canada: NB, NF, NS, ON, QC 

KY, MA, MD, ME, MI, MN, NC, NH, NY, OH, PA, TN, VA, VT, 

Striatura ferrea E.S. Morse, 1864 G5 WI; Canada: AB, NS, ON, QC 

AL, AR, DE, FL, GA, IL, IN, KS, KY, LA, MD, MI, MO, MS, NC, 

Striatura meridionalis (Pilsbry and Ferriss, 1906) G5 NJ, NM, OH, OK, PA, SC, TN, TX, VA, WV 

CT, DE, IA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, NE, NJ, 

Striatura milium (E.S. Morse, 1859) G5 NY, OH, OK, PA, VA, VT, WI, WV; Canada: NF, NS, ON, QC 

Striatura pugetensis (Dall, 1895) G5 AK, CA, HI, ID, MT, OR, WA; Canada: BC 

Ventridens acerra (J. Lewis, 1870) G4 AL, GA, KY, NC, TN, VA, WV 

Ventridens arcellus Hubricht, 1976 G4 GA, MD, NC, TN, VA, WV 

Ventridens brittsi (Pilsbry, 1892) G3 AR, MO, OK 

Ventridens cerinoideus (Anthony, 1865) G4 AL, DE, FL, GA, MD, NC, SC, VA 

Ventridens coelaxis (Pilsbry, 1899) G3 KY, NC, TN, VA 

Ventridens collisella (Pilsbry, 1896) G4 AL, GA, KY, NC, TN, VA, WV 

Ventridens decussatus (Walker and Pilsbry, 1902) G3 GA, NC, SC, TN, VA 

AL, AR, FL, GA, IL, IN, KY, LA, MD, MO, MS, NC, OK, PA, TN, 

Ventridens demissus (A. Binney, 1843) G5 TX, VA, WV; Canada: ON 

Ventridens eutropis Pilsbry, 1946 G2G3 TN 

Ventridens gularis (Say, 1822) G5 AL, GA, IN, KY, LA, MS, NC, OH, PA, SC, TN, VA, WV 

AL, AR, DE, FL, GA, IN, KY, LA, MD, MI, MS, NC, NY, OH, PA, 

Ventridens intertextus (A. Binney, 1843) G5 SC, TN, TX, VA, WV; Canada: ON 

Ventridens lasmodon (Phillips, 1841) G4 AL, KY, NC, TN, VA, WV 

Ventridens lawae (W.G. Binney, 1892) G4 AL, GA, KY, NC, TN, VA 

AL, AR, DE, IL, IN, KS, KY, LA, MD, MI, MO, MS, NC, NJ, NY, 

Ventridens ligera (Say, 1821) G5 OH, OK, PA, TN, VA, WV; Canada: ON 

Ventridens monodon Hubricht, 1964 G2 AL 

Ventridens percallosus (Pilsbry, 1898) G3 KY, TN 

Ventridens pilsbryi Hubricht, 1964 G4 AL, GA, KY, LA, MS, NC, SC, TN, VA 

Ventridens suppressus (Say, 1829) G5 DE, KY, MD, MI, NC, NJ, NY, OH, PA, TN, VA, WV; Canada: ON 

Ventridens theloides (Walker and Pilsbry, 1902) G4G5 AL, GA, KY, NC, SC, TN, VA, WV 

Ventridens virginicus (Vanatta, 1936) G4 IL, MD, NY, PA, VA, WV 

Ventridens volusiae (Pilsbry, 1900) G2G3 FL 

Vitrinizonites latissimus (J. Lewis, 1875) G4 AL, GA, KY, NC, TN, VA 

AL, AR, AZ, CA, CO, CT, DE, FL, GA, HI, IA, ID, IL, IN, KS, KY, 

LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, 

NM, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, 

Zonitoides arboreus (Say, 1816) G5 WA, WI, WV, WY; Canada: AB, BC, NB, NF, NS, ON, QC 

Zonitoides elliotti (Redfield, 1856) G4 AL, GA, KY, NC, SC, TN, VA, WV 

Zonitoides kirbyi R.W. Fullington, 1974 G2 IN, MO, OK, TX 

Zonitoides lateumbilicatus (Pilsbry, 1895) G3G4 AL, KY, MS, TN 

Zonitoides limatulus (A. Binney, 1840) G4G5 IA, IL, IN, KY, ME, MI, MN, MO, NY, OH, VT, WI 

AK, DE, IA, IL, IN, KY, LA, MA, MD, ME, MI, MN, NE, NJ, NY, 

OH, OK, OR, PA, SD, TN, UT, VT, WA, WI, WV; Canada: AB, BC, 

Zonitoides nitidus (Muller, 1774) G5 NF, NS, ON, QC 

Zonitoides patuloides (Pilsbry, 1895) G3 GA, NC, SC, TN 

65

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Sciences of Philadelphia. 52:561-567. 

Pilsbry, H.A. 1939-1948. Land Mollusca of North America (north of Mexico). Monographs of the Academy of 

Natural Sciences of Philadelphia, 3(parts 1-4): 2215 pp. 

Pilsbry, H. A. & J. H. Ferriss. 1906. Mollusca of the southwestern states, II. Proceedings of the Academy of Natural 

Sciences of Philadelphia, 58:123-175. 

68

Ponder, W. F. and D. R. Lindberg. 1997. Towards a phylogeny of gastropod mollusks: an analysis using 

morphological characters. Zoological Journal of the Linnean Society. 119:83-265. 

Richardson, L. 1980. Helicidae: catalog of species. Tryonia, 3(1): 1-697. 

Roth, B. 1987. Identities of two Californian land mollusks described by Wesley Newcomb. Malacological Review, 

20: 129-132. 

Roth, B. 1996. Homoplastic loss of dart apparatus, phylogeny of the genera, and a phylogenetic taxonomy of the 

Helminthoglyptidae (Gastropoda: Pulmonata). The Veliger, 39(1): 18-42. 

Regoli, F., S. Gorbi, D. Fattorini, S. Tedesco, A. Notti, N. Machella, R. Bocchetti, M. Benedetti, F. Piva, 2006. Use 

of the land snail Helix aspersa as sentinel organism for monitoring ecotoxicologic effects of urban pollution: an 

integrated approach. Environmental Health Perspectives, 114(1): 63-69. 

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Natural History Contributions in Science, 3: 1-81. 

Roth, B. 1997. [Review of] Guamampa n.g. (Gastropoda, Pulmonata), a bradybaenid land snail with monadeniid 

characters, by A. A. Schileyko. The Veliger, 40(4): 368-370. 

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Pp. 29-51. In: J. Taylor Ed. Origin and evolutionary radiation of the Mollusca. The Malacological Society of 

London, London. 

Slapcinsky, J. and B. Coles. 2004. Revision of the genus Pilsbryna (Gastropoda: Pulmonata: Gastrodontidae) and 

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21-46. 

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and Vestia turgida (Rossmässler, 1836) (Gastropoda: Clausiliidae). Journal of Molluscan Studies, 71: 105-112. 

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the affinities of the genus (Gastropoda: Pulmonata). Journal of Molluscan Studies. 58-135-155. 

Turgeon, D.D., J.F. Quinn, Jr., A.E. Bogan, E.V. Coan, F.G. Hochberg, W.G. Lyons, P.M. Mikkelsen, R.J. Neves, 

C.F.E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.G. Thompson, M. Vecchione, and J.D. Williams. 1998. 

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Edition. American Fisheries Society Special Publication 26, Bethesda, Maryland: 526 pp. 

Wade, C. M., P. B. Mordan, and F. Naggs. 2006. Evolutionary relationships among the Pulmonate land snails and 

slugs (Pulmonata: Stylommatophora). Biological Journal of the Linnean Society 87:593-610. 

Wade, C. M., P. B. Mordan, and B. Clarke. 2001. A phylogeny of the landsnails (Gastropoda: Pulmonata). 

Proceedings of the Royal Society of London Series B 268:413-422. 

Waggoner, J., S.A. Clark, K.E. Perez, and C. Lydeard. 2006. A survey of terrestrial gastropods of the Sipsey 

Wilderness (Bankead National Forest), Alabama. Southheastern Naturalist, 5(1): 57-68. 

Walker, Bryant, 1928. The terrestrial shell-bearing Mollusca of Alabama. Michigan University Museum of 

Zoology, Miscellaneous Publication No. 18, 180 pp. 

Walsh, G.E. and B.F. Coles. 2006. Daedalochila lithica and Daedalochila dorfeuilliana (Gastropoda: Polygyridae) 

in Arkansas, USA: morphology, distribution, and habitat. The Nautilus, 120(4): 131-138. 

Guide suggestions: 

Baker, F. C., 1939. Fieldbook of Illinois Land Snails. Illinois Natural History Survey, Urbana, Illinois. 166 pp. 

Burch, J. B., 1962. How to Know the Eastern Land Snails. Wm. C. Brown Company Publishers, Dubuque, Iowa. 

214 pp. 

Burch, J. B. & T. A. Pearce, 1990. Chapter 9: Terrestrial gastropods. Pp 201-309 in D. L. Dindal (ed.) Soil Biology 

Guide. John Wiley and Sons, New York. 1349 pp. 

Hubricht, L., 1985. The distributions of the native land mollusks of the eastern United States. Fieldiana: Zoology, 

N.S. 24: i-viii, 1-191. 

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Pearce, T.A. and A. Örstan, 2006. Terrestrial gastropoda. Pp. 261-285 in Sturm, C.F., T.A. Pearce and A. Valdés 

(Eds.). The Mollusks: a Guide to their Study, Collection, and Preservation. Universal Publishers, Inc., Boca 

Raton, FL. 445 pp. 

Pilsbry, H.A, 1939-1948. Land Mollusca of North America (North of Mexico). Monographs of the Academy of 

Natural Sciences of Philadelphia, (parts 1-4): 2215 pp. 

Appendix 

Species Synonymy for Burch’s (1962) “How to Know the Eastern Land Snails 

Compiled by James (Jay) R. Cordeiro, NatureServe 

Listed with each name is the authority (usually the most recent) on which the synonymic placement is based. 

Names alphabetical by species and not italicized for readability. 

albolabris (Say, 1816), Triodopsis = Neohelix albolabris (Say, 1816) (fide Emberton, 1988) 

aldrichianus (G.H. Clapp, 1907), Clappiella = Helicodiscus aldrichianus (G.H. Clapp, 1907) (fide Hubricht, 1985) 

alticola (Ingersoll, 1875), Columella = Columella columella alticola (Ingersoll, 1875) (fide Turgeon et al., 1998) 

appressus (Say, 1821), Mesodon = Patera appressa (Say, 1821) (fide Emberton, 1991) 

approxima (Walker and Pilsbry, 1902), Retinella = Glyphyalinia clingmani (Dall, 1890) (fide Hubricht, 1985) 

archeri Pilsbry, 1940, Mesodon = Fumonelix archeri (Pilsbry, 1940) (fide Emberton, 1991) 

auriculata Say, 1818, Polygyra = Daedalochila auriculata (Say, 1818) (fide Emberton, 1994) 

auriformis (Bland, 1859), Polygyra = Daedalochila auriformis (Bland, 1859) (fide Emberton, 1994) 

avara Say, 1818, Polygyra = Daedalochila avara (Say, 1818) (fide Emberton, 1994) 

avara (Say, 1824), Catinella = nomen dubium; most occurrences attributed to Catinella vermeta (Say, 1829) (fide 

Hoagland and Davis, 1987) 

bicolor (Hutton, 1834), Gulella = Huttonella bicolor (Hutton, 1834) (there is no consensus in assigning this species 

to Gulella nor in the status of the subgenus Huttonella, of which it is the type species (Naggs, 1989), however 

Turgeon et al. (1998) list it in Huttonella) 

binneyana (E.S. Morse, 1864), Retinella = Nesovitrea binneyana (E.S. Morse, 1864) (fide Hubricht, 1985) 

binneyanus (Pilsbry, 1899), Mesodon = Patera binneyana (Pilsbry, 1899) (fide Emberton, 1991) 

burringtoni (Pilsbry, 1928) Retinella = Glyphyalinia wheatleyi (Bland, 1883) (fide Hubricht, 1985) 

campi G.H. Clapp and Ferriss, 1919, Praticolella = Praticolella taeniata Pilsbry, 1940 (fide Hubricht, 1985) 

clenchi (Rehder, 1932), Mesodon = Patera clenchi (Rehder, 1932) (fide Emberton, 1991) 

carolinensis (Cockerell, 1890), Retinella = Glyphyalinia carolinensis (Cockerell, 1890) (fide Hubricht, 1985) 

caroliniensis (I. Lea, 1834), Triodopsis = Xolotrema carolinense (I. Lea, 1834) (fide Emberton, 1988) 

catskillensis (Pilsbry, 1896), Discus cronkhitei = Discus catskillensis (Pilsbry, 1896) (fide Hubricht, 1985) 

chilhoweensis (J. Lewis, 1870), Mesodon = Appalachina chilhoweensis (J. Lewis, 1870) (fide Emberton, 1994) 

chisosensis Pilsbry, 1936, Polygyra = Daedalochila chisosensis (Pilsbry, 1936) (fide Emberton, 1994) 

christyi (Bland, 1860), Mesodon = Fumonelix christyi (Bland, 1860) (fide Emberton, 1991) 

circumstriata (Taylor, 1908), Retinella = Glyphyalinia wheatleyi (Bland, 1883) (fide Hubricht, 1985) 

clarki (Vanatta, 1924), Anguispira = Anguispira alternata (Say, 1816) (fide Hubricht, 1985) 

clarki (I. Lea, 1858), Mesodon = Patera clarki (I. Lea, 1858) (fide Emberton, 1991) 

clavulinus (Potiez and Michaud, 1838), Lamellaxis = Allopeas clavulinum (Potiez and Michaud, 1838) (fide 

Turgeon et al., 1998, who follow Naggs, 1994, and Cowie et al., 1995) 

clingmani (Dall, 1890), Retinella = Glyphyalinia clingmani (Dall, 1890) (fide Hubricht, 1985) 

cronkhitei (Newcomb, 1865), Discus = Discus whitneyi (Newcomb, 1864) (fide Roth, 1987) 

cumberlandiana (G.H. Clapp, 1919), Retinella = Glyphyalinia cumberlandiana (G.H. Clapp, 1919) (fide Hubricht, 

1985) 

cryptomphala (G.H. Clapp, 1915), Retinella = Glyphyalinia cryptomphala (G.H. Clapp, 1915) (fide Hubricht, 1985) 

dalliana (Pilsbry and Simpson, 1889), Retinella = Nesovitrea dalliana (Pilsbry and Simpson, 1889) (fide Hubricht, 

1985) 

dealbatus (Say, 1830), Bulimulus = Rabdotus dealbatus (Say, 1830) (fide Hubricht, 1985) 

denotata (Ferussac, 1821), Triodopsis = Xolotrema denotatum (Ferussac, 1821) (fide Emberton, 1988) 

dentifera (A. Binney, 1837), Triodopsis = Neohelix dentifera (A. Binney, 1837) (fide Emberton, 1988) 

70

divesta (Gould, 1848), Triodopsis = Neohelix divesta (Gould, 1848) (fide Emberton, 1988) 

dorfeuilliana I. Lea, 1838, Polygyra = Daedalochila dorfeuilliana (I. Lea, 1838) (fide Walsh and Coles, 2006) 

downieanus (Bland, 1861), Mesodon = Inflectarius downieanus (Bland, 1861) (fide Emberton, 1991) 

electrina (Gould, 1841), Retinella = Nesovitrea electrina (Gould, 1841) (fide Hubricht, 1985) 

elegans (Gmelin, 1791), Helicella = Trochoidea elegans (Gmelin, 1791) (fide Gude and Woodward, 1921) 

elliotti (Redfield, 1856), Ventridens = Zonitoides elliotti (Redfield, 1856) (fide Hubricht, 1985) 

fatigiata Say, 1829, Polygyra = Daedalochila fatiagata (Say, 1829) (fide Coles and Walsh, 2006) 

ferrissi (Pilsbry, 1897), Mesodon = Inflectarius ferrissi (Pilsbry, 1897) (fide Emberton, 1991) 

floridana (Leidy, 1851), Veronicella = Leidyula floridana (Leidy, 1851) (fide Thome, 1975) 

floridana (Morrison, 1937), Retinella = Glyphyalinia floridana (Morrison, 1937) (fide Hubricht, 1985) 

floridana Pilsbry, 1907, Varicella gracillima = Melaniella gracillima floridana (Pilsbry, 1907) (fide Turgeon et al., 

1998, who place this taxon in the family Oleacinidae fide Burch and Pearce, 1990, but provide no explanation 

for their inclusion in the genus Melaniella Pfeiffer, 1859, instead of Varicella Pfeiffer, 1855; note Pilsbry, 1946, 

lists Melaniella as a subgenus of Varicella) 

floridanus (Pilsbry, 1898), Microceramus = Microceramus pontificus (Gould, 1848) (fide Hubricht, 1978) 

floridanum G.H. Clapp, 1918, Carychium = Carychium mexicanum Pilsbry, 1891 (fide Hubricht, 1985) 

fosteri (F.C. Baker, 1921), Triodopsis = Xolotrema fosteri (F.C. Baker, 1921) (fide Emberton, 1988) 

fraternum (Say, 1824), Stenotrema = Euchemotrema fraternum (Say, 1824) (fide Turgeon et al., 1998 who elevated 

the genus Euchemotrema from subgenus without providing justification, however fide Emberton (1994) for 

monophyletic support of genus Euchemotrema) 

gracilis (Hutton, 1834), Lamellaxis = Allopeas gracilis (Hutton, 1834) (fide Turgeon et al., 1998 who follow Naggs, 

1994, and Cowie et al., 1995) 

gularis theloides (Walker and Pilsbry, 1902), Ventridens = Ventridens theloides (Walker and Pilsbry, 1902) (fide 

Hubricht, 1985) 

hippocrepis (Pfeiffer, 1848), Polygyra = Daedalochila hippocrepis (Pfeiffer, 1848) (fide Emberton, 1994) 

hispida (Linnaeus, 1758), Hygromia = Trochulus hispida (Linnaeus, 1758) (fide Dundee, 1974 for synonymy with 

Trichia striolata (Pfeiffer, 1828) and ICZN, 2004 for retention of Trochulus over Trichia) 

hubrichti Pilsbry, 1940, Stenotrema = Euchemotrema hubrichti (Pilsbry, 1940) (fide Turgeon et al., 1998 who 

elevated the genus Euchemotrema from subgenus without providing justification, however fide Emberton 

(1994) for monophyletic support of genus Euchemotrema) 

indentata (Say, 1823), Retinella = Glyphyalinia indentata (Say, 1823) (fide Hubricht, 1985) 

inflectus (Say, 1821), Mesodon = Inflectarius inflectus (Say, 1821) (fide Emberton, 1991) 

indianorum (Pilsbry, 1899), Mesodon = Patera indianorum (Pilsbry, 1899) (fide Emberton, 1991) 

leai (A. Binney, 1821), Stenotrema = Euchemotrema leai (A. Binney, 1821) (fide Turgeon et al., 1998 who elevated 

the genus Euchemotrema from subgenus without providing justification, however fide Emberton (1994) for 

monophyletic support of genus Euchemotrema) 

leporina (Gould, 1848), Polygyra = Daedalochila leporina (Gould, 1848) (fide Emberton, 1994) 

jacksoni (Bland, 1866), Polygyra = Daedalochila jacksoni (Bland, 1866) (fide Coles and Walsh, 2006) 

jonesianus (Archer, 1938), Mesodon = Fumonelix jonesiana (Archer, 1938) (fide Emberton, 1991) 

kiowaensis (Simpson, 1888), Mesodon = Patera kiowaensis (Simpson, 1888) (fide Emberton, 1991) 

lewisiana (G.H. Clapp, 1908), Retinella = Glyphyalinia lewisiana (G.H. Clapp, 1908) (fide Hubricht, 1985) 

latispira Pilsbry, 1896, Polygyra = Triodopsis vultuosa (Gould, 1848) (fide Hubricht, 1985) 

limpida Gould, 1850, Vitrina = Vitrina angelicae limpida Gould, 1850 (fide Forcart, 1955) 

lubrica (Müller, 1774), Cionella = Cochlicopa lubrica (Müller, 1774) (fide Roth, 2003) 

magazinensis (Pilsbry and Ferriss, 1907), Mesodon = Inflectarius magazinensis (Pilsbry and Ferriss, 1907) (fide 

Emberton, 1991) 

mariae (Albers, 1850), Bulimulus alternatus = Rabdotus alternatus (Say, 1830) (fide Hubricht, 1985) 

mauritianum (Pfeiffer, 1852), Lamellaxis = Allopeas mauritianum (Pfeiffer, 1852) (fide Turgeon et al., 1998 who 

follow Naggs, 1994, and Cowie et al., 1995) 

mooreana (W.G. Binney, 1857), Polygyra = Daedalochila mooreana (W.G. Binney, 1857) (fide Coles and Walsh, 

2006) 

multilineata (Say, 1821), Triodopsis = Webbhelix multilineata (Say, 1821) (fide Emberton, 1988) 

mutabilis Hubricht, 1951, Pallifera = Megapallifera mutabilis (Hubricht, 1951) (fide Hubricht, 1985) 

obstricta (Say, 1821), Triodopsis = Xolotrema obstrictum (Say, 1821) (fide Emberton, 1988) 

orbiculata (Say, 1818), Helicina = Oligyra orbiculata Say, 1818 (fide Turgeon et al., 1998, who apparently follow 

Baker, 1922) 

71

pennsylvanicus (Green, 1827), Mesodon = Patera pennsylvanica (Green, 1827) (fide Emberton, 1991) 

pentadelphia (Pilsbry, 1900), Retinella = Glyphyalinia pentadelphia (Pilsbry, 1900) (fide Hubricht, 1985) 

peregrina Rehder, 1932, Polygyra = Daedalochila peregrina (Rehder, 1932) (fide Coles and Walsh, 2006) 

perigraptus Pilsbry, 1894, Mesodon = Patera perigrapta (Pilsbry, 1894) (fide Emberton, 1991) 

plicata Say, 1821, Polygyra = Daedalochila plicata (Say, 1821) (fide Coles and Walsh, 2006) 

poirieri (Mabille, 1883), Lehmannia = Limax marginatus Müller, 1774, however U.S. specimens often are misidentified 

introductions of Lehmannia valentiana (Ferussac, 1821) (fide Branson, 1980) with which it has often 

been mistakenly synonymized (e.g. Dundee, 1974) 

praecox H.B. Baker, 1930, Retinella = Glyphyalinia praecox (H.B. Baker, 1930) (fide Hubricht, 1985) 

pustuloides (Bland, 1858), Polygyra = Lobosculum pustuloides (Bland, 1858) (fide Emberton, 1994) 

pustula (Ferussac, 1832), Polygyra = Lobosculum pustula (Ferussac, 1832) (fide Emberton, 1994) 

raderi (Dall, 1898), Retinella = Glyphyalinia raderi (Dall, 1898) (fide Hubricht, 1985) 

rhoadsi (Pilsbry, 1899), Retinella = Glyphyalinia rhoadsi (Pilsbry, 1899) (fide Hubricht, 1985)roundyi Morrison, 

1935, Paravitrea = Helicodiscus roundyi (Morrison, 1935) (fide Hubricht, 1985) 

roemeri (Pilsbry and Ferriss, 1906), Retinella = Glyphyalinia roemeri (Pilsbry and Ferriss, 1906) (fide Hubricht, 

1985) 

roemeri (Pfeiffer, 1848), Holospira = Metastoma roemeri (Pfeiffer, 1848) (fide Hubricht, 1985) 

roemeri (Pfeiffer, 1848), Mesodon = Patera roemeri (Pfeiffer, 1848) (fide Emberton, 1991) 

sayanus (Pilsbry, 1906), Mesodon = Appalachina sayana (Pilsbry, 1906) (fide Emberton, 1994) 

saludensis (Morrison, 1937), Clappiella = Helicodiscus saludensis (Morrison, 1937) (fide Hubricht, 1985) 

sargentianus (C.W. Johnson and Pilsbry, 1892), Mesodon = Patera sargentianus (C.W. Johnson and Pilsbry, 1892) 

(fide Emberton, 1991) 

schiedeanus (Pfeiffer, 1841), Bulimulus = Rabdotus mooreanus (Pfeiffer, 1868) (fide Hubricht, 1985) 

sculptilis (Bland, 1858), Retinella = Glyphyalinia sculptilis (Bland, 1858) (fide Hubricht, 1985) 

smithi (G.H. Clapp, 1905), Mesodon = Inflectarius smithi (G.H. Clapp, 1905) (fide Emberton, 1991) 

striolata (Pfeiffer, 1828), Hygromia = Trochulus striolata (Pfeiffer, 1828) (fide Dundee, 1974 for synonymy with 

Trichia striolata (Pfeiffer, 1828) and ICZN, 2004 for retention of Trochulus over Trichia) 

subpalliatus (Pilsbry, 1893), Mesodon = Inflectarius subpalliatus (Pilsbry, 1893) (fide Emberton, 1991) 

texasiana (Moricand, 1833), Polygyra = Linisa texasiana (Moricand, 1833) (fide Emberton, 1995) 

tholus (W.G. Binney, 1857), Polygyra = Daedalochila mooreana (W.G. Binney, 1857) (fide Pilsbry, 1940) 

tridens Morrison, 1985, Pilsbryna = Helicodiscus tridens (Morrison, 1935) (fide Hubricht, 1985) 

tridens Morrison, 1935, Pilsbryna = Helicodiscus tridens (Morrison, 1935) (fide Hubricht, 1985) 

triodontoides (Bland, 1861), Polygyra = Daedalochila triodontoides (Bland, 1861) (fide Emberton, 1994) 

troostiana I. Lea, 1839, Polygyra = Daedalochila troostiana (I. Lea, 1839) (fide Coles and Walsh, 2006) 

uvidermis Pilsbry, 1890, Vitrinizonites = Vitrinizonites latissimus (J. Lewis, 1875) (fide Hubricht, 1985) 

uvulifera (Shuttleworth, 1852), Polygyra = Daedalochila uvulifera (Shuttleworth, 1852) (fide Emberton, 1994) 

vanattai (Walker and Pilsbry, 1902), Retinella = Pilsbryna vanattai (Walker and Pilsbry, 1902) (fide Slapcinsky and 

Coles, 2004) 

varians (Menke, 1829), Cepolis = Hemitrochus varians (Menke, 1829) (fide Emberton, 1991b; Miller and Naranjo- 

Garcia, 1991) 

virginica Morrison, 1937, Retinella = Glyphyalinia virginica (Morrison, 1937) (fide Hubricht, 1985) 

vermiculata Müller, 1774, Otala = Eobania vermiculata (Müller, 1774) (type of genus Eobania Hesse, 1913, fide 

Richardson, 1980) 

walkeri (Pilsbry, 1900), Paravitrea = Paravitrea umbilicaris (Ancey, 1887) (fide Hubricht, 1985) 

weatherbyi W.G. Binney, 1874, Pallifera = Megapallifera weatherbyi (W.G. Binney, 1874) (fide Hubricht, 1985) 

wetherbyi (Bland, 1873), Mesodon = Fumonelix wetherbyi (Bland, 1873) (fide Emberton, 1991) 

wheatleyi (Bland, 1883), Retinella = Glyphyalinia wheatleyi (Bland, 1883) (fide Hubricht, 1985) 

wheatleyi (Bland, 1860), Mesodon = Fumonelix wheatleyi (Bland, 1860) (fide Emberton, 1991) 

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