Tiny Jewels: an Introduction to Pupillid Taxonomy, Ecology, and Collection
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Page 22 AMERICAN CONCHOLOGIST Vol. 37, No. 1 Tiny Jewels: An introduction to pupillid taxonomy, ecology, and collection Jeff Nekola Introduction Across most of North America roughly 50% of all land 2000 illustrated, with only 11 of these being pupillids. The large snail species and almost 90% of all individuals are less than 5mm shells which dominate this book in actuality represent less than 2% in maximum dimension, or approximately the size of Lincoln’s chin of all individuals across not only North America, but also New on a penny (Fig. 1). Of these, the Pupillidae make an especially Zealand, eastern Australia, western Africa, and southeast Asia as important contribution to biodiversity. This family (here used in well. the expansive, historical sense as outlined by H.A. Pilsbry) includes Adequate documentation of land snail diversity thus the genera Bothriopupa, Chaenaxis, Columella, Gastrocopta, requires investigators to efficiently collect and accurately identify Pupilla, Pupisoma, Sterkia, and Vertigo, and encompasses almost pupillids. Unfortunately, neither has been common. Two major 10% of the entire USA and Canadian fauna (~125 species), with an reasons for this exist. First, as none of the taxa exceeds 6mm in unknown number of additional taxa residing in Mexico. North maximum dimension, accurate identification requires critical America represents the global biodiversity center for Chaenaxis, examination at 20-40x magnification. This has made even Gastrocopta, Pupilla, and Vertigo, harboring at least 2/3 of all laboratory identifications suspect, with there being a high incidence known global taxa and all known morphological groups. The of misidentification and mixed lots in museum collections. For Pupillidae are the third most diverse family east of the continental instance, I recently found that over 90% of the Vertigo collections divide (12% of the total fauna), exceeded only by the Polygyridae in the National Museum of Canada were incorrectly identified. (30%) and Zonitidae (22%). Pupillids alone make up approximately Second, most species are cryptic, being found primarily in one-third of both total species diversity and number of individuals decomposed leaf litter. As a consequence they tend to be for site-scale faunas throughout North America, with this level undersampled by those relying on locating individuals by eye. This approaching 80% - 100% within the northern taiga and tundra, acidic has led to the lack of documentation of not only the normal range wetlands and savannas of the southeastern coastal plain, and fog- of morphological variation within and between populations and belt chaparral of the California coast. In some places, such as the taxa but also the true geographic and ecological ranges for most pocosins of the North Carolina coastal plain, the number of pupillid species. As a result, hasty conclusions concerning specific identity, land snails can exceed 5,000 individuals per square meter. biogeography, and ecology in this family has been unfortunately commonplace. Over the last decade, colleague Brian Coles of the Welsh National Museum and I have observed pupillid communities across most of North America, ranging from central Quebec, Hudson’s Bay and the Alaskan north slope to Florida, Texas, the desert southwest, and coastal California. In this time, we have collected from over 1,700 stations, including almost 75% of North American pupillid taxa and over 200,000 total individuals. In the process we have described two new species, with perhaps another 5-6 waiting in the wings. This experience provides a unique perspective with which to introduce this important group of mollusks to the shell collecting community. Important Concological Features for Shell Identification Identification of pupillid shells requires careful observation and comparison of approximately two-dozen shell characteristics. While a number of these are also commonly used in identification of taxa from other families, use of some is almost completely limited to pupillids. The more important of these features are: Figure 1: Representative micro-snails from a northeastern Wisconsin wet meadow. Apex: the uppermost 2-3 whorls of the shell Body Whorl: the final full whorl in an adult shell Yet, in spite of their ecological importance, pupillids and Callus: calcified thickening of the palatal wall of the aperture, other microsnails have been almost completely ignored by often deposited between lamellae collectors. For instance, in Abbott’s “Compendium of Landshells” only a few dozen microsnail species are included out of the roughly March 2009 AMERICAN CONCHOLOGIST Page 23 Crest: a bowing out of the shell immediately in back of the aperture as seen in side view Palatal Depression: indentation of the shell surface at the location of the palatal lamellae Penultimate Whorl: the next to the last whorl in an adult shell Sinulus: indentation of the aperture margin along the palatal wall Suture: indentation of the shell surface where two whorls meet Figure 3: Location of the major apertural lamellae used in pupillid identification, illustrated through use of a Figure 2: Major shell features used to identify pupillid taxa, Vertigo ovata SEM image. illustrated through use of SEM images of Vertigo elatior (left), V. bollesiana (upper right) and V. cristata (lower right). In the genus Gastrocopta, the angular and parietal lamellae are fused into a single body, which is often referred to as an ‘angulo- Overall shell shape is also often diagnostic, ranging from parietal lamella.’ It is important to note that, depending upon the conical to ovoid to cylindrical. Also of considerable importance in species, the strength of a given lamella, or the total number of pupillid taxonomy is the position and appearance of a variable lamellae, may vary between individuals both within and between number of shell thickenings often found in the shell aperture. While populations. This level of variability can only be assessed when often colloquially referred to as ‘teeth’ this term is misleading as large numbers of shells are observed over a broad ecological and/ they are in no way related to the snail’s gastrointestinal track. In or geographic range. fact, it remains unclear what function (if any) they serve. Because of their thin, plate-like appearance, it is best to refer to these as Species Concepts and Taxonomic Issues ‘lamellae’. It should be noted, however, that H.A. Pilsbry made Some believe that because their critical distinguishing the terminology unnecessarily complex by referring to the palatal features are often no larger than a tenth of a mm (or less) in size, lamellae as ‘lobes.’ The up to seven or more lamellae in the shell they must be of no taxonomic merit. These researchers have tended aperture are designated by their position: to recommend the lumping of many taxa. On the other hand, others have felt that even the slightest differences must have taxonomic Angular: the plate on the parietal wall of the aperture to the right merit, and have advocated the elevation of many pupillid forms to of the parietal lamella in dextral shells species-level rank. Basal: the plate on the bottom left side (in dextral shells) of the Until recently, there has been no objective way to assess aperture below the columellar lamella these opposing viewpoints, as pupillids often demonstrate a high Columellar: the plate on the columellar wall of the aperture degree of aphallism and limited levels of anatomical variation. As Infraparietal: the plate on the parietal wall to the left of the parietal a result, the level of taxonomic discourse often descended into name- lamella in dextral shells calling and appeals to authority. Thanks to DNA sequencing, Parietal: major plate in the middle of the parietal wall of the independent information can now be obtained. We recently aperture completed an initial survey of DNA sequences from 25 pupillid Lower Palatal: lowermost of the two major plates often found on taxa, focusing on the Vertigo gouldii group (Figure 4). While these the palatal wall analyses demonstrate that shell features should not be used to infer Upper Palatal: uppermost of the of the two major plates often evolutionary relationships, they usually function well in assigning found on the palatal wall species-level differences. The beliefs of some of the most ardent lumpers can thus be shown to be false, with the former western ‘V. Page 24 AMERICAN CONCHOLOGIST Vol. 37, No. 1 Figure 4: Phylogenetic tree of the Vertigo gouldii group and other various pupillids. Nodes with strong to moderate support based on four different analytical approaches have been provided to the left of each node. March 2009 AMERICAN CONCHOLOGIST Page 25 gouldii’ subspecies actually representing five highly distinct with it being possible for small taxa to be blown across the open species-level taxa. Ardent splitters were also shown to be wrong, ocean from one island to the next. Recent genetic evidence also particularly in the case of the Vertigo arthuri clade which some had demonstrates that small land snails have been able to migrate from suggested to represent up to six species. This did not surprise us Western Europe to Tristan da Cuna (isolated islands in the South as we had already observed complete conchological intergradation Atlantic) and back, likely on migrating shore birds. Such long- between these forms across much of North America. These analyses range passive dispersal is aided by their adhesive mucus, which also demonstrated that some species assumed to be shared between makes it easy for them to adhere to passing vertebrates. This can North America and Eurasia actually represent species complexes. be easily demonstrated for pupillids by noting the individuals that In the case of Pupilla muscorum the distinct Eurasian lineages have commonly stick to a hand run through damp sedge or grass leaves. escaped and become naturalized across eastern North America. As a result, North American pupillids are characterized These analyses also help us understand more about their origin.