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Invasive Land Snails and Slugs in North America

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Invasive Land Snails and Slugs in North America Invasive Land Snails and Slugs in North America National Plant Diagnostic Network Malacology Workshop UC Davis 16-17 June 2015 David G. Robinson USDA APHIS PPQ National Malacology Laboratory Academy of Natural Sciences of Drexel University, Philadelphia, PA INTRODUCTION Most snails and slugs found associated with human habitation and agricultural areas, and other areas of human disturbance (= synanthropic species) belong to invasive (non-native) species Native terrestrial species are generally found in undisturbed areas, and may be used as indicators of the relative health of a natural environment; native species as part of a high biodiversity indicate a stable habitat that more likely to be resistant to invasives Do invasive species outcompete native species, or merely occupy disturbed areas where the native species have already been driven out ? The external characters of a slug, or the snail’s shell has traditionally been used to identify the species Increasingly, the genitalia are required to identify the snail’s species Molecular techniques are also being used to further separate cryptic species that are inseparable using anatomical or shell characters Selected quarantine-significant species that are invasive in the continental United States and their diagnostic characters will be discussed here Pulmonate slug (from Kerney & Cameron, 1979) Traditionally slugs were described using external morphological characters Introduced terrestrial slug families AGRIOLIMACIDAE LIMACIDAE MILACIDAE Pulmonate slugs (from Herbert, 1997) ARIONIDAE VERONICELLIDAE Pulmonate slugs (from Herbert, 1997) As malacologists began to study the internal anatomy, especially the genitalia, this led to the recognition that there were many more species than previously thought, while external characters (e.g. color) became less important Basic dissection of terrestrial slug (from Kerney & Cameron, 1979) In 1999, the USDA initiated the North American Slug Project, to document all the invasive slugs and their distributions in the United States and Canada Slugs are being submitted from across the country, with the ultimate goal of documenting all invasives and their distribution , on a county by county basis Dissection of all specimens is required, with molecular analysis (using CO1 and 16S genes) being used to check dubious identifications Deroceras laeve (Müller, 1774) Genitalia of D. laeve (from Barker, 1999) Deroceras invadens (Reise et al., 2011) (photo courtesy H. Reise) (Reproductive anatomy from G. Barker, 1999) Lehmannia valentiana (A. de Férussac, 1821) Unfortunately, the molecular analysis of the Palearctic Arionidae has made identification of the species more complicated Arion s.s. consists of 3 anatomically-defined taxa: Arion ater (Linné, 1758), A. rufus (Linné, 1758), and A. vulgaris Moquin-Tandon, 1855 (= A. lusitanicus non Mabille, 1868); from molecular evidence, there are 4 distinct clades, designated A,B,C, & D. However, the 3 anatomical taxa do not correspond exactly with the molecular clades: A. ater consists of clade A A. rufus consists of clades A, B & D A. vulgaris consists of clades A & C Arion vulgaris Moquin-Tandon, 1855 from Germany (photo courtesy of I. Richling) In the United States and Canada, we have found no evidence of any anatomical A. ater ALL of the large Arion slugs (i.e. Arion s.s.) analyzed from North America are anatomically A . rufus, almost all belonging to clades B & D; only a single clade A specimen has been found to date (in OR) Arion rufus (Linné, 1758) (photo at top courtesy of M. Hitchcox; middle photo courtesy of P. Marquez; bottom photo courtesy of I. Richling) Similarly, the anatomical species A. subfuscus (Draparnaud, 1801) consists of at least two species, including the cryptic A. fuscus (Müller, 1744) , plus at least 5 subclades, S1-S5 In North America, we have confirmed the presence of A. fuscus and A. subfuscus – subclades S1 and S2, with the latter restricted to the northeast USA At present we are far from documenting all the invasive slugs in the United States, and our molecular studies are only at the early stages of the project Pulmonate snail shell (from Kerney & Cameron, 1979) Similarly, in the “GOOD OL’ DAYS,” conchologists identified snails described the shell using diagnostic characters Species descriptions were often based on long dead (faded) museum specimens Little or nothing was known of the living animal or its anatomy Currently malacologists dissect the snail’s body to find diagnostic characters Initial preparation of snail (drowning in water for 12 hours, then preserving in 75- 80% ethanol) [Optionally anesthetize using tobacco, menthol crystals, etc.] Removal of the snail from its shell Exposure of organs by opening the body cavity between the head and the mantle [Optionally access the body cavity through the lung area, above the mantle] Genitalia usually pure white in color, on the right side of Basic dissection of terrestrial pulmonate snail the body (from Kerney & Cameron, 1979) Male and female tracts unite at the atrium, which opens through the genital pore (= gonopore) The “Brown garden snail” – Cornu aspersum (Müller, 1774) (= Helix aspersa, Cryptomphalus aspersus, Cantareus aspersus) An edible snail widely (and intentionally) introduced worldwide, in temperate, subtropical and tropical zones Introduced into the continental United States in the 1850s A pest of a wide range of agricultural and horticultural plants, especially Citrus spp. Currently costs $7-10 million in control/suppression programs in California alone Currently widespread along the West Coast (CA, OR, WA), the southern tier of states, with periodic infestations elsewhere (e.g. NJ, PA, NC, etc.) with control programs in place As an example of pulmonate Cornu aspersum (Müller, 1774) genitalic anatomy: Genitalia of C. aspersum (Müller, 1774) (from Kerney & Cameron, 1979) FEMALE: One large dart sac (= stylophore) containing a large calcareous dart Two mucus glands, with numerous branches (“spaghetti”) Very long bursa copulatrix duct that branches into two, one leading to the diverticulum, the other to the bursa copulatrix reservoir MALE: The phallus extends from the atrium to (usually) where the penial retractor muscle is attached The epiphallus extends from (usually) the penial retractor muscle attachment to where the vas deferens connects The flagellum extends beyond where the vas deferens connects The vas deferens connects to the prostate (not shown) The relative lengths and shape of male genitalia can be diagnostic Introduced species: Theba pisana (Müller, 1774) - HELICIDAE Theba pisana is considered by most agricultural authorities worldwide to be the worst snail pest It is believed to be originally from western North Africa (Morocco area); it spread throughout the Mediterranean Basin during Roman times Introduced to California in 1914 (Hanna, 1966), and subsequently re-introduced several times after being “successfully eradicated” (or perhaps it was never completely eradicated ?) In the mid-1990’s, the last program by the California Department of Food and Agriculture (CDFA) was stopped just short of a complete eradication; the remaining population was reportedly in an isolated area in San Diego County, and presumably not considered a threat to agriculture A USDA PPQ survey in June 2005 showed that T. pisana is staging a “come-back” and was found throughout San Diego County Introduced species: Theba pisana (Müller, 1774) (continued) Theba pisana is a very serious pest in southern Australia causing severe losses in wheat, barley, and oilseed rape It also reaches such high numbers that it clogs up harvesting machinery T. pisana is a pest as (a) a voracious plant feeder and (b) due to its massing behavior Heavy infestations of T. pisana in the York Peninsula, South Australia; snails massing on fence posts, and on wheat stubble after harvest (photos courtesy of M. Leyson and A. Lush) Introduced species: Theba pisana (Müller, 1774) (continued) 2012-2014 infestations of Theba pisana and other species introduced into Australia from the Mediterranean species are the worst in Australia’s history and many farmers are selling or abandoning their farms Introduced species: Theba pisana (Müller, 1774) (continued) Infestations of Theba pisana on ornamental fruit trees in San Diego County Infestations of Theba pisana on grass on low shrubs in San Diego County Introduced species: Theba pisana (Müller, 1774) (continued) FEMALE: One dart sac (=stylophore) containing a large calcaerous dart Two mucus glands, not branched with distinct alveolar structure MALE: The phallus is swollen; internally with a double verge, the first very broad, the second protruding into atrial section of phallus The epiphallus is well developed, but flagellum is rudimentary (barely visible) Genitalia of Theba pisana (Müller, 1774) (from Schileyko, 2006) Introduced species: Theba pisana (Müller, 1774) (continued) Distribution of Theba pisana (Müller, 1774) in the continental United States: San Diego County, California Introduced species: Trochoidea elegans (Gmelin, 1791) - HYGROMIIDAE Established in a graveyard in Charleston, SC, in the late 1800s, and still surviving well into the 1940s USDA PPQ surveys in 2002, 2004, and 2005 did not find any specimens Introduced into Sunny Point, NC, possibly in November 1999; detected May 2000; eradicated within a year Up to 8 mm in height, and 10 mm in maximum diameter. Shell conical above, very flattened below, with sharply keeled, somewhat overlapping, flattened whorls. Genitalia of T. elegans (Gmelin,
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