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Taxonomy and Biogeography of Hemphillia (Gastropoda: Pulmonata: Arionidae) in North American Rain Forests, with Description of a New Species (Hemphillia Skadei Sp

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Taxonomy and Biogeography of Hemphillia (Gastropoda: Pulmonata: Arionidae) in North American Rain Forests, with Description of a New Species (Hemphillia Skadei Sp Canadian Journal of Zoology Taxonomy and biogeography of Hemphillia (Gastropoda: Pulmonata: Arionidae) in North American rain forests, with description of a new species (Hemphillia skadei sp. nov., Skade’s Slug) Journal: Canadian Journal of Zoology Manuscript ID cjz-2017-0260.R1 Manuscript Type: Article Date Submitted by the Author: n/a Complete List of Authors: Lucid, Michael;Draft Idaho Department of Fish and Game Rankin, Andrew; University of Idaho, Department of Biological Sciences Espindola, Anahi; University of Idaho, Department of Biological Sciences Chichester, Lyle; Independent Ehlers, Shannon; Idaho Department of Fish and Game; Missouri Department of Conservation Robinson, Lacy; Idaho Department of Fish and Game; Yellowston to Yukon Conservation Initiative Sullivan, Jack; University of Idaho, Department of Biological Sciences BIODIVERSITY < Discipline, GASTROPODA < Taxon, TEMPERATE < Keyword: Habitat, TAXONOMY-SYSTEMATICS < Discipline, PHYLOGENY < Discipline, MORPHOLOGY < Discipline, FOREST < Habitat https://mc06.manuscriptcentral.com/cjz-pubs Page 1 of 33 Canadian Journal of Zoology 1 1 Title: Taxonomy and biogeography of Hemphillia (Gastropoda: Pulmonata: Arionidae) in 2 North American rain forests, with description of a new species (Hemphillia skadei sp. nov., 3 Skade’s Slug). 4 5 Authors: Lucid M. K.1*, Rankin A.2, Espíndola A.3, Chichester L.4, Ehlers S.5, Robinson L.6, 6 Sullivan J.7 7 1Idaho Department of Fish and Game, 2885 Kathleen Ave. Coeur d’Alene, Idaho, 83815, 8 USA. [email protected] 9 2Department of Biological Sciences, Life Sciences South 252, 875 Perimeter Drive MS3051, 10 University of Idaho, Moscow, Idaho, 83844-3051, USA. [email protected] 11 3Department of Biological Sciences, Life Sciences South 252, 875 Perimeter Drive MS3051, 12 University of Idaho, Moscow, Idaho, Draft83844-3051, USA. [email protected] 13 4209 Chestnut Springs Way, Williamston, South Carolina, 29697, USA. 14 [email protected] 15 5Idaho Department of Fish and Game, 2885 Kathleen Ave. Coeur d’Alene, Idaho, 83815, 16 USA. Current address: Missouri Department of Conservation, 2929 County Road 618, 17 Ellington, MO 63638, USA. [email protected] 18 6Idaho Department of Fish and Game, 2885 Kathleen Ave. Coeur d’Alene, Idaho, 83815, 19 USA. Current address: Yellowstone to Yukon Conservation Initiative; PO Box 733, 20 Sandpoint, Idaho 83864, USA. [email protected] 21 7Department of Biological Sciences, Life Sciences South 252, 875 Perimeter Drive MS3051, 22 University of Idaho, Moscow, Idaho, 83844-3051, USA. [email protected] 23 Corresponding author: Lucid M. K. Idaho Department of Fish and Game, 2885 Kathleen 24 Ave. Coeur d’Alene, Idaho, 83815, USA. Email: [email protected]. Phone: +1 25 208 830-1451, Fax: +1 208 769-1418. https://mc06.manuscriptcentral.com/cjz-pubs Canadian Journal of Zoology Page 2 of 33 2 Lucid M. K., Rankin A., Espíndola A., Chichester L., Ehlers S., Robinson L., Sullivan J. “Taxonomy and biogeography of Hemphillia (Gastropoda: Pulmonata: Arionidae) in North American rain forests, with description of a new species (Hemphillia skadei, Skade’s Slug)”. 26 Abstract 27 Hemphillia (jumping slugs) species diversity was studied across its range in western North 28 America’s inland temperate rainforests. The taxonomic relationships among jumping slug 29 populations were clarified by integrating morphological, molecular, and biogeographic 30 approaches. A new species, Hemphillia skadei was discovered in this process and is 31 described herein. We base this taxonomic decision on molecular comparison of 32 representatives from other Hemphillia species and three morphological characters which 33 distinguish H. skadei from its sister species H. camelus. The distribution of H. skadei and H. 34 camelus is described along with the notable lack of detection of H. danielsi within the 35 primary survey area. 36 Draft 37 Keywords: biodiversity, forest, Gastropoda, morphology, phylogeny, temperate, taxonomy- 38 systematics https://mc06.manuscriptcentral.com/cjz-pubs Page 3 of 33 Canadian Journal of Zoology 3 39 Introduction 40 The demographic and distribution data necessary for landscape level species conservation are 41 only usable in the context of accurate taxonomic frameworks (Wilson 2000). Within this 42 context, spatially expansive inventory programs are a necessary component of species status 43 assessments, particularly for data-deficient species (e.g., Bland et al. 2017), to determine if 44 they should be targeted for conservation action. North American terrestrial gastropods are a 45 case-in-point. Mollusk species comprise the second largest phylum (Mollusca) of Animalia, 46 and this highlights their ecological impact on the biosphere. Mollusks serve a wide variety of 47 ecological roles including decomposing organic matter, hosting parasites, vectoring disease, 48 soil building, and providing a prey base for many species (Jordan and Black 2012). Although 49 mollusks are listed as the third largest group of International Union of Conservation of Nature 50 (IUCN) threatened animal species globallyDraft and although they are the most numerous group of 51 IUCN threatened species in North America (http://www.iucnredlist.org/, accessed 27 52 February 2017), biological knowledge is limited and much remains to be discovered. Without 53 thorough knowledge of the ecology, evolution, and systematics of these animals it will be 54 difficult to describe and conserve the diversity they mollusks represent. 55 Up to 25% of North American terrestrial gastropods await description (Nekola 2014) 56 and new species and genera from the Pacific Northwest are described regularly (e.g., Leonard 57 et al. 2003; Leonard et al. 2011). Some areas of the Pacific Northwest have exceptionally 58 high terrestrial mollusk diversity and endemism (Burke 2013; Frest and Johannes 1995), 59 some of which may still be undescribed. Jumping slugs, gastropods in the genus Hemphillia, 60 are endemic to North America’s Pacific Northwest. The genus currently consists five 61 recognized species (Burke 2013) distributed in a mesic forest disjunct pattern (Brunsfeld et 62 al. 2001). https://mc06.manuscriptcentral.com/cjz-pubs Canadian Journal of Zoology Page 4 of 33 4 63 Hemphillia dromedarius Branson, 1972, H. glandulosa Bland & W. G. Binney, 1872, 64 and H. malonei Pilsbry, 1917, have coastal distributions while H. camelus Pilsbry & Vanatta, 65 1898, and H. danielsi Vanatta, 1914, are distributed in inland forests. Hemphillia burringtoni 66 Branson, 1972, a sixth purported species, is not supported by molecular analysis (Wilke & 67 Ziegltrum 2004). Hemphillia pantherina Branson, 1975, a seventh purported species, was 68 described from a single coastal specimen (Branson 1975) but is now widely viewed as not 69 warranting specific recognition (Burke 2015; Burke pers. comm.). 70 The two inland Hemphillia species (H. camelus and H. danielsi) were included as part 71 of a multi-taxa inventory program in the inland Pacific Northwest (Lucid et al. 2016). 72 Specifically, H. camelus was known to occur within the study area (Burke 2013), and H. 73 danielsi was known to occur adjacent to (Hendricks 2003), and predicted to occur within 74 (Burke 2013), the study area. Our objectivesDraft were to 1) identify the Hemphillia species 75 diversity from northern Idaho and the surrounding regions, 2) clarify the taxonomic 76 relationships of those entities integrating morphological, molecular, and biogeographic 77 approaches, 3) describe anatomical, molecular, distributional, and natural history 78 characteristics of a new taxon discovered in this process. 79 80 Methods 81 Study Area 82 Centered on northern Idaho’s Panhandle region, the study area encompassed portions of 83 northeastern Washington, and northwestern Montana, in the USA (Fig. 1). It included parts of 84 the Selkirk, Purcell, West Cabinet, Coeur d' Alene, and Saint Joe mountain ranges. The 85 topography is mountainous, ranging in elevation from 702 to 2326 m, with a climate 86 characterized by mild summers and wet and moderately cold winters. The heavily forested https://mc06.manuscriptcentral.com/cjz-pubs Page 5 of 33 Canadian Journal of Zoology 5 87 area is dominated by a diverse mix of conifer species and is characterized as supporting 88 inland temperate rainforest (DellaSala et al. 2011). 89 90 Field Methods 91 We stratified our study area into 920 5x5 km sampling cells and conducted 992 surveys in 92 879 cells. Our survey sites were either randomly selected but biased to roads and trails (n = 93 842) or sub-selected from random Forest Inventory and Analysis plots (n = 150) (Bechtold 94 and Patterson 2005) based on site characteristics (stand age, elevation, and distance to road; 95 see Lucid et al. 2016 for details). Each gastropod survey site was visited twice at approximate 96 14 day intervals from May-November, 2010-2014. Each survey consisted of 1) three 30x30 97 cm cardboard cover board traps (Boag 1982) from which specimens were collected after a 14 98 day deployment period, 2) collection,Draft freezing, and drying of one liter of leaf litter (Coney et 99 al. 1981) from which gastropods were sorted, and 3) two 15 minute timed visual searches 100 conducted 14 days apart. Air temperature was recorded every 90 minutes at survey sites for 101 one year as described in Lucid et al. (2016). Field personnel collected all terrestrial 102 gastropods encountered, drowned each specimen in water, and then preserved specimens in 103 70% ethanol. 104 105 Morphological
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