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Do Philomycus carolinianus (Gastropoda: Philomycidae) prefer to congregate? PDF

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THE NAUTILUS 125(2):83-85, 2011 Page 83 Do Philomycus ccirolinianus (Gastropoda: Philomycidae) prefer to congregate? TimothyA. Pearce Katherine A. Porter Section of Mollusks Carnegie Museum ofNatural History Pittsburgh, PA 15213 USA [email protected] ABSTRACT Huddling and congregation are not unique to Philomycus species. Orstan (2007) noted huddling by Field observations of land slugs Philomycus carolinianus individuals ofthe same and ofdifferent species ofslugs, (Bose, 1802) together at shelters led us to ask whether they but we do not know whether the same mechanisms were attracted to conspecifics versus attracted to a limited trherseoeurscheelt(ee.rgs.,toshtehlrteeer)s.luOgsurinleaabcohrarteoprlyicaetxep,ewriitmhenttheoefxfpeerce-d atiroen,acatlitnhgouignhiwnteer-noawndhaivnetrab-estpteecrifuincdehurdsdtlaensd.inIgnoaldtdhie- tation that slugs would occurtogether under asingle shelter ii systematic^ of Philomycidae (Fairbanks, 1986; 1989; they preferred to be with conspecifics, while slugs would 1990; 1993; 1998; Tsai et ah, 2005; Tsai and Wu, 2008), occur singly under each shelter if they avoided conspecifics. we still know relatively little about their biology. In this We found that slugs chose shelters regardless ofthe presence note, we examine congregation and huddling behavior of conspecifics, suggesting that their occurrence together in among individuals ofa single species, the eastern North nature is due to the sharing of limited shelter resources. American slug Philomycus carolinianus (Bose, 1802). Additionalkey words: landslugs, shelter, limited resource MATERIALS AND METHODS Slugs were wild-caught in spring 2009 from the Patux- INTRODUCTION ent National Wildlife Refuge, Prince George’s County, Maryland (39o03' N, 76°49' W). The slugs were used in Animals congregate for many reasons. In some cases, another experiment over the summer, then provided theybenefit from the presence ol others. In other cases, to us for this study, which was conducted from 27 they occur together in an area because a resource is October to 21 November 2009. Voucher specimens, presentthere. Explanations forcongregation behavior in preserved in 80% EtOH within a day post mortem, terrestrial mollusks include prevention of water loss are deposited at Carnegie Museum of Natural History (Dundee et ah, 1975; Cook, 1981; Waite, 1988) and (CM104587). escape from ground-level heat (Cook, 2001). Wepreparedthree arenas. Eachwas 44 x 28 x 22.5cm In nature, gastropods are commonly associated with high. The arenas were corrugated cardboard boxes, each mm woody debris, and can be found congregated in areas lined with a plastic bag and covered with a 1.4 mesh with large amounts ofsuch debris (Kappes et ah, 2009). screen. To maintain humidity in the arenas, each screen Individuals of species ol slugs in the genus Philomycus was covered with a stack of 12 moistened paper towels. Rafinesque, 1820 are sometimes seen in close proximity Each paper towel had a surface area ol 276 cirri. to each other, especially near coarse woody debris. In Each arenafloorwas coveredwith soil approximately 1 cm some cases, these slugs are found in physical contactwith deep. In each arena,weplacedthreebarkshelters measur- eachother(aphenomenon known as huddling). However, ing approximately 12 cm long, 10 cm wide, and 1.5 cm Philomycus individuals do not always exhibit huddling in thick. Each arena also contained a water dish and a food thewild, and thepresence ofmultiple slugs inclose prox- dish that contained fish food flakes and mushroom slices. imityisyetto be explained. Twopossible explanations are The amounts offood and water provided were ample for thatindividualsofPhilomycus species aregregarious (pre- the number of slugs in each arena, ensuring that neither fer to associate with each other) and that they tend to food nor moisture was limiting. Shelter was the limiting congregate at the locations oflimiting resources, regard- factor; other than the bark, food dish, and water dish, the less ofthepresenceofotherslugs. arenawas devoidofobjects thatcouldfunction as shelters. . Page 84 THE NAUTILUS, Vol. 125, No. 2 We placed three mature slugs in each arena. For each Because these slugs appear to assort randomly, replicate, we observed slug positions once daily for six congregations of slugs found in nature may indicate days. Observations were made during the daytime when the presence of a scarce resource (e.g., shelter) in that slugs would more likely be resting under shelters. Time particular location. Therefore, observations ofslug hud- ofday when observations were made ranged from 7:42- dling could be used as evidence for resource scarcity. 21:44 with a mean of 12:55 (95%, Cl 9:00-16:50). For example, some land slugs appear to cluster to con- Because we had a limited number of slugs, we re-used serve moisture (Dundee et al., 1975; Cook, 1981; slugs in subsequent trials to maximize the number of Waite, 1988; although see Welsford et ah, 1990). In replicates. The assigning of individuals to arenas was this experiment, food and moisture were non-limiting. not strictly random. Rather, with the exception of 2 A repetition of the experiment in which moisture, individuals in the last trial, the slugs were grouped in a rather than shelter, was the limiting factor might manner ensuring that the individuals together in one allow us to study whether individuals of Philomycus trial would not be together in a subsequent trial. carolinianiis do huddle to conserve moisture. Investiga- We compared the observed slug positions with a null tions in which other resources were limiting could also model, which assumed a random distribution of slugs address whether huddling occurs due to the scarcity of under the shelters. We generated the null model by other resources. determining the 18 possible arrangements of three All ofthe slugs used in this investigation were mature. slugs under three shelters. This null model predicts An investigation of the effects of age on congregation nine instances of singletons (slugs alone under shel- behavior could provide additional data on the be- ters), six instances of pairs (two slugs under the same havior of Philomycus species. Studies involving species shelter), and three instances of triplets (three slugs other than P. carolinianiis could determine whether under the same shelter). Triplets would be expected to other species show similarbehaviors. be more common if slugs are gregarious and less com- mon if they prefer to be solitary. We used a chi-square ACKNOWLEDGMENTS test to determine the statistical significance of the observations. We are grateful to Megan Paustian for lending us the slugs to use inthese trials. Cagin Unalhelpeddiscuss the design ofthe experimentand helped set up the trials. An RESULTS AND DISCUSSION anonymous reviewer provided helpful comments that improved the manuscript. Table shows the observed and expected numbers of 1 shelters with different numbers of slugs. With eight replicates (a total of 83 observations), we observed 46 LITERATURE CITED instances of slugs alone under shelters, 31 instances of Cook, A. 1981. Huddling and the control ofwater loss by the tthwroeeslsulgusgsunudnedrerthethesasmaemesheslhteelrt,era.ndWesixdiinssctaarndceeds 1o8f s29l:ug289U-m29a8x. pseudoflavus Evans. Animal Behaviour observations in which at least one sing was not under a Cook, A. 2001. Behavioralecology: on doingthe right thing, in shelter or in which a slug was using the food or water therightplaceattherighttime. In: G.M. Barker(ed.)The dish as a shelter. We also discarded two replicates in Biology of Terrestrial Molluscs. CABI Publishing, New which one ofthe slugs died. The observed and expected York, NewYork, pp. 447—487. values are not significantly different from random [yj = Dundee, D.S., M. Tizzard, and M. Traub. 1975. Aggregative 5.33, 2 degrees offreedom, 0.1>P>0.05). behaviorinveronieellid slugs. The Nautilus 89: 69-71 This result suggests that slugs choose shelters ran- Fairbanks, H.L. 1986. The taxonomic status of Philomycus domly,withoutregardtowhetheranotherslugispresent. togatus (Pulmonata: Philomycidae): a morphological and Of particular interest is the observed trend for fewer eMlaelcatcroolpohgoiraet2i7c:c2o7m1p-a2r8i0s.onwithPhilomycuscarolinianiis. triplets than predictedbythe random model (6vs. 13.8). Fairbanks, H.L. 1989. The reproductive anatomy and taxo- Although this trend is not statistically significant, it does nomic status of Philomycus venustus Hubricht, 1953 suggestthatthese slugs are not gregarious. and Philomycus bisdosus Branson, 1968 (Pulmonata: Philomycidae). The Nautilus 103: 20-23. Fairbanks, H.L. 1990. Morphological comparisons ofthe spe- Table 1. Number of shelters having 1, 2, or 3 slugs cies of Megapallifera (Gastropoda: Philomycidae). The underneath. y“ analysis indicates the slugs were behaving Nautilus 104: 71-75. randomly (0.1>P>0.05). Fairbanks H.L. 1993. The reproductive anatomyofPhilomycus sellatus Hubricht, 1972 and Philomycus virginicus Numberofslugsinshelter Observed Expected XOp Hubricht, 1953 (Gastropoda: Philomycidae). The Nautilus 107: 9-13. 1 46 41.5 0.49 Fairbanks, H.L. 1998. Clarificationofthetaxonomicstatusand 2 31 27.7 0.40 reproductiveanatomyofPhilomycusbatchi Branson, 1968 3 6 13.8 4.44 (Gastropoda: Pulmonata: Philomycidae). The Nautilus Totals 83 83 5.33 1 12: 1-5. T.A. Pearce and K.A. Porter, 2011 Page 85 Kappes, H., M. Jabin, Kulfan, P. Zach, and W. Topp. 2009. Tsai, C.L. and S.K. Wu. 2008. A new Meghimatium slug J. Spatial patterns of litter-dwelling ta\a in relation to the (Pulmonata: Philomycidae) from Taiwan. Zoological Stud- amounts ofcoarse woody debris in European temperate ies47: 759-766. deciduous forests. Forest Ecology and Management Waite,T.A. 1988. Huddlingandposturaladjustmentsinresponse 257: 1255-1260. todesiccatingconditionsinDeroceras reticulatum (Muller). Orstan, A. 2007. Contactwith an alien. Tentacle 15: 14-15. Journalof Molluscan Studies54: 249-250. Tsai, C.L., II.II. Lin, and S.K. Wu. 2005. Comparison of Welsford, I.G., PA. Banta, and D.J. Prior. 1990. Size- four philomycid slugs (Gastropoda: Stylommatophora: dependent responses to dehydration in the terrestrial Philomycidae) of Taiwan. Endemic Species Research slug,LimaxmaximusL.:locomotoractivityandhuddlingbe- 7:41-49. havior. Journalof ExperimentalZoology253: 229-234.

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