Bradley, Joseph G.

Eason, Perri K.

English

Cadernos Espinosanos (E-Journal)

279Phyllomedusa - 16(2), December 2017Received 22 September 2017Accepted 30 October 2017Distributed December 2017Short CommuniCationNovel interaction between a pisaurid spider (Araneae: Pisauridae) and an adult Eurycea lucifuga (Caudata: Plethodontidae)Joseph G. Bradley and Perri K. Eason1 Department of Biology, University of Louisville, Louisville, Kentucky 40292, USA. E-mail: jgbrad02@louisville.edu.Keywords: aggression, Cave Salamander, caves, defense.Palavras-chave: agressão, cavernas, defesa, salamandra cavernícola.Aggression can be common in the behavioral repertoire of many salamander species (Hairston 1987). This is in part because many species are VGTTKVQTKCN NGCFKPI VQ KPVTCURGEKſE GURGEKCNN[intrasexual, aggression; however, non-territorial species may still exhibit aggression during courtship periods (Mathis GV CN 1995). Some species also display aggressive behavior (e.g., biting) toward potential predators. For example, the Allegheny Mountain Dusky Salamanders (&GUOQIPCVJWU QEJTQRJCGWU Cope, 1859) and Black-bellied Salamanders [&SWCFTCOCEWNCVWU (Holbrook, 1840)] sometimes respond by biting when attacked by Garter Snakes [6JCOPQRJKUUKTVCNKU (Linnaeus, 1758)] (Brodie GVCN 1989). Aggressive behavior has seldom been reported for the Cave Salamander ('WT[EGCNWEKHWIC 4CſPGUSWG  6JKU URGEKGU FQGUnot generally appear to be aggressive to EQPURGEKſEUCPFKPHCEVECPEQOOQPN[DGHQWPFresting in close proximity in groups of up to 10 individuals in holes and on ledges of rock walls (Howard 1985). It is possible that at least males become more aggressive during breeding season, perhaps only at or near breeding sites; however, such behavioral changes have not been reported, either because they are rare or because breeding habitats may occur deep in cave passages that CTG KPCEEGUUKDNG VQ JWOCPU +PVGTURGEKſEaggressive behaviors toward other salamanders are similarly poorly known for this species. 'WT[EGC NWEKHWIC often occur in multi-species salamander assemblages (Myers 1958, Camp and Jensen 2007), suggesting that competitive interactions may ensue, particularly given that many salamander species are generalist, opportunistic predators with similar diets. However, inactive Cave Salamanders sometimes TGUVXGT[PGCTKGEOCRCTVQVJGTURGEKGUQHsalamanders, including Zig-zag Salamanders (2NGVJQFQP FQTUCNKU Cope, 1889), Dusky Salamanders [&GUOQIPCVJWUHWUEWU4CſPGUSWG1820)], and Streamside Salamanders (#OD[UVQOCDCTDQWTK Kraus and Petranka, 1989) (Joseph G. Bradley, pers. obs.), and aggressive interactions have not been reported in these circumstances.Phyllomedusa 16(2):279–282, 2017© 2017 Universidade de São Paulo - ESALQ ISSN 1519-1397 (print) / ISSN 2316-9079 (online)doi: http://dx.doi.org/10.11606/issn.2316-9079.v16i2p279-282280Phyllomedusa - 16(2), December 2017Salamanders and large carnivorous inverte-brates often share common resources (e.g., microhabitat and invertebrate prey), and either resource competition or the potential of intra-guild predation could result in aggressive encounters between these organisms. Some few such interactions have been observed. For example, Red-backed Salamanders [2NGVJQFQPEKPGTGWU (Green, 1818)] exhibited aggressive postures toward carabid beetles (Gall GV CN 2003), centipedes (Hickerson GV CN 2004), and large spiders (Hickerson GV CN 2017), and they bit centipedes and large spiders that approached them in resident-intruder encounters. Interactions between ' NWEKHWIC and large predacious invertebrates have never been reported to our knowledge. However, Cave Salamanders often co-occur with large spiders in caves. Herein, we provide a description of a novel interaction between a Cave Salamander and a Nursery Web Spider, 2KUCWTKPC sp. (Pisauridae), KP 5CWGTMTCWV %CXG ' 2 ő6QOŒ 5CY[GT 5VCVGPark, Louisville, Kentucky, USA, on 19 June 2016. This interaction took place ca. 3 m into the cave and 2 m high on a brick wall, which was constructed in the cave in the 1800s. There was an egg sac in a nursery web near the spider, so the initiation of the interaction was likely a display of aggression to guard the area around the sac rather than a predation attempt by the spider. In the interaction, the spider aggressively advanced toward the salamander with its front legs in the air, to which the salamander retorted by quickly advancing toward the spider, whereupon the spider withdrew around the nearby corner of the wall. The salamander stood on the tips of its digits on the front limbs, raised its head and chest nearly to the vertical plane exposing the gular region, and kept its hind feet ƀCVQPVQRQHVJGYCNNFKURNC[KPIVJKURQUVWTGKPthe direction of the spider (Figure 1). Advancing again, the salamander moved toward and looked around the corner for the spider, which had TGVTGCVGFDG[QPFKVUYGD#HVGTCUJQTVVKOGmin), the salamander returned to near its original observed position, thus concluding the interaction.Hickerson GV CN (2017) provide the only description we are aware of concerning aggressive encounters between adult spiders and adult salamanders, and they suggest these interactions to be competitive. The initial behavior by the spider appeared to be in defense of the egg sac and/or itself. Cave Salamanders are known to predate large spiders, such as Wolf Spiders (Lycosidae) (Peck and Richardson 1976), but it is uncertain whether this salamander was attempting predation of the URKFGT QT UKORN[ TGCEVKPI VQ VJG URKFGTŏUapproach with aggression; however, the raised-trunk posture may suggest aggression or defense. Smith and Balch (1985) described a similar posture by an individual ' NWEKHWIC when released after being contained for several hours, postulating this to be a defensive stance similar to that observed in Black Salamanders [#PGKFGU ƀCXKRWPEVCVWU (Strauch, 1870)]. Although aggressive behavior has not been documented for 'NWEKHWICVQYCTFEQPURGEKſEUor other species of salamanders, aggressive encounters do apparently occur between Cave Salamanders and spiders, although they may be uncommon. Large lycosid and pisaurid spiders Figure 1. Aggressive encounter between a Nursery Web Spider (Pisaurina sp.) and an adult Cave Salamander (Eurycea lucifuga) in the twilight zone of Sauerkraut Cave. Photo by Joseph G. Bradley.$TCFNG[CPF'CUQP281Phyllomedusa - 16(2), December 2017can be predators of amphibians, and although predation on frogs appears to be more common, TGRQTVUJCXGURGEKſECNN[PQVGFURKFGTRTGFCVKQPon certain life stages of salamanders. Rubbo GVCN (2003) found that adult Wolf Spiders [()NCFKEQUC RWNEJTC (Keyserling, 1877)] attacked juvenile Spotted Salamanders [#OD[UVQOC OCEWNCVWO (Shaw, 1802)], sometimes killing and con suming them. Similarly, Crane and Mathis (2015) witnessed a Fishing Spider, &QNQOGFGU sp. (Pisauridae), predate a larval Ringed Sala mander (#OD[UVQOCCPPWNCVWO Cope, 1886), and interestingly, juvenile European Cave Salamanders, *[FTQOCPVGU (5RGNGQOCPVGU) spp. (Plethodontidae), have been predated by large spiders in caves (Ficetola GV CN 2013). To our knowledge, predation of adult salamanders by spiders has not been documented, but given that spiders can be important predators of adult anurans (Gunzburger and Travis 2005, Menin GV CN 2005), they may also be able to capture adult salamanders.The relationships between salamanders and large predacious invertebrates are not well understood and may be complex and ecologically important, given that these taxa commonly compete for food resources and may predate each other (Hickerson GVCN 2012). Smaller life stages of salamanders (i.e., larval and juvenile forms) may be more susceptible to predation by these carnivorous invertebrates due to both their smaller size and their lack of experience in predatory situations (but see Anthony GV CN 2007). Conversely, larger salamanders may be able to withstand aggressive encounters with large carnivorous invertebrates. Observation of aggressive interactions between salamanders and invertebrates may be relatively rare given the paucity of reported encounters, or they may be OQTG FKHſEWNV VQ QDUGTXG KP PCVWTG KP EQPVTCUVnumerous anecdotal reports of predation of anurans by spiders have been published (Menin GVCN 2005). Clearly, these types of interactions CPF VJGKTGEQNQIKECNUKIPKſECPEGYCTTCPV HWTVJGTinvestigation.#EMPQYNGFIOGPVU.—We thank the staff of E. 2ő6QOŒ5CY[GT5VCVG2CTMHQTHCEKNKVCVKPIVJKUresearch. We also thank the anonymous reviewers who provided comments and suggestions on this manuscript. This observation was made during a study of the natural history and ecology of ' NWEKHWIC in Sauerkraut Cave and was approved by the Kentucky Department and Fish and Wildlife (permit # SC1611045) and the Kentucky Department of Parks (permit # 1412).  ReferencesAnthony, C. D., C. A. M. Hickerson, and M. D. Venesky. 2007. Responses of juvenile terrestrial salamanders to introduced (.KVJQDKWU HQTſECVWU) and native centipedes (5EQNQRQET[RVQRU UGZURKPQUWU). ,QWTPCN QH <QQNQI[  54–62.Brodie Jr., E. D., T. G. Dowdey, and C. D. Anthony. 1989. Salamander antipredator strategies against snake attack: biting by &GUOQIPCVJWU. *GTRGVQNQIKEC  167–171.Camp, C. D. and J. B. Jensen. 2007. Use of twilight zones of caves by plethodontid salamanders. %QRGKC  594–604.Crane, A. and A. Mathis. 2015. Observation of predation by a lycosid spider on a captive-reared salamander larva (#OD[UVQOCCPPWNCVWO). *GTRGVQNQI[0QVGU  455–457.Ficetola, G. F., R. Pennati, and R. Manenti. 2013. Spatial segregation among age classes in cave salamanders: habitat selection or social interactions? 2QRWNCVKQP'EQNQI[  217–226.Gall, S. B., C. D. Anthony, and J. A. Wicknick. 2003. Do behavioral interactions between salamanders and beetles indicate a guild relationship? #OGTKECP /KFNCPF0CVWTCNKUV  363–374.Gunzburger, M. S. and J. Travis. 2005. Critical literature review of the evidence for unpalatability of amphibian eggs and larvae. ,QWTPCNQH*GTRGVQNQI[  547–571.Hairston, N. G., Sr. 1987. %QOOWPKV[ 'EQNQI[ CPF5CNCOCPFGT )WKNFU Cambridge Studies in Ecology. Cambridge, UK. Cambridge University Press. 230 pp.Hickerson, C.-A. M., C. D. Anthony, and A. M. Figura. 2017. Behavioral interactions between terrestrial salamanders and spiders: competition or intraguild predation? 'VJQNQI['EQNQI['XQNWVKQP in press.0QXGNKPVGTCEVKQPDGVYGGPCRKUCWTKFURKFGTCPFEurycea lucifuga282Phyllomedusa - 16(2), December 2017Hickerson, C.-A. M., C. D. Anthony, and B. M. Walton.  +PVGTCEVKQPUCOQPI HQTGUVƀQQTIWKNFOGODGTU KPstructurally simple microhabitats. #OGTKECP /KFNCPF0CVWTCNKUV  30–42.Hickerson, C.-A. M., C. D. Anthony, and J. A. Wicknick. 2004. Behavioral interactions between salamanders and centipedes: competition in divergent taxa. $GJCXKQTCN'EQNQI[  679–686.Howard, R. S. 1985. 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An experimental assessment of invertebrate/vertebrate predation: the interaction between wolf spiders ()NCFKEQUCRWNEJTC) and terrestrial salamanders (#OD[UVQOC OCEWNCVWO). ,QWTPCN QH<QQNQI[  1–5.Smith, D. D. and L. C. Balch. 1985. 'WT[EGCNWEKHWIC (Cave Salamander). Behavior. *GTRGVQNQIKECN4GXKGY  77–78.'FKVQT ,CKOG$GTVQNWEK$TCFNG[CPF'CUQP

Novel interaction between a pisaurid spider (Araneae: Pisauridae) and an adult Eurycea lucifuga (Caudata: Plethodontidae)

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Novel interaction between a pisaurid spider (Araneae: Pisauridae) and an adult Eurycea lucifuga (Caudata: Plethodontidae)

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