Carl Anthony, Associate Professor
University Heights, OH 44118 USA

Academic Background
BA Biology - North Central College, 1987
PhD Environmental and Evolutionary Biology - University of Louisiana, Lafayette, 1995




Research Projects
Ecological separation in a polymorphic terrestrial salamander - When studying speciation, researchers commonly examine reproductive isolation in recently diverged populations. Polymorphic species provide an opportunity to examine the role of reproductive isolation in populations that may be in the process of divergence. We examined a polymorphic population of terrestrial salamanders (Plethodon cinereus) for evidence of sympatric ecological separation by color morphology. Recent studies have correlated temperature and climate with color morphology in this species, but no studies have looked at differences in diet or mate choice between color morphs. We used artificial cover objects to monitor salamander diet, mating preference and surface activity over a two year period. We detected differences in diet and mate preference between striped and unstriped color morphs. The diets of striped individuals were significantly more diverse and were made up of more profitable prey than the diets of unstriped salamanders. Opposite sex pairs tended to be made up of individuals of the same color morph and striped males were able to attract larger females than were unstriped males. We corroborate findings of earlier studies suggesting that the unstriped form is adapted to warmer conditions. Unstriped individuals were the first to withdraw from the forest floor as temperatures fell in late fall and they tended to be found at warmer temperatures than did striped salamanders. We conclude that the two color morphs exhibit some degree of ecological separation and tend to mate assortatively, but are unlikely to be undergoing divergence given the observed frequency of intermorph pairings.
Edge effects and intraguild predation in native and introduced centipedes: evidence from the field and from laboratory microcosms – Human alteration of habitat has increased the proportion of forest edge in areas of previously continuous forest. This edge habitat facilitates invasion of exotic species into remaining fragments. The ability of native species to resist invasion varies and may depend on intrinsic variables such as dispersal and reproductive rates as well as external factors such as rate of habitat change and the density of populations of introduced species in edge habitat. We examined the distributional and competitive relationships of two members of the class Chilopoda, Scolopocryptops sexspinosus, a centipede native to the eastern US, and Lithobius forficatus, an exotic centipede introduced from Europe. We found that L. forficatus was most abundant in edge habitat and S. sexspinosus was most abundant in the interior habitat at our field sites. Although L. forficatus was present in habitat interiors at 11 of 12 sites, there was no correlation between fragment size and numbers of L. forficatus in interior habitat. The native centipede was rarely found occupying fragment edges. We used laboratory microcosms to examine potential competitive interactions and to indirectly assess prey preferences of the two species. In microcosms both species consumed similar prey, but the native centipede, S. sexspinosus, acted as an intraguild predator on the introduced centipede. Native centipedes were competitively superior in both intraspecific and interspecific pairings. Our results suggest that intraguild predation may aid native centipedes in resisting invasion of introduced centipedes from edge habitat.
Ecological interactions across diverse taxa I: interactions between salamanders and predatory beetles – Guilds consist of groups of species that use mutual resources or collections of overlapping resources. These groups of species are often members of the same family or genus, but they can span larger taxonomic categories. Predatory arthropods, such as carabid beetles, centipedes and spiders, and small carnivorous vertebrates, such as woodland salamanders, overlap extensively in microhabitat and prey types. These types of species, though unrelated, may form important guilds of forest floor predators. Stephanie Gall and I examined the behavioral interactions between a carabid ground beetle (Platynus tenuicollis) and a small woodland salamander (the red-backed salamander, Plethodon cinereus) in laboratory encounters. These species occur syntopically and are thought to feed on similar prey species. Individuals of P. cinereus are known to be territorial and to defend cover objects against conspecific and congeneric intruders. Because both species require moist conditions, it seems probable that they might compete for resources, such as cover objects and their associated prey, as the forest floor and leaf litter dries. We posed two general hypotheses in this study: (1) that salamanders would display territorial behavior toward intruding ground beetles and (2) that ground beetles might display similar types of territorial behaviors as those described previously for P. cinereus. In all treatments, residency status had an effect on behavior. Resident beetles and salamanders behaved more aggressively than did intruders. When paired with beetles, resident salamanders exhibited territorial behavior similar to that reported in other studies, but no biting was observed. Resident beetles exhibited similar levels of aggressive behavior across treatments. When physical attacks by beetles were observed, they were brief and often escalated to biting and chasing. Platynus tenuicollis bit intruders of both species in a significant number of trials and this often resulted in the production of adhesive secretions by P. cinereus which successfully immobilized P. tenuicollis. Anti-predatory behaviors observed in P. cinereus and biting by P. tenuicollis of P. cinereus, suggest that intraguild predation may be as important as interspecific resource competition in these species of forest floor predators.
Ecological interactions across diverse taxa II: interactions between salamanders and centipedes – Red-backed salamanders, Plethodon cinereus, use territorial advertisement in the form of agonistic displays and pheromonal scent marking as a mechanism for intraspecific interference competition. Although ecological and behavioral interactions among species of salamanders have been well studied, little is known about the interactions between territorial P. cinereus and other ecologically similar species, such as large predatory invertebrates. Our field data indicate that P. cinereus and a large syntopic centipede, Scolopocryptops sexspinosus, exhibit negative spatial associations in natural habitats, possibly indicating interspecific territoriality. Only seven instances of salamander/ centipede co-occurrence were recorded from a field sample of 247 occupied cover objects. Cover object size was positively correlated with salamander SVL (tip of the snout to the anterior end of the cloaca), but there was no correlation of cover object size to centipede length. Data on the ability of P. cinereus to differentiate among chemicals on the substrate suggest that visual cues are not necessary to elicit a territorial response from intruding salamanders. Although in laboratory trials salamanders behaved similarly toward intruders of both species, biting was directed only toward centipedes. Salamanders spent significantly more time approaching centipedes than they did approaching other salamanders. Approach behavior was often associated with nose tapping and may be an investigative, rather than aggressive, behavior. We suggest that territorial P. cinereus respond similarly to intruding salamanders and centipedes, but that they escalate more readily to biting centipedes because S. sexspinosus is sightless and thus unable to respond to visual signals.
Filial cannibalism (cannibalism of offspring by parents) – Recent studies have called into question the role of Wright’s coefficient of relatedness (r) in the interactions among relatives. Kin selection theory predicts a positive relationship between relatedness and frequency of “altruistic” acts, but a number of researchers have reported the opposite relationship. I used a Lycosid spider (Pardosa milvina) to test the hypothesis that genetic relatedness would affect the propensity of a cannibalistic species to prey on genetic relatives. I considered lack of predation to be a form of “altruism” where the predator incurs a cost (loss of a meal) that benefits potential prey. Specifically, I questioned whether direct genetic offspring would be avoided as prey items and whether sex or reproductive condition of a cannibalistic predator would affect the likelihood of predation on conspecific juveniles. As predicted by kin selection theory, spiderling mothers ate significantly fewer of their own offspring than they did of non-kin spiderlings of the same age. Adult virgin female and adult male spiders ate significantly more spiders than females that had recently carried spiderlings. Females with egg sacs consumed significantly fewer spiderlings than did virgin female spiders. These findings support Hamilton’s rule and suggest that in some systems, genetic relatedness plays a strong role in governing altruistic behavior toward relatives.
Aggressive behavior and territoriality – In terrestrial plethodontid salamanders, aggressive behavior is thought to function in the spacing of territorial residents among contested cover objects on the forest floor. Such behavior, when exhibited toward heterospecifics, plays an important role in the competitive interactions between species. I compared levels of aggressive in intra- and interspecific contexts in two species of sympatric salamanders (Plethodon albagula and P. ouachitae) that have similar ecological requirements but differ in adult size. I also tested the effectiveness of such behavior in holding cover objects (territorial foci) in the laboratory and on the forest floor. We predicted that if one species were more aggressive than the other, then that species would have greater success in obtaining and holding cover objects. In laboratory trials, residents of P. ouachitae (the smaller species) were extremely aggressive in both intra and interspecific contexts. Individuals of P. ouachitae delivered bites at a rate 14 times that of previously studied species of Plethodon and were significantly more likely to escalate to biting when tested as territorial residents (in intra- and interspecific trials) and as intruders (in intraspecific trials). Plethodon albagula exhibited a lower level of aggression, similar to other species of Plethodon studied previously. In laboratory trials, in which salamanders competed for cover objects of differing quality, residents of P. ouachitae were effective in expelling conspecific intruders and they were somewhat effective at expelling larger intruding P. albagula. Residents of P. albagula were less effective at expelling conspecific intruders and did not expel intruding P. ouachitae. The extreme aggression exhibited by P. ouachitae enabled it to expel intruders from artificial cover objects and to invade cover objects held by larger heterospecific residents. Field data supported intraspecific defence of cover objects by P. ouachitae, but results for P. albagula were inconclusive. These results are consistent with the geographic distributions of these species (P. ouachitae typically outnumbers P. albagula in the Ouachita Mountains) and provide an example of a behavioral mechanism overcoming a size-related disadvantage.


Collaborators

Cari-Ann M. Hickerson – Cleveland State University
B. Michael Walton – Cleveland State University
Dean C. Adams – Iowa State University

Graduate Students

Megan Acord - Mate choice and polymorphism in the red-backed salamander, Plethodon cinereus

Marcie Reiter - Territorial behavior in polymorphic red-backed salamanders

George Meszaros - Niche expansion in depauperate terrestrial salamander communities

Angela Stuzcka - The influence of diet on territorial behavior of a polymorphic salamander

Kim Thompson – Competitive interactions between red-backed and two-lined salamanders

Past Students

Anna Figura – Interactions between salamanders and forest floor spiders

Matthew Venesky – Antipredator responses of striped and un-striped red-backed salamanders

Rebecca Forrest Pinder – Stream health, parasitism, and reproductive condition of streamside salamanders

Cari Hickerson – Behavioral interactions between centipedes and woodland salamanders

Stephanie Gall – Competitive interactions between red-backed salamanders and carabid beetles

Jordan Schaul – Parasitism in hybrid salamanders relative to parental genotypes

Jaret Reblin – Effects of pesticide exposure on the development of chorus frog larvae

Lori Waltonen – Predatory strategies of ringneck snakes

Publications (please email me - canthony@jcu.edu for pdf files of papers)
Anthony, C.D., Venesky, M.D., and C.M. Hickerson. 2008. Ecological separation in a polymorphic terrestrial salamander. Journal of Animal Ecology 77:646-653.

Venesky, M.D. and C.D. Anthony. 2007. Antipredator adaptations and predator avoidance for two color morphs of the Eastern Red-backed Salamander, Plethodon cinereus. Herpetologica 63:450-458.

Anthony, C.D., Hickerson C.M., and M. D. Venesky. 2007. Responses of juvenile terrestrial salamanders to introduced (Lithobius forficatus) and native centipedes (Scolopocryptops sexspinosus). Journal of Zoology 271:54-62.

Hickerson, C.M., C.D. Anthony, and B.M. Walton. 2005. Edge effects and intraguild predation in native and introduced centipedes: evidence from the field and from laboratory microcosms. Oecologia 146:100-119.

Wicknick, J.A., C.D. Anthony, and J.S. Reblin. 2005. An amphibian survey of Killbuck Marsh Wildlife Area. Ohio Journal of Science 105:2-7.

Hickerson, C.M., C.D. Anthony, and J.A. Wicknick. 2004. Behavioral interactions between salamanders and centipedes: competition in divergent taxa. Behavioral Ecology 15:679-686.

Gall, S.B., C.D. Anthony, and J.A. Wicknick. 2003. Behavioral interactions between salamanders and beetles indicate a guild relationship. American Midland Naturalist 149:363-374.

Anthony, C.D. 2003. Kinship influences cannibalism in the wolf spider, Pardosa milvina. Journal of Insect Behavior 16:23-36.

Anthony, C.D., J.A. Wicknick, and R.G. Jaeger. 2002. Site tenacity and homing in the Rich Mountain salamander, Plethodon ouachitae (Caudata: Plethodontidae). The Southwestern Naturalist 47:401-408.

Anthony, C.D., J.A. Wicknick, and R.G. Jaeger. 1997. Social interactions in two sympatric salamanders: effectiveness of a highly aggressive strategy. Behaviour 134:71-88.

Mitchell, J.C., J.A. Wicknick, and C.D. Anthony. 1996. Do timber harvesting practices affect Peaks of Otter salamander (Plethodon hubrichti) populations? Amphibian and Reptile Conservation 1:15-19.

Adams, D.C. and C.D. Anthony. 1996. Using randomization techniques to analyze behavioural data. Animal Behaviour 51:733-738.

Jaeger, R.G., J.A. Wicknick, M.R. Griffis, and C.D. Anthony. 1995. Sociobiology of a terrestrial salamander: juveniles enter adult territories during stressful foraging periods. Ecology 76:533-543.

Anthony, C.D., J.R. Mendelson, III, and R.R. Simons. 1994. Differential parasitism by sex on plethodontid salamanders and histological evidence for structural damage to the nasolabial groove. American Midland Naturalist 132:302-307.

Williams, T.A. and C.D. Anthony. 1994. A technique to isolate salamander granular gland products with a comment on the evolution of adhesiveness. Copeia 1994:540-541.

Anthony, C.D. 1993. Recognition of conspecific odors by Plethodon caddoensis and P. ouachitae. Copeia 1993:1028-1033.

Anthony, C.D. and J.A. Wicknick. 1993. Aggressive interactions and chemical communication between adult and juvenile salamanders. Journal of Herpetology 27:261-264.

Anthony, C.D., D.R. Formanowicz, Jr., and E.D. Brodie, Jr. 1992. The effect of prey availability on the search behavior of two species of Chinese salamanders. Herpetologica 48:287-292.

Anthony, C.D., D.R. Formanowicz, Jr., and E.D. Brodie, Jr. 1991. Effect of mine reclamation on ant community structure in eastern Texas. The Texas Journal of Science 43:261-272.

Ducey, P.K., C.D. Anthony, and E.D. Brodie, Jr. 1991. Thresholds and escalation of antipredator responses in the Chinese salamander Cynops cyanurus: inter- and intra-individual variation. Behavioral Processes 23:181-191.

Brodie, E.D., Jr., T.G. Dowdy, and C.D. Anthony. 1989. Salamander antipredator strategies against snake attack: biting by Desmognathus. Herpetologica 45:167-171.

Species Accounts, Geographic Distribution, and Life History Notes
Anthony, C.D, M.D. Venesky, and J. Spetz. 2007. Life History Note: Eurycea longicauda longicauda. Predation. Herpetological Review 38:175-176.

Anthony, C.D. 2005a. Plethodon albagula, Western Slimy Salamander. Pp. 788-789. In Lannoo, M. J. (Ed.), Amphibian Declines: The Conservation Status of United States Species. University of California Press, Berkeley, California.

Anthony, C.D. 2005b. Plethodon caddoensis, Caddo Mountain Salamander. Pp. 792-793. In Lannoo, M. J. (Ed.), Amphibian Declines: The Conservation Status of United States Species. University of California Press, Berkeley, California.

Anthony, C.D. 2005c. Plethodon fourchensis, Fourche Mountain Salamander. Pp. 808. In Lannoo, M. J. (Ed.), Amphibian Declines: The Conservation Status of United States Species. University of California Press, Berkeley, California.

Anthony, C.D. 2005d. Plethodon kiamichi, Kiamichi Slimy Salamander. Pp. 819-820. In Lannoo, M. J. (Ed.), Amphibian Declines: The Conservation Status of United States Species. University of California Press, Berkeley, California.

Anthony, C.D. 2005e. Plethodon kisatchie, Louisiana Slimy Salamander. Pp. 820-821. In Lannoo, M. J. (Ed.), Amphibian Declines: The Conservation Status of United States Species. University of California Press, Berkeley, California.

Anthony, C.D. 2005f. Plethodon ouachitae, Rich Mountain Salamander. Pp. 831-833. In Lannoo, M. J. (Ed.), Amphibian Declines: The Conservation Status of United States Species. University of California Press, Berkeley, California.

Anthony, C.D. 2005g. Plethodon sequoyah, Sequoyah Slimy Salamander. Pp. 838. In Lannoo, M. J. (Ed Amphibian Declines: The Conservation Status of United States Species. University of California Press, Berkeley, California.

Reblin, J. and C.D. Anthony. 2001. Life History Note: Eurycea longicauda longicauda. Predation. Herpetological Review 32:245-246.

Anthony, C.D. and J.A. Wicknick. 1993. Geographic distribution. Plethodon ouachitae. Herpetological Review 24:152.

Teaching
BL 222 – General Ecology
BL 370 – Evolution
BL 444 – Advanced Ecology
BL 540 – Behavior
BL 521 – Herpetology

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