Jocelyn Behm’s Publications

Matthew Helmus’s Publications


  • Lewkiewicz, SM, B Seibold, MR Helmus. 2023. Quantifying population resistance to climatic variability: The invasive spotted lanternfly grape pest is buffered against temperature extremes in California. bioRxiv, 2023.12.21.572841.
  • Behm, JE, J Ellers, WAM Jesse, TJ Tran, MR Helmus. 2023. Predicting and quantifying coexistence outcomes between resident and invading species using trait and abundance data. bioRxiv. 2023.01.12.523647.
  • Gleditsch, J., JE Behm, MR Helmus. 2023. The great acceleration of island saturation by species introductions in the Anthropocene has altered species-area relationships. bioRxiv. 2023.01.10.523426.
  • Huron, NA, SB Hedges, & MR Helmus. 2022. Detecting stabilizing, directional, and disruptive patterns of anthropogenic species loss with general models of nonrandom extinction. bioRxiv.
  • Huron, NA & MR Helmus. 2022. Predicting host associations of the invasive spotted lanternfly on trees across the USA. bioRxiv.
  • Jesse, WAM, J Ellers, JE Behm, GC Costa, SB Hedges, MR Helmus. 2022. Elevated human footprint on islands promotes both introduction and extinction probability of insular reptiles at opposite ends of geographic, evolutionary and ecological continua. bioRxiv.


  • Belouard, N & JE Behm. 2023. Multiple paternity in the invasive spotted lanternfly (Hemiptera: Fulgoridae). Environmental Entomology, 1-7.
  • De Bona, S, L Barringer, P Kurtz, J Losiewicz, GR Parra, & MR Helmus. 2023. lydemapr: an R package to track the spread of the invasive Spotted Lanternfly (Lycorma delicatula, White 1845) (Hemiptera, Fulgoridae) in the United States. NeoBiota, 86:151-168.
  • Swartz, TM, JM Gleditsch, JE Behm. 2023. A functional trait approach reveals the effects of landscape context on ecosystem services provided by urban birds. Landscape and Urban Planning. 234
  • Gleditsch, JM, JE Behm, J Ellers, WAM Jesse, MR Helmus. 2023. Contemporizing island biogeography theory with anthropogenic drivers of species richness. Global Ecology and Biogeography. 32(2):233-249.
  • Ramirez, VA, S De Bona, MR Helmus & JE Behm. 2023. Multiscale assessment of oviposition habitat associations and implications for management in the spotted lanternfly (Lycorma delicatula), an emerging invasive pest. Journal of Applied Ecology.
  • Belouard, N & JE Behm. 2023. Spotted! Computer-aided individual photo-identification allows for mark-recapture of invasive spotted lanternfly (Lycorma delicatula). Frontiers in Insect Science. 3:9  
  • Liu, S, Y Meng, JE Behm, W Zhang, S Fu. 2023. Canopy nitrogen addition affects ground-dwelling spider assemblages and trophic position. Insect Conservation and Diversity. 16:10-121.


  • Turner, DB, JE Behm, PM Phillips, VA Ramirez, MR Helmus. 2022. Coding for broader impact: leveraging coding skills for stakeholder communication. Frontiers in Ecology and the Environment.
  • Behm, JE, N Bélouard, JM Gleditsch, PM Phillips, TM Swartz. 2022. Trait-based approaches for understanding how biodiversity generates sustainable benefits in urban vegetated green infrastructure. Current Opinion in Environmental Sustainability 57:1-11.
  • Huron, NA, JE Behm, MR Helmus. 2022. Paninvasion severity assessment of a US grape pest to disrupt the global wine market. Communications Biology 5, 655 .
  • Behm, JE, GM Busala, MR Helmus. 2022. First records of three new lizard species and a range expansion of a fourth lizard species introduced to Aruba. BioInvasions Records 11(1): 296–306.
  • Lewkiewicz, SM, S De Bona, MR Helmus, B Seibold. 2022. Temperature sensitivity of pest reproductive numbers in age-structured PDE models, with a focus on the invasive spotted lanternfly. Journal of Mathematical Biology. 85(3): 1-37
  • Gutierrez-Velez, VH, MR Gilbert, D Kinsey, JE Behm. 2022. Beyond the ‘urban’ and the ‘rural’: conceptualizing a new generation of infrastructure systems to enable rural–urban sustainability. Current Opinion in Environmental Sustainability. 56, 101177
  • Liu, S, X Shangwe, D Wu, JE Behm, Y Meng, H Yuan, P Wen, AC Hughes, X Yang. 2022. Understanding global and regional patterns of termite diversity and regional functional traits. iScience. 2022.105538
  • Liu, S, Y Meng, JE Behm, W Zhang, S Fu. 2022. Nitrogen addition enhances the bottom-up effects in the detrital food web. Global Ecology and Conservation. 39:e02299.
  • Cavender-Bares, JM, E Nelson, JE Meireles, JR Lasky, DA Miteva, DJ Nowak, WD Pearse, MR Helmus, AE Zanne, WF Fagan, C Mihiar, NZ Muller, NJB Kraft, S Polasky. 2022. The hidden value of trees: Quantifying the ecosystem services of tree lineages and their major threats across the contiguous US. PLOS Sustainability and Transformation. 1(4).
  • Jarzyna, MA, KEA Norman, JM LaMontagne, MR Helmus, D Li, SM Parker, M Perez Rocha, S Record, ER Sokol, PL Zarnetske, TD Surasinghe. 2022. Community Stability is Related to Animal Diversity Change. Ecosphere. 13( 3): e3970.
  • Li, D, S Record, ER Sokol, ME Bitters, MY Chen, YA Chung, Y. Anny, MR Helmus, et al. 2022. Standardized NEON Organismal Data for Biodiversity Research. Ecosphere137): e4141.


  • Nagy, RCBalch, JKBissell, EKCattau, … MR Helmus, et. al. 2021Harnessing the NEON data revolution to advance open environmental science with a diverse and data-capable communityEcosphere 1212):e03833.
  • Prinzing, A, S Pavoine, H Jactel, … MR Helmus, et al. (2021), Disturbed habitats locally reduce the signal of deep evolutionary history in functional traits of plants. New Phytologist, 232: 1849-1862.
  • Meng, Y., H Chen, JE Behm, S Xia, B Wang, S Liu, and X Yang2021Effects of different tea plantation management systems on arthropod assemblages and network structureEcosphere 127):e03677.
  • Pearsall, H., VH Gutierrez-Velez, MR Gilbert, … JE Behm, et al. 2021. Advancing equitable health and well-being across urban–rural sustainable infrastructure systems. npj Urban Sustainability. 126 (2021).
  • Ordway, EM, AJ Elmore, S Kolstoe, … MR Helmus et al. 2021Leveraging the NEON Airborne Observation Platform for socio-environmental systems researchEcosphere, 126):e03640.
  • Liu, S., JE Behm, S Wan, J Yan, Q Ye, W Zhang, X Yang, S Fu. 2021. Effects of canopy nitrogen addition on soil fauna and litter decomposition rate in a temperate forest and a subtropical forest. Geoderma. 


  • Li, D, R Dinnage, LA Nell, MR Helmus, AR Ives. 2020. phyr: An R package for phylogenetic species-distribution modelling in ecological communities. Methods in Ecology and Evolution, 00:1-9.
  • Jesse, WAM,  J Molleman, O Franken, M Lammers, MP Berg,  JE Behm, MR Helmus, J Ellers. 2020. Disentangling the effects of plant species invasion and urban development on arthropod community compositionGlobal Change Biology, 26 (6), 3294-3306.
  • Behm, JE. 2020. Is biodiversity needed for sustainability? A spotlight on urban landscapes. American Journal of Botany, 107 (5), 703-706. 
  • Perella, CD and JE Behm. 2020. Understanding the spread and impact of exotic geckos in the greater Caribbean region. Biodiversity and Conservation, 29, 1109–1134.
  • TJ Tran, MR Helmus, JE Behm. 2020. Green infrastructure space and traits (GIST) model: Integrating green infrastructure spatial placement and plant traits to maximize multifunctionality. Urban Forestry & Urban Greening, 49, 126635.


  • Assour, H.R. and JE Behm. 2019. First Occurrence of Cheilomenes sexmaculata (Fabricius)(Coleoptera: Coccinellidae) to the Caribbean Island of Curaçao. Neotropical Entomology. 48(5):863-865.
  • Helmus, MR 2019. Book Review: Phylogenetic Diversity Applications and Challenges in Biodiversity Science. Quarterly Review of Biology. 94(3):308-309.
  • Helmus, MR and JE Behm. 2019. Island Biogeography Revisited. In: Earth Systems and Environmental Sciences. Elsevier. 1-6. Book chapter.
  • Liu, S., X. Lin, JE Behm, H. Yuan, P. Stiblik, J. Šobotník, J. Gan, S. Xia, X. Yang. In Press. Comparative responses of termite functional and taxonomic diversity to land-use change. Ecological Entomology.
  • Rapacciuolo, G, CH Graham, J Marin, JE Behm, GC Costa, SB Hedges, MR Helmus, VC Radeloff, BE Young, TM Brooks. 2019. Species diversity as a surrogate for conservation of phylogenetic and functional diversity in terrestrial vertebrates across the Americas. Nature Ecology & Evolution 3:53–61.
  • Behm, JE, G van Buurt, BM DiMarco, J Ellers, CG Irian, KE Langhans, K McGrath, TJ Tran, MR Helmus. 2019. First records of the mourning gecko (Lepidodactylus lugubris Duméril and Bibron, 1836), common house gecko (Hemidactylus frenatus Schlegel, 1836), and Tokay gecko (Gecko gecko Linnaeus, 1758) on the island of Curaçao, Dutch Antilles, and remarks on their distributions in the Caribbean region. BioInvasion Records 8(1), 34-44


  • Behm, JE, BR Waite, ST Hsieh, MR Helmus. 2018. Benefits and limitations of three-dimensional printing technology for ecological research. BMC Ecology 18(32) 1:13 
  • Helmus, MR and JE Behm. 2018. Anthropocene Island Biogeography. In: Encyclopedia of the Anthropocene. (eds. D. DellaSala, M. Goldstein). Elsevier. pp 37-43. Book chapter. 
  • Jesse, WAM, JE Behm, MR Helmus, J Ellers. 2018. Human land use promotes the abundance and diversity of exotic species on Caribbean islands. Global Change Biology 24(10):4784-4796 
  • Liu, S, J Hu, JE Behm, X He, J Gan, X Yang. 2018. Nitrogen addition changes the trophic cascade effects of spiders on a detrital food web. Ecosphere 9(10):e02466 
  • Behm, JE, CF Behm, GM Behm, MC Behm, SP Behm, MR Helmus2018. Hemidactylus turcicus (Mediterranean Gecko) Geographic update. Herpetological Review 49(1): 73. 
  • Marin, J, G Rapacciuolo, GC Costa, CH Graham, TM Brooks, BE Young, VC Radeloff, JE Behm, MR Helmus, SB Hedges. 2018. Evolutionary time drives global tetrapod diversity. Proceedings of the Royal Society B 285 (1872), 1-8.


  • Rapacciuolo, G, J Marin, GC Costa, MR Helmus, JE Behm, TM Brooks, SB Hedges, VC Radeloff, BE Young, CH Graham. 2017. The signature of human pressure history on the biogeography of body mass in tetrapods. Global Ecology and Biogeography 26:1022-1034.
  • Tucker, CM, MW Cadotte, SB Carvalho, TJ Davies, S Ferrier, SA Fritz, R Grenyer, MR Helmus, LS Jin, AO Mooers, S Pavoine, O Purschke, DW Redding, DF Rosauer, M Winter, F Mazel. 2017. A guide to phylogenetic metrics for conservation, community ecology and macroecologyBiological Reviews 92 (2), 698-715. 
  • Helmus, MR, JE Behm, WAM Jesse, JJ Kolbe, J Ellers, JB Losos. 2017. Exotics exhibit more evolutionary history than natives: A comparison of the ecology and evolution of exotic and native anole lizardsInvasion Genetics: The Baker and Stebbins Legacy, 122-138. 


  • Nelson, EJ, MR Helmus, J Cavender-Bares, S Polasky, JR Lasky, AE Zanne, WD Pearse, NJB Kraft, DA Miteva, WF Fagan. 2016. Commercial Plant Production and Consumption Still Follow the Latitudinal Gradient in Species Diversity despite Economic GlobalizationPloS One 11 (10), e0163002. 
  • Liu, S, JE Behm, J Chen, S Fu, X He, J Hu, D Schaefer, J Gan, X Yang. 2016. Functional redundancy dampens the trophic cascade effect of a web-building spider in a tropical forest floorSoil Biology and Biochemistry 98, 22-29.
  • Yguel, B, H Jactel, IS Pearse, D Moen, M Winter, J Hortal, MR Helmus, I Kühn, S Pavoine, O Purschke, E Weiher, C Violle, W Ozinga, M Brändle, I Bartish, A Prinzing. 2016. The evolutionary legacy of diversification predicts ecosystem functionThe American Naturalist 188 (4), 398-410. 
  • Behm, JE and MR Helmus. 2016. Microhyla butleri (Tubercled Pygmy frog) predation. Herpetological Review, 47(4): 643-644. 


  • Jamshidi, S, JE Behm, D Eveillard, ET Kiers, P Vandenkoornhuyse. 2015. Using hybrid automata modelling to study phenotypic plasticity and allocation strategies in the plant mycorrhizal mutualismEcological Modelling 311, 11-19. 
  • Pearse, WD, MW Cadotte, J Cavender-Bares, AR Ives, CM Tucker, SC Walker, MR Helmus. 2015. pez: phylogenetics for the environmental sciencesBioinformatics 31 (17), 2888-2890. 
  • Vermaat, JE, J Ellers, MR Helmus. 2015. The role of biodiversity in the provision of ecosystem servicesEcosystem Services: From Concept to Practice, 25. Email for pdf.


  • Helmus, MR, DL Mahler, JB Losos. 2014. Island biogeography of the AnthropoceneNature 513 (7519), 543-546. 
  • Behm, JE, R Geurts, ET Kiers. 2014. Parasponia: a novel system for studying mutualism stabilityTrends in Plant Science 19 (12), 757-763. 
  • Behm, JE, ET Kiers. 2014. A phenotypic plasticity framework for assessing intraspecific variation in arbuscular mycorrhizal fungal traitsJournal of Ecology 102 (2), 315-327. 
  • Engelmoer, DJP, JE Behm, ET Kiers. 2014. Intense competition between arbuscular mycorrhizal mutualists in an in vitro root microbiome negatively affects total fungal abundanceMolecular Ecology 23 (6), 1584-1593. 
  • Vasseur, DA, JW Fox, A Gonzalez, R Adrian, BE Beisner, MR Helmus, C Johnson, P Kratina, C Kremer, C de Mazancourt, E Miller, WA Nelson, M Paterson, JA Rusak, JB Shurin, CF Steiner. 2014. Synchronous dynamics of zooplankton competitors prevail in temperate lake ecosystemsProc. R. Soc. B 281 (1788). 
  • Pearse, WD, A Purvis, J Cavender-Bares, MR Helmus. 2014. Metrics and Models of Community PhylogeneticsModern Phylogenetic Comparative Methods and Their Application in Evolutionary Biology. Springer Berlin Heidelberg, pp. 451-464. 


  • Blanchet S, MR Helmus, S Brosse, G Grenouillet. 2013. Regional vs local drivers of phylogenetic and species diversity in stream fish communitiesFreshwater Biology 59 (3), 450-462. 
  • Behm, JE, X Yang, J Chen. 2013. Slipping through the cracks: rubber plantation is unsuitable breeding habitat for frogs in Xishuangbanna, ChinaPloS One 8 (9), e73688. 
  • Behm, JE, DA Edmonds, JP Harmon, AR Ives. 2013. Multilevel statistical models and the analysis of experimental dataEcology 94 (7), 1479-1486. 
  • Helmus, MR, N Mercado-Silva, MJ Vander Zanden. 2013. Subsidies to predators, apparent competition and the phylogenetic structure of prey communitiesOecologia 173 (3), 997-1007. 
  • Behm, JE, WKV Fu. 2013. Field guide to the frogs of the Xishuangbanna Tropical Botanic Garden. Xishuangbanna Tropical Botanic Garden Education Center Publication. Book description (in Chinese).


  • Kua, CS, J Ruan, J Harting, CX Ye, MR Helmus, J Yu, CH Cannon. 2012. Reference-Free Comparative Genomics of 174 Chloroplasts. PloS One 7 (11), e48995. 
  • Helmus, MR, AR Ives. 2012. Phylogenetic diversity–area curvesEcology 93:S1-S43. 
  • Behm, JE, BE Knaebe, VWK Fu, DA Edmonds. 2012. Polypedates leucomystax (Java Whipping Frog) communal nestingHerpetological Review 42 (4), 587-588.
  • Edmonds, DA, JE Behm. 2012. Raorchestes menglaensis Geographic UpdateHerpetological Review 43 (1), 125-126.
  • Edmonds, DA, JE Behm. 2012. Raorchestes menglaensis (Mengla small treefrog) colorationHerpetological Review 43 (1), 125-126.


  • Eklöf, A, MR Helmus, M Moore, S Allesina. 2011. Relevance of evolutionary history for food web structureProceedings of the Royal Society of London B: Biological Sciences 279 (1733), 1588-1596. 
  • Ives, AR, MR Helmus. 2011. Generalized linear mixed models for phylogenetic analyses of community structureEcological Monographs 81 (3), 511-525. 
  • Scheiner, SM, A Chiarucci, GA Fox, MR Helmus, DJ McGlinn, MR Willig. 2011. The underpinnings of the relationship of species richness with space and timeEcological Monographs 81 (2), 195-213. 
  • Quan, RC, G Ren, JE Behm, L Wang, Y Huang, Y Long, J Zhu. 2011. Why does Rhinopithecus bieti prefer the highest elevation range in winter? A test of the sunshine hypothesisPloS One 6 (9), e24449. 
  • Gilman, RT, JE Behm. 2011. Hybridization, species collapse, and species reemergence after disturbance to premating mechanisms of reproductive isolationEvolution 65 (9), 2592-2605. 


  • Ives, AR, MR Helmus. 2010. Phylogenetic metrics of community similarity. The American Naturalist 176, E128-E142. 
  • Kembel, SW, PD Cowan, MR Helmus, WK Cornwell, H Morlon, DD Ackerly, SP Blomberg, CO Webb. 2010. Picante: R tools for integrating phylogenies and ecologyBioinformatics 26 (11), 1463. 
  • Helmus, MR, WB Keller, MJ Paterson, ND Yan, CH Cannon, JA Rusak. 2010. Communities contain closely related species during ecosystem disturbanceEcology Letters 13 (2), 162-174. 
  • Behm, JE, AR Ives, JW Boughman. 2010. Breakdown in postmating isolation and the collapse of a species pair through hybridizationThe American Naturalist 175 (1), 11-26. 

2009 and earlier

  • Ahrenstorff, TD, GG Sass, MR Helmus. 2009. The influence of littoral zone coarse woody habitat on home range size, spatial distribution, and feeding ecology of largemouth bass (Micropterus salmoides)Hydrobiologia 623 (1), 223-233. 
  • Helmus, MR, LB Allen, O Dominguez-Dominguez, ED Pardo, P Gesundheit, J Lyons, N Mercado-Silva. 2009. Threatened fishes of the world: Allotoca goslinei Smith and Miller, 1987 (Goodeidae)Environmental Biology of Fishes 84 (2), 197-198. 
  • Mercado‐Silva, N, MR Helmus, M Zanden. 2008. The effects of impoundment and non‐native species on a river food web in Mexico’s central plateauRiver Research and Applications 25 (9), 1090-1108. 
  • Helmus, MR, GG Sass. 2008. The rapid effects of a whole‐lake reduction of coarse woody debris on fish and benthic macroinvertebratesFreshwater Biology 53 (7), 1423-1433. 
  • Newton, RJ, SE Jones, MR Helmus, KD McMahon. 2007. Phylogenetic ecology of the freshwater Actinobacteria acI lineageApplied and Environmental Microbiology 73 (22), 7169-7176. 
  • Helmus, MR, K Savage, MW Diebel, JT Maxted, AR Ives. 2007. Separating the determinants of phylogenetic community structureEcology Letters 10 (10), 917-925. 
  • Helmus, MR, TJ Bland, CK Williams, AR Ives. 2007. Phylogenetic measures of biodiversityThe American Naturalist 169 (3), E68-E83. 
  • Helmus, MR, DE Dussourd. 2005. Glues or poisons: which triggers vein cutting by monarch caterpillars? Chemoecology 15 (1), 45-49. 
  • Drake, DL, MA Hagerty, JE Behm, SJ Goldenberg, JR Spotila. 2003. Marine turtle nesting activity at Playa Naranjo Santa Rosa National Park, Costa Rica for the 1998-99 season. Chelonian Conservation and Biology 4 (3), 675-678.
  • Drake, DL, JE Behm, SJ Goldenberg, MA Hagerty. 2001. Lepidochelys olivacea (olive ridley sea turtle) predation. Herpetological Review 32, 104.


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