Huw Lloyd

Wildlife ecology and conservation (birds, bats)

Passive acoustic monitoring of wildlife populations

Fitness consequences of animal decision-making strategies (animal personalities)

Factors influencing conservation translocation outcomes

My research and engagement is focused on generating new scientific evidence on the ecology and behavior of animal species, their populations, and communities. I conduct research primarily through building international research collaborations (e.g., in Peru, Brazil, Bolivia, China, Tonga and the UK). The primary focus of my wildlife ecology and conservation science research is to understand the distribution and abundance of living organisms, and the environmental factors that influence their lives. Much of this research examines the responses of populations and communities to habitat change, using bird and mammal communities from both temperate and tropical forests, and wetland landscapes as model systems. My work encompasses both wider landscape responses by communities and populations, and a relatively narrow disciplinary focus on single species.

Wildlife Ecology and Conservation

Ornithology

Behavioral Ecology

Conservation Biology

Xiao-Yuan Wu, Huw Lloyd, Lu Dong, Yan-Yun Zhang, & Nan Lyu. (2024). Plasticity and the

structural characteristics of personality traits in captive-reared Japanese quail during ontogeny. 

Behavioural Ecology and Sociobiology 

Wu, Xiao-Yuan; Lloyd, Huw; Dong, Lu; Zhang, Yanyyun; Lyu, Nan. (2024). Moving beyond defining

animal personality traits: the importance of behavioural correlations and plasticity for conservation

translocations. Global Ecology and Conservation. 49: e02784.

P. Liu., Y. Lou., LIoyd, H., Y-H, Sun. (2023). Multiple field study approaches reveal the functions of

female song and duets in a temperate songbird. Animal Behaviour 206: 1-11

Y. Lou^., Q. Zhao., Y. Hu., L. Chen., P. Liu., Y. Fang., H. Lloyd, & Y-H. Sun. (2023). Personality

-dependent nest site selection and nest success during incubation in a wild population of chestnut

 thrush Turdusrubrocanus. iScience. https://doi.org/10.1016/j.isci.2023.107419  

Bai, Y., Liu, Y., Lei, W., Que, P., Liu, Y., Pagani-Nunez, E., Li, J., Lloyd, H., Zhang, ZZ. & Li, D. (2023).

Mammalian predators and vegetated nesting habitat reduce the breeding success of Kentish plovers

within a protected area: a case study in the Yellow Sea. Ecology and Evolution 13:e9884.

Pengfei Liu, Nan Lyu, Huw LIoyd, Yuehua Sun. (2023). Soft-song rather than broadcast song best

predicts physical attack in plain laughingthrush. Ibis 165 (2): 561-570.

Lloyd. H. (2023). Avian Ecology and Diversity, Population Monitoring and Conservation: Introduction

to the Special Issue. Diversity.

https://www.mdpi.com/journal/diversity/special_issues/Avian_EcoDiversity

M. G. Delaunay, C. Brassey, C. Larsen, H. Lloyd, & R. A. Grant. (2022). The evolutionary origin of avian

rictal bristles and their likely role in feeding ecology. Scientific Reports. 12:21108.

Nan Lyu, Justin Yeh, Huw Lloyd and Sun Yue-Hua. (2022). The influence of male mate choice on the

evolution of male preferences and female traits. Communications Biology 5:4 

Nan Lyu, Yunbiao Hu, Jiahua Zhang, Huw Lloyd, Yue-Hua Sun and Yi Tao. (2021). Switching costs in

stochastic environments drive the emergence of matching behaviour in animal decision‑making

through the promotion of reward learning strategies. Scientific Reports 11:23593     

Chen Yang, Jie Wang, Nan Lyu, & Huw Lloyd. (2021). Comparison of digital photography and

spectrometry for evaluating colour perception in humans and other trichomatic species. Behavioural

Ecology and Sociobiology 75: 151

Zhang, J., Zhang, Y., Lloyd, H., Zhang, Z. & Li, D. (2021). Rapid reclamation and degradation of Suaeda

salsa saltmarsh along coastal China’s northern Yellow Sea. Land. 10, 835.

Zhang, j., Zhang, H., Liu, Y., Lloyd, H., Zhang, Z. & Donglai Li. (2021). Saltmarsh vegetation and social 

environment influence flexible seasonal vigilance strategies for two sympatric migratory curlew species

in adjacent coastal habitats. Avian Research 12:39.

Bedson, CPE., Thomas, L., Wheeler, PM., Reid, N., Harris, WE., Lloyd, H., Mallon, D. & Preziosi, R.

(2021). Estimating density of mountain hares using distance sampling: a comparison of daylight visual 

surveys, night-time thermal imaging and camera traps. Wildlife Biology.

Mahamued, BA., Donald, PF., Collar, NJ., Marsden, SJ., Ndang’ang’a, PK., Wondafrash, M., Dellengen

Abebe, Y., Bennett, J., Wotton, SR., Gornall, D. & Lloyd, H. (2021). Rangeland loss and population

decline of the critically endangered Liben Lark Heteromirafra archeri in southern Ethiopia. Bird 

Conservation International. pp.1-14. 

F. Wu, W. Lei, H. Lloyd, Z. Zhang (2020). Predictors of Gull-billed tern (Gelochelidon nilotica) nest

survival in artificial coastal saltpans, Bohai Bay, China. PeerJ. 8, pp.e10054-e10054.

D. Li, J. Zhang, Y. Liu, H. Lloyd, E. Pagani-Núñez, et al. (2020). Differences in dietary specialization, 

habitat use and susceptibility to human disturbance influence feeding rates and resource partitioning 

between two migratory Numenius curlew species. Estuarine, Coastal and Shelf Science. 245, 

pp.106990-106990.

D. Li, J. Zhang, L. Chen, H. Lloyd, Z. Zhang (2020). Burrow ambient temperature influences Helice crab

activity and availability for migratory Red‐crowned cranes Grus japonensis. Ecology and Evolution. 10

(20), pp.11523-11534.

MG. Delaunay, C. Larsen, H. Lloyd, M. Sullivan, RA. Grant (2020). Anatomy of avian rictal bristles in

Caprimulgiformes reveals reduced tactile function in open-habitat, partially diurnal foraging 

species. Journal of Anatomy 272 (2): 355-366.

Mendez, D., Marsden, S. J. & Lloyd, H. (2019). Assessing population size and structure of Andean

Condor Vultur gryphus in Bolivia using a photographic ‘capture-recapture’ method. Ibis 161 (4): 867-

877.

Xia, C., Deng, Z., Lloyd, H., Møller, A.P., Zhao, X., & Zhang, Y.Y. (2019). The function of three main call types in common cuckoo. Ethology 125(9): 652-659.

Deng, Z., Lloyd, H., Xia, C., Li, D. & Zhang, Y.Y. (2019). Within-season decline in call consistency of individual male Common Cuckoos (Cuculus canorus). Journal of Ornithology 160(2): 317-327.

Deng, Z., Lloyd, H., Xia, C., Møller, A.P., Liang, W. & Zhang, Y.Y. (2019). Components of variation in female common cuckoo calls. Behavioural Processes 158: 106-112.

Xia, C., Lloyd, H., Shi, J, Wei, C. & Zhang, Y. (2018). Dawn singing of the Brownish-Flanked Bush Warbler influences dawn chorusing of bird communities. Ethology 124 (6):400-409.

Li, D., Yu L., Lloyd, H., Shuyu, Z., Shuyan, Z, Dongmei, W. & Zhang, Z. (2017). Habitat-dependent changes in Red-crowned Crane vigilance behaviour influenced by wildlife tourism. Scientific Reports 7: 16614.

Nan, L., Servedio, M. R., Lloyd, H. & Sun, Y-H. (2017). The evolution of post-pairing male mate choice. Evolution71(6): 1465-1477. 

Li, Y., Xia, C., Lloyd, H., Li, D. & Zhang, Y. (2017). Identification of vocal individuality in male cuckoos using different analytical techniques.Avian Research. 8 (1).

Nan, Y., Moermond, T. C., Lloyd, H. Yu, X., Dou, L., Zhang, K., Bisong, Y. & Jianghong, R. (2016). Effects of supplementary feeding on the breeding ecology of Buff-throated Partridge at a Tibetan Sacred Site. 

PLoS ONE doi:10.1371/journal.pone.0146568 

Liu, B., Li, L., Lloyd, H., Xia, C. & Zhang, Y.Y. (2016). Comparing post-release survival and habitat use by captive-bred Cabot’s Tragopan (Tragopan caboti) in an experimental test of soft-release reintroduction strategies. Avian Research 7, 19.

Nan, L., Lloyd, H. & Yue-Hua, S (2015). The evolution of delayed plumage maturation in birds and the significance of condition-dependent parental care. Behavioural Ecology and Sociobiology 69 (6): 1003-1010

Li, D., Sun, X., Lloyd, H., Wei, H., Liu, Y. & Zhang, Z. (2015). Reed Parrotbill nest predation by Tidal mudflat crabs: evidence for an ecological trap? Ecosphere 6 (1): 20

Xia, C., Wang, P-D., Liu, B-Y., Lloyd, H. & Zhang, Y. Y. (2015). Reliability of the brownish-flanked bush warbler's soft song in male-male conflict. Avian Research 6 (6)

Li, D., Ding, Y., Yuan, Y., Lloyd, H. & Zhang, Z. (2014). Female tidal mudflat crabs represent a critical food resource for migratory Red-crowned Cranes in the Yellow River Delta, China. Bird Conservation International 24: 416-428.

Torres-Sovero, C., Lloyd, H. & Brightsmith, D.J. (2014). Uso de collpa y otras observaciones del Maracana Cabeciazul (Primolius couloni) en el Sureste del Peru. Ornitologìa Neotropical 25: 87-94

Xia, C., Wei, C., Lloyd, H., Wu, Q., Liu, J., & Zhang, Y. (2014). Dawn singing intensity of the male Brownish-flanked Bush Warbler: effects of territorial insertions and number of neighbours. Ethology 120:1-7

Li, D. L., Chen, S, H., Lloyd, H., Zhu, S. Y., Shan, K. & Zhang, Z. Z. (2013). A preliminary assessment of the importance of artificial habitats to migratory waterbirds within a novel wetland landscape, Yellow River Delta, China. Bird Conservation International 23 (2): 184-198

Xia, C., Lin, X., Liu, W., Lloyd, H. & Zhang, Y. (2012). Acoustic identification of individuals within large avian populations: A case study of the Brownish-flanked Bush Warbler, south-central China. PLoS ONE 7(8): e42528. doi:10.1371/journal.pone.0042528 

Nan, L., Chen-Xi, J., Lloyd, H. & Yue-Hua, S. (2012). Species-specific habitat fragmentation assessment, considering ecological niche requirements and dispersal capability. Biological Conservation 152: 102-109.

Nan, L., Yu, J., Lloyd, H. & Yue-Hua, S. (2012). Assessing the distribution and potential risks from climate change for the Sichuan Jay Perisoreus internigrans. The Condor 114 (2): 365-376. 

Lloyd, H., Sevillano Ríos, S., Marsden, S. J. & Valdez-Velásquez, A. (2012). Bird community composition and abundance across an Andean tree-line ecotone. Austral Ecology 37: 470-478

Zhou, Z., Sun, Y., Lu, D., Xia, C., Lloyd, H. & Zhang, Y. (2012). Breeding biology of Asian House Martin Delichon dasypus in a high-elevation area. Forktail 28: 1-6.

Bernardo, C. S. S., Lloyd, H., Olmos, F., Cancian, L. F. & Galetti, M. (2011). Using post-release monitoring data for optimizing avian reintroduction programs: a two-year case study from the Brazilian Atlantic Rainforest. Animal Conservation 14 (6):676-686

Bernardo, C. S. S., Lloyd, H., Baily, N.; & Galetti, M. (2011). Modelling post-release survival of reintroduced Red-billed Curassows Crax blumenbachii, Brazil. Ibis 153: 562-572

Lloyd, H. & Marsden S. J. (2011). Between-patch bird movements within a high-Andean Polylepis woodland/matrix landscape: implications for habitat restoration. Restoration Ecology 19 (1): 74-82.

Li, D., Chen, S., Guan, L., Lloyd, H., Liu Y., Juanzhang, L. & Zhang, Z. Z. (2011). Patterns of waterbird community composition across a natural and restored wetland landscape mosaic, Yellow River Delta, China. Estuarine, Coastal, and Shelf Science 91 (2): 325-332.

Bernardo, C. S. S., Creswell, B., Lloyd, H., Azeredo, R. & Simpson, J. (2011) Selection of radio-transmitter and post-release monitoring of reintroduced Red-billed Curassow Crax blumenbachii in Brazil. European Journal of Wildlife Research 57 (3): 689-694

Wei, M., Lloyd, H. & Zhang, Y. (2011). Neighbour-stranger discrimination by Yellow-bellied Tit Parus venustulus: evidence for the dear-enemy effect. Journal of Ornithology 152: 431-438.

Nan, Y., Kai, Z., Jianghong, R., Lloyd, H. Yu, X., Ying, W. & Bisong, Y. (2011). Group size does not influence territory size and overlap in a cooperative breeding gamebird. Ardea 99:199-206

Sevillano Ríos, S., Lloyd. H. & Valdez-Velásquez, A. (2011). Bird species richness, diversity and abundance in Polylepis woodlands, Huascaran Biosphere Reserve, Perú. Studies on Neotropical Fauna and Environment 46 (1): 69-76.

Dong, L., Niu, W., Zhou, Z., Yu-Cheng, H., Sun, Y., Lloyd, H. & Zhang, Y. & (2011). Assessing the genetic integrity of captive and wild populations for reintroduction programs: the case of Cabots Tragopan in China. Chinese Birds 2 (2): 65-71.

Kai, Z., Nan, Y., Yu, X., Jianghong, R., Lloyd, H. & Bisong, Y. (2011) Nesting behaviour of Szechenyi Monal-Partridge in tree-line habitats, China. Wilson Journal of Ornithology 123 (1): 93-96.

Lloyd, H. (2008). Influence of within-patch habitat quality on Polylepis bird abundance. Ibis 150: 735-745.

Lloyd, H. & Marsden, S. J. (2008). Bird community variation across Polylepis woodland fragments and matrix habitats: implications for conservation within a high Andean landscape. Biodiversity and Conservation 17: 2645-2660

Lloyd, H. (2008). Abundance and patterns of rarity of Polylepis birds in the Cordillera Vilcanota, southern Perú: implications for habitat management strategies. Bird Conservation International 18: 164-180. 

Lloyd, H. (2008). Foraging ecology of high-Andean insectivorous birds in remnant Polylepis forest patches Wilson Journal of Ornithology 120 (3): 531-544.

Lloyd, H. (2004). Habitat and population estimates of some threatened lowland forest bird species in Tambopata, south-east Peru. Bird Conservation International 14 (4): 261-277.

Lloyd, H. (2003). Population densities of some nocturnal raptors (Strigidae) in southeast Peru. Journal of Field Ornithology 74 (4): 376-380

Lloyd, H. (2000). Population densities of the Black-faced Cotinga Conioptilon mcilhennyi in southeast Perú. Bird Conservation International 10: 277-285.

Harrison, C.H., Lloyd, H. & Field, C. (2017). Evidence review of the impacts of solar farms on birds, bats and general ecology. Natural England, Manchester Metropolitan University.

Lloyd, H., Cahill, A. J., Jones, M. & Marsden, S. J. (1998) Estimating densities using distance sampling. In Bird surveys (Eds) C. J. Bibby, M. Jones & S. J. Marsden. Expedition Advisory Centre of the Royal Geographic Society and BirdLife International

Walker, B., Lloyd, H. & Cheshire, G. (2008). Peruvian Wildlife. Bradt Publishing, UK.