Environmental and Geographic Conditions on the Breeding Grounds Drive Bergmannian Clines in Nightjars

Environmental and Geographic Conditions on the Breeding Grounds Drive Bergmannian Clines in Nightjars

Journal of Biogeography, 2025

Citation

Skinner, A.A., Korpach, A.M., Åkesson, S., Bakermans, M.H., Bayne, E.M., Benson, T.J., Boano, G., Brigham, R.M., Christiansen, S.S., Conway, G.J., Davy, C.M., Evens, R., Fraser, K.C., Harrison, A.-L., Hedenström, A., Henderson, I.G., Honkala, J., Jacobsen, L.B., Lathouwers, M., Marra, P.P., Ng, J.W., Norevik, G., Scarpignato, A.L., Thorup, K., Tonra, C.M., Van Wilgenburg, S.L., Vitz, A.C., Ward, M. & Knight, E. 2025. Environmental and Geographic Conditions on the Breeding Grounds Drive Bergmannian Clines in Nightjars. Journal of Biogeography doi:10.1111/jbi.15176
GPS-tagged Nightjar being released, by Jody Lawrence / BTO

Overview

Bergmann’s rule (or ‘law’) is one of the oldest in ecology and evolutionary biology, first presented in 1847. This rule states that animals inhabiting colder climates/latitudes are larger, with greater volume than individuals of the same species occupying warmer climates/latitudes.

In more detail

Typically, we think of this in the context of relatively sedentary cold-adapted species like wolves or bears, where the more northern individuals are bigger and rounder than their more southern relatives.  However, for long distance migrant birds, like nightjars, which breed in the north but winter in warm, equatorial conditions, it is unclear which part of the annual cycle has most influence on Bergmann’s rule.

The main aim of this paper was to  investigate how variation in body size and migration, in Nightjars, is influenced by environmental and geographic conditions on the  breeding grounds. Three migrant northern hemisphere nesting species were studied, including  European Nightjar (Caprimulgus europaeus), Eastern Whip-poor-will (Antrostomus vociferus) and Common Nighthawk (Chordeiles minor).  GPS-tracking data were used to link individuals to breeding and wintering sites, from which geographic and environmental data were extracted, along with migration distance and morphometric data.

The findings showed that environmental and location variables within the  breeding range were the strongest predictors of body size, rather than at  wintering range variables. These showed that body mass (and wing length) within the breeding grounds increased northward for all three species, thus conforming to Bergmann’s rule. However, latitude and longitude correlated strongly with environmental variables and migratory distance; thus, these geographical variables likely encompass many factors that influence body size in nightjars. 

The present study is among the first to use tracking data from individual birds to understand how environmental pressures across the annual cycle are related to body size. The findings highlight the critical role of geographic breeding-ground factors in shaping Bergmannian patterns, offering robust evidence to support nearly two centuries of research.

Abstract

Aim

To evaluate (1) whether three migratory nightjar species (Family Caprimulgidae) adhere to Bergmann’s rule, (2) whether environmental factors on the breeding or wintering grounds determine body size, and (3) which mechanistic hypotheses best explain Bergmannian patterns in body size.

Location

North and South America; Europe and Africa.

Taxon

Eastern whip-poor-will (Antrostomus vociferus), Common nighthawk (Chordeiles minor) and European nightjar (Caprimulgus europaeus).

Methods

We used GPS tracking and morphometric data to assess competing hypotheses explaining variation in body size for each species, based on their breeding (n = 3388) and wintering (n = 189) locations.

Results

All three species exhibited Bergmannian patterns in body size, providing the first evidence that nightjars conform to Bergmann's rule despite adaptations to severe environmental conditions. Environmental and geographic variables at breeding sites were stronger predictors of body size than wintering-site variables. Although we found partial support for Bergmann’s temperature regulation hypothesis, geographic variables, rather than specific environmental factors, emerged as the strongest predictors of body size variation.

Main Conclusions

Latitude and longitude correlated strongly with environmental variables and migratory distance; thus, these geographical variables likely encompass many factors that influence body size in nightjars. The present study is among the first to use tracking data from individual birds to understand how environmental pressures across the annual cycle are related to body size. Our findings highlight the critical role of geographic breeding-ground factors in shaping Bergmannian patterns, offering robust evidence to support nearly two centuries of research since Bergmann's rule was first described in 1847.

This work was supported by Ohio Agricultural Research and Development Center, British Trust for Ornithology, Environment and Climate Change Canada, EACOM Timber Corporation, Association of Field Ornithologists, Illinois Department of Natural Resources, Natural Sciences and Engineering Research Council of Canada, U.S. Fish and Wildlife Service, Mitacs, Conoco Phillips, Mark Constantine, Weyerhauser Company Limited, Svenska Forskningsrådet Formas, University of Manitoba, The British Birds Charitable Trust and Canadian Foundation for Innovation.