title: Energetic constraint of non-monotonic mass change during offspring growth: a general hypothesis and application of a new tool.
abstract: Post-natal growth is an important life-history trait and can be a sensitive indicator of ecological stress. For over 50 years, monotonic (never-decreasing) growth has been viewed as the predominant trajectory of post-natal mass change in most animal species, notably among birds. However, prevailing analytical approaches and energetic constraints may limit detection of non-monotonic (or multiphasic), determinate growth patterns, such as mass recession in birds (weight loss prior to fledging, preceded by overshooting adult mass), which is currently believed to be restricted to few taxa. Energetic surplus and shortfall are widespread conditions that can directly influence the degree of mass overshooting and recession. Thus, we hypothesize that in many species, prevailing energetic constraints force mass change away from a fundamental non-monotonic trajectory to instead follow a monotonic curve. We observed highly non-monotonic, mass change trajectories (overshooting adult mass by up to almost 20%) among common tern Sterna hirundo chicks, a well-studied species long-established as growing monotonically. We quantified the prevalence and magnitude of non-monotonic mass change prior to fledging for 313 common tern chicks that successfully fledged from two discrete populations in multiple years. We used a new approach for analysing non-monotonic curves to examine differences in mass change trajectories between populations under contrasting abiotic (freshwater vs. saltwater) and biotic stresses (low rates of food provisioning). Some degree of mass recession occurred in 73% of all study chicks. Overshooting adult mass followed by extensive mass recession was most prevalent at our freshwater colony, being detected among 34-38% of chicks annually. Non-monotonic trajectories were less marked in populations experiencing ecological stress and among lower quality individuals. Chicks that were provisioned at higher rates were more likely to both overshoot adult mass and experience subsequent mass recession. Our results in common terns provide strong support for the hypothesis that non-monotonic trajectories are the fundamental pattern of mass change but are constrained to be monotonic under energetic shortfall. This justifies future tests of the generality of this hypothesis across a broad range of taxa. We also demonstrate a recent analytical tool that prevents routine fitting of monotonic curves without prior investigation of non-monotonic trends.
authors.affiliation: Division of Science, Pennsylvania State University, Berks Campus, 2080 Tulpehocken Road, Reading, PA, 19610, USA.
authors.forename: Jennifer M
authors.initials: JM
authors.lastname: Arnold
authors.affiliation: I.C.T. Nisbet & Company, 150 Alder Lane, North Falmouth, MA, 02556, USA.
authors.forename: Ian C T
authors.initials: IC
authors.lastname: Nisbet
authors.affiliation: Division of Science, Pennsylvania State University, Berks Campus, 2080 Tulpehocken Road, Reading, PA, 19610, USA.
authors.forename: Stephen A
authors.initials: SA
authors.lastname: Oswald
mesh_terms: D000818:Animals; D046091:Charadriiformes / Q000254:growth & development / Q000502:physiology*; D002149:Energy Intake; D008404:Massachusetts; D008954:Models, Biological; D009864:Ontario; D012621:Seasons
keywords: FlexParamCurve; energetic constraints; growth analysis; mass overshooting; mass recession; nlme; non-monotonic growth curve; nonlinear mixed effects models