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"Harrod, Chris"
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Identifying knowledge gaps in hypersaline systems supporting the global electrical transition: invertebrate community structure in salars from the lithium triangle
2025
Following decades of mining impacts, South America’s hypersaline wetlands (salars) face increasing threats from lithium extraction to support global decarbonisation. Although globally important, salars are understudied and information needed to understand environmental impacts is lacking. Modern ecological studies on salars have focused on microbial community composition and function but other taxa are less studied, including resident and migratory reptiles and birds and their aquatic invertebrate prey. Given the scale and immediate nature of the threats associated with lithium exploitation, we must deepen our understanding of salar biology, but this is impeded by logistic/financial restrictions given the heightened costs of sampling in these often remote, extreme environments. Given the pressing demand for information, we collated/analysed information from the literature. We generated lists of invertebrate taxa present in 63 hypersaline environments from Argentina, Bolivia, Chile and Peru, and examined relationships between invertebrate community structure and physicochemical factors. We recorded 46 different taxa, with the Centropagidae being the most frequently recorded taxon across systems. Multivariate analyses of community structure showed significant clustering among sites. Variation in community structure was best explained by maximum salinity (18%). Geographical location or ecosystem size had no obvious effect on community structure. We provide a useful broad view of aquatic invertebrate diversity in the hypersaline salars but highlight the general lack of information regarding the ecology of these ecosystems.
Journal Article
Phenotype-environment correlations in a putative whitefish adaptive radiation
by
Harrod, Chris
,
Mallela, Jennie
,
Kahilainen, Kimmo K.
in
Adaptation
,
Adaptation, Physiological
,
Adaptation, Physiological - genetics
2010
1. The adaptive radiation of fishes into benthic (littoral) and pelagic (lentic) morphs in post-glacial lakes has become an important model system for speciation. Although these systems are well studied, there is little evidence of the existence of morphs that have diverged to utilize resources in the remaining principal lake habitat, the profundal zone. 2. Here, we tested phenotype-environment correlations of three whitefish (Coregonus lavaretus) morphs that have radiated into littoral, pelagic and profundal niches in northern Scandinavian lakes. We hypothesized that morphs in such trimorphic systems would have a morphology adapted to one of the principal lake habitats (littoral, pelagic or profundal niches). Most whitefish populations in the study area are formed by a single (monomorphic) whitefish morph, and we further hypothesized that these populations should display intermediate morphotypes and niche utilization. We used a combination of traditional (stomach content, habitat use, gill raker counts) and more recently developed (stable isotopes, geometric morphometrics) techniques to evaluate phenotype-environment correlations in two lakes with trimorphic and two lakes with monomorphic whitefish. 3. Distinct phenotype-environment correlations were evident for each principal niche in whitefish morphs inhabiting trimorphic lakes. Monomorphic whitefish exploited multiple habitats, had intermediate morphology, displayed increased variance in gillraker-counts, and relied significantly on zooplankton, most likely due to relaxed resource competition. 4. We suggest that the ecological processes acting in the trimorphic lakes are similar to each other, and are driving the adaptive evolution of whitefish morphs, possibly leading to the formation of new species.
Journal Article
How strongly does diet variation explain variation in isotope values of animal consumers?
by
Harrod, Chris
,
Bortoluzzi, Jenny Rose
,
Rowland, Hugh
in
Animals
,
Biology and Life Sciences
,
Biomass
2024
Analysis of stable isotopes in consumers is used commonly to study their ecological and/or environmental niche. There is, however, considerable debate regarding how isotopic values relate to diet and how other sources of variation confound this link, which can undermine the utility. From the analysis of a simple, but general, model of isotopic incorporation in consumer organisms, we examine the relationship between isotopic variance among individuals, and diet variability within a consumer population. We show that variance in consumer isotope values is directly proportional to variation in diet (through Simpson indices), to the number of isotopically distinct food sources in the diet, and to the baseline variation within and among the isotope values of the food sources. Additionally, when considering temporal diet variation within a consumer we identify the interplay between diet turnover rates and tissue turnover rates that controls the sensitivity of stable isotopes to detect diet variation. Our work demonstrates that variation in the stable isotope values of consumers reflect variation in their diet. This relationship, however, can be confounded with other factors to the extent that they may mask the signal coming from diet. We show how simple quantitative corrections can recover a direct 1:1 correlation in some situations, and in others we can adjust our interpretation in light of the new understanding arising from our models. Our framework provides guidance for the design and analysis of empirical studies where the goal is to infer niche width from stable isotope data.
Journal Article
Clarifying a trophic black box: stable isotope analysis reveals unexpected dietary variation in the Peruvian anchovy Engraulis ringens
2019
Small fishes play fundamental roles in pelagic ecosystems, channelling energy and nutrients from primary producers to higher trophic levels. They support globally important fisheries in eastern boundary current ecosystems like the Humboldt Current System (HCS) of the SE Pacific (Chile and Peru), where fish catches are the highest in the world (per unit area). This production is associated with coastal upwelling where fisheries target small pelagic fishes including the Peruvian anchovy (
). The elevated biomass attained by small pelagics is thought to reflect their low trophic position in short/simple food chains. Despite their global importance, large gaps exist in our understanding of the basic ecology of these resources. For instance, there is an ongoing debate regarding the relative importance of phytoplankton versus animal prey in anchovy diet, and ecosystem models typically assign them a trophic position (TP) of ∼2, assuming they largely consume phytoplankton. Recent work based on both relative energetic content and stable isotope analysis (SIA) suggests that this value is too low, with
N values indicating that anchovy TP is ca. 3.5 in the Peruvian HCS.
We characterised the trophic ecology of adult anchovies (
= 30), their putative prey and carnivorous jack mackerel (
= 20) captured from N Chile. SIA (
C and
N) was used to estimate the relative contribution of different putative prey resources.
N was used to estimate population level trophic position.
Anchovies showed little variability in
C (-18.7 to -16.1‰) but varied greatly in
N (13.8 to 22.8‰)-individuals formed two groups with low or high
N values. When considered as a single group, mixing models indicated that anchovy diet was largely composed of zooplankton (median contribution: 95% credibility limits), with major contributions of crustacean larvae (0.61: 0.37-0.77) and anchovy (preflexion) larvae (0.15: 0.02-0.34), and the assimilation of phytoplankton was negligible (0.05: 0-0.22). The modal (95% credibility limits) estimate of TP for the pooled anchovy sample was 3.23 (2.93-3.58), overlapping with recent SIA-based estimates from Peru. When the two
N groups were analysed separately, our results indicate that the lower
N group largely assimilated materials from crustacean larvae (0.73: 0.42-0.88), with a TP of 2.91 (2.62-3.23). Mixing models suggested high
N anchovies were cannibalistic, consuming anchovy preflexion larvae (0.55: 0.11-0.74). A carnivorous trophic niche was supported by high TP (3.79: 3.48-4.16), mirroring that of carnivorous juvenile jack mackerel (
; 3.80: 3.51-4.14). Our results support recent conclusions regarding high TP values of anchovy from Peru and reveal new insights into their trophic behaviour. These results also highlight the existence of cryptic trophic complexity and ecosystem function in pelagic food webs, classically considered as simple.
Journal Article
Adaptive Radiation along a Thermal Gradient: Preliminary Results of Habitat Use and Respiration Rate Divergence among Whitefish Morphs
by
Harrod, Chris
,
Patterson, William Paul
,
Kahilainen, Kimmo Kalevi
in
Adaptation
,
Adaptation, Physiological
,
Adaptive radiation
2014
Adaptive radiation is considered an important mechanism for the development of new species, but very little is known about the role of thermal adaptation during this process. Such adaptation should be especially important in poikilothermic animals that are often subjected to pronounced seasonal temperature variation that directly affects metabolic function. We conducted a preliminary study of individual lifetime thermal habitat use and respiration rates of four whitefish (Coregonus lavaretus (L.)) morphs (two pelagic, one littoral and one profundal) using stable carbon and oxygen isotope values of otolith carbonate. These morphs, two of which utilized pelagic habitats, one littoral and one profundal recently diverged via adaptive radiation to exploit different major niches in a deep and thermally stratified subarctic lake. We found evidence that the morphs used different thermal niches. The profundal morph had the most distinct thermal niche and consistently occupied the coldest thermal habitat of the lake, whereas differences were less pronounced among the shallow water pelagic and littoral morphs. Our results indicated ontogenetic shifts in thermal niches: juveniles of all whitefish morphs inhabited warmer ambient temperatures than adults. According to sampling of the otolith nucleus, hatching temperatures were higher for benthic compared to pelagic morphs. Estimated respiration rate was the lowest for benthivorous profundal morph, contrasting with the higher values estimated for the other morphs that inhabited shallower and warmer water. These preliminary results suggest that physiological adaptation to different thermal habitats shown by the sympatric morphs may play a significant role in maintaining or strengthening niche segregation and divergence in life-history traits, potentially contributing to reproductive isolation and incipient speciation.
Journal Article
Comparing trophic position estimates using bulk and compound specific stable isotope analyses: applying new approaches to mackerel icefish Champsocephalus gunnari
2024
Quantifying the tropic position (TP) of an animal species is key to understanding its ecosystem function. While both bulk and compound-specific analyses of stable isotopes are widely used for this purpose, few studies have assessed the consistency between and within such approaches. Champsocephalus gunnari is a specialist teleost that predates almost exclusively on Antarctic krill Euphausia superba . This well-known and nearly constant trophic relationship makes C. gunnari particularly suitable for assessing consistency between TP methods under field conditions. In the present work, we produced and compared TP estimates for C. gunnari and its main prey using a standard bulk and two amino acid-specific stable isotope approaches (CSI-AA). One based on the difference between glutamate and phenylalanine (TP Glx-Phe ), and the other on the proline-phenylalanine difference (TP Pro-Phe ). To do that, samples from C. gunnari, E. superba and four other pelagic invertebrate and fish species, all potential prey for C.gunnari, were collected off the South Orkney Islands between January and March 2019, analyzed using standard isotopic ratio mass spectrometry methods and interpreted following a Bayesian approach. Median estimates (CI 95% ) for C. gunnari were similar between TP bulk (3.6; CI 95% : 3.0-4.8) and TP Glx-Phe (3.4; CI 95% :3.2-3.6), and lower for TP Pro-Phe (3.1; CI 95% :3.0-3.3). TP differences between C. gunnari and E. superba were 1.4, 1.1 and 1.2, all compatible with expectations from the monospecific diet of this predator (ΔTP=1). While these results suggest greater accuracy for Glx-Phe and Pro-Phe, differences observed between both CSI-AA approaches suggests these methods may require further validation before becoming a standard tool for trophic ecology.
Journal Article
Diet of Andean Leaf‐Eared Mice (Phyllotis) Living at Extreme Elevations on Atacama Volcanoes: Insights From Metagenomics, DNA Metabarcoding, and Stable Isotopes
by
Harrod, Chris
,
Quiroga‐Carmona, Marcial
,
Quezada‐Romegialli, Claudio
in
Andes
,
aolian zone
,
Arthropods
2024
On the flanks of > 6000 m Andean volcanoes that tower over the Atacama Desert, leaf‐eared mice (Phyllotis vaccarum) live at extreme elevations that surpass known vegetation limits. The diet of these mice in these barren, hyperarid environments has been the subject of much speculation. According to the arthropod fallout hypothesis, sustenance is provided by windblown insects that accumulate in snowdrifts (“aolian deposits”). Mice may also feed on saxicolous lichen or forms of cryptic vegetation that have yet to be discovered at such high elevations. We tested hypotheses about the diet of mice living at extreme elevations on Atacama volcanoes by combining metagenomic and DNA metabarcoding analyses of gut contents with stable isotope analyses of mouse tissues. Genomic analyses of contents of the gastrointestinal tract of a live‐captured mouse from the summit of Volcán Llullaillaco (6739 m) revealed an opportunistic but purely herbivorous diet, including lichens. Although we found no evidence of animal DNA in gut contents of the summit mouse, stable isotope data indicate that mice from elevations at or near vegetation limits (~5100 m) include a larger fraction of animal prey in their diet than mice from lower elevations. Some plant species detected in the gut contents of the summit mouse are known to exist at lower elevations at the base of the volcano and in the surrounding Altiplano, suggesting that they may occur at higher elevations beneath the snowpack or in other cryptic microhabitats. On the flanks of > 6000 m Andean volcanoes that tower over the Atacama Desert, leaf‐eared mice (Phyllotis vaccarum) live at extreme elevations that surpass known vegetation limits. We used a combination of metagenomics, DNA metabarcoding, and stable isotope analyses to investigate the diet of mice living in these extreme environments. Results revealed several surprises and suggest that mice living at elevations that exceed apparent vegetation limits may feed on plants beneath the snowpack or in other cryptic microhabitats.
Journal Article
Ecological plasticity of the European eel Anguilla anguilla in a tidal Atlantic lake system in Ireland
by
Harrod, Chris
,
Arai, Takaomi
,
McCarthy, T. Kieran
in
Anguilla anguilla
,
Aquatic habitats
,
Brackish water
2019
Recent studies have shown that anguillid eel populations in habitats spanning the marine–freshwater ecotone can display extreme plasticity in the range of catadromy expressed by individual fish. The apparent use of marine and freshwater habitats by the European eel Anguilla anguilla was examined by analysing the strontium (Sr) and calcium (Ca) concentrations in otoliths of eels collected from a tidal Atlantic lake system in Ireland. Variations of the Sr:Ca ratio in the otoliths indicated that a variety of environmental salinities had been experienced in the habitats that were occupied during the growth phase of these individuals in the tidal Atlantic lake system. The otolith microchemistry of these eels indicated that most of the eels had entered each salinity environment (freshwater (FW); brackish water (BW); marine-dominated water (MW) and full seawater (SW)) from fresh water to full seawater just after recruitment and had stayed in each environment until maturation without movement to other salinity environments. Only 2 of 93 (2%) eels had shifted their habitat once in their lives. This result suggests that each individual might have an environmental habitat preference, although each individual could move along a short (<2 km) salinity gradient.
Journal Article
Rapid sympatric ecological differentiation of crater lake cichlid fishes within historic times
by
Harrod, Chris
,
Meyer, Axel
,
Kautt, Andreas F
in
Adaptation (Biology)
,
Adaptation, Biological - genetics
,
Adaptation, Biological - physiology
2010
Background
After a volcano erupts, a lake may form in the cooled crater and become an isolated aquatic ecosystem. This makes fishes in crater lakes informative for understanding sympatric evolution and ecological diversification in barren environments. From a geological and limnological perspective, such research offers insight about the process of crater lake ecosystem establishment and speciation. In the present study we use genetic and coalescence approaches to infer the colonization history of Midas cichlid fishes (
Amphilophus
cf.
citrinellus
) that inhabit a very young crater lake in Nicaragua-the
ca
. 1800 year-old Lake Apoyeque. This lake holds two sympatric, endemic morphs of Midas cichlid: one with large, hypertrophied lips (~20% of the total population) and another with thin lips. Here we test the associated ecological, morphological and genetic diversification of these two morphs and their potential to represent incipient speciation.
Results
Gene coalescence analyses [11 microsatellite loci and mitochondrial DNA (mtDNA) sequences] suggest that crater lake Apoyeque was colonized in a single event from the large neighbouring great lake Managua only about 100 years ago. This founding in historic times is also reflected in the extremely low nuclear and mitochondrial genetic diversity in Apoyeque. We found that sympatric adult thin- and thick-lipped fishes occupy distinct ecological trophic niches. Diet, body shape, head width, pharyngeal jaw size and shape and stable isotope values all differ significantly between the two lip-morphs. The eco-morphological features pharyngeal jaw shape, body shape, stomach contents and stable isotopes (δ
15
N) all show a bimodal distribution of traits, which is compatible with the expectations of an initial stage of ecological speciation under disruptive selection. Genetic differentiation between the thin- and thick-lipped population is weak at mtDNA sequence (
F
ST
= 0.018) and absent at nuclear microsatellite loci (
F
ST
< 0.001).
Conclusions
This study provides empirical evidence of eco-morphological differentiation occurring very quickly after the colonization of a new and vacant habitat. Exceptionally low levels of neutral genetic diversity and inference from coalescence indicates that the Midas cichlid population in Apoyeque is much younger (
ca
. 100 years or generations old) than the crater itself (
ca
. 1 800 years old). This suggests either that the crater remained empty for many hundreds of years after its formation or that remnant volcanic activity prevented the establishment of a stable fish population during the early life of the crater lake. Based on our findings of eco-morphological variation in the Apoyeque Midas cichlids, and known patterns of adaptation in Midas cichlids in general, we suggest that this population may be in a very early stage of speciation (incipient species), promoted by disruptive selection and ecological diversification.
Journal Article
Habitat Mosaic Limits Gene Flow and Promotes Morphological Adaptation in a Generalist Mammal
by
Harrod, Chris
,
Montgomery, W. Ian
,
Booth, Warren
in
Adaptation
,
Agricultural land
,
Apodemus sylvaticus
2025
Many habitat generalist species exploit habitat patches of differing types and quality, yet the influence of such habitat mosaics on genetic structure remains poorly understood. Here, we tested whether fine‐scale habitat heterogeneity affects the population structure of the European wood mouse (Apodemus sylvaticus) by sampling three matched forest parks in Northern Ireland across hedgerow, forest edge, and inner forest habitats. Microsatellite analysis revealed strong genetic differentiation among sites and consistent divergence between habitat types within sites. Stable isotope data showed that hedgerow mice fed at a higher trophic level than inner forest individuals, with forest edge mice intermediate. Mandible shape also differed by habitat and was correlated with δ15N, though differences were subtle and may reflect both drift and plasticity. Together, these results indicate that habitat mosaics can promote repeated, fine‐scale population divergence even in the absence of physical barriers. This highlights the role of ecological heterogeneity in structuring genetic variation in widespread generalists and cautions against assuming panmixia in continuous landscapes. We investigated how fine‐scale habitat heterogeneity influences genetic structure in the European wood mouse by sampling individuals across hedgerow, forest edge, and inner forest habitats in three Northern Ireland forest parks. Microsatellite and stable isotope analyses revealed consistent genetic divergence and dietary differences among habitat types, with subtle but significant variation in mandible shape. These findings suggest that ecological mosaics can drive fine‐scale population divergence in habitat generalists, even without physical barriers.
Journal Article