# Non-exchangeable hydrogen (δ2H) stable isotope ratios in fauna provide enhanced dietary, isotopic niche and home range reconstruction at Aqaba Castle, Jordan

**Authors:** Gene T. Shev, Bea De Cupere, Anastasia Brozou, Benjamin T. Fuller, Marcello A. Mannino, Joris Peters, Wim Van Neer, Steven Bouillon, Claudio Ottoni

PMC · DOI: 10.1371/journal.pone.0328991 · PLOS One · 2025-08-01

## TL;DR

This study shows that hydrogen isotope ratios in animal bones from Aqaba Castle help better understand diets, habitats, and water sources in a dry region.

## Contribution

The study introduces δ2H as a more effective isotope proxy for niche differentiation and geographic provenancing in arid environments.

## Key findings

- δ2H values show less overlap than δ13C and δ15N, improving isotopic niche differentiation.
- Camels and goats have smaller δ2H offsets from rainfall compared to chickens and cattle.
- δ2H values suggest gazelles and chukars inhabited highland, water-stressed environments near Aqaba Castle.

## Abstract

Bone collagen of terrestrial and marine animals (n = 218) recovered from Ottoman period contexts at Aqaba Castle, Jordan (16th–19th centuries CE), were analyzed for δ13C, δ15N, and δ2H isotope ratios. While δ13C and δ15N values showed considerable overlap among species in the hyperarid environment, δ2H values exhibited less overlap, enhancing stable isotopic niche differentiation. In domesticates, δ2H values show trophic enrichments of +18.4‰ from herbivores to omnivores (dogs), and +26‰ to cats which had the highest δ2H values. Fish δ2H values show a positive relationship with increasing trophic level but also moderately correlate with body size (r = 0.61, R2 = 0.37). The offset between collagen δ2H and rainfall (δ2Hmw) values is smaller for camels (−1.4‰), sheep (−4.5‰), and goats (+6.8%), than for chickens (−18.5‰) and cattle (−27.0‰) due to more frequent consumption of 2H-depleted groundwater by the latter species, because of their higher water requirements. Similarities between local precipitation and bone collagen δ2H values for most terrestrial herbivores suggest the utility of δ2H values for geographic provenancing. This is explored by overlapping gazelle and chukar collagen δ2H values over a regional δ2Hmw isoscape, tentatively suggesting these species inhabited the water-stressed highland environments surrounding Aqaba Castle. This study demonstrates the advantages of incorporating bone collagen δ2H values alongside δ13C and δ15N values as a useful environmental proxy, enhancing interpretations of animal dietary behaviour, trophic levels, water sources, and wild animal home ranges.

## Full-text entities

- **Chemicals:** CO2 (MESH:D002245), amino acid (MESH:D000596), Distilled Water (MESH:D014867), N (MESH:D009584), strontium (MESH:D013324), Cr (MESH:D002857), H (MESH:D006859), arginine (MESH:D001120), essential amino acids (MESH:D000601), caffeine (MESH:D002110), lipid (MESH:D008055), HCl (MESH:D006851), humic acid (MESH:D006812), 87Sr (-), ammonia (MESH:D000641), 2H (MESH:D003903), C (MESH:D002244), oxygen (MESH:D010100), salt (MESH:D012492), NaOH (MESH:D012972), 13C (MESH:C000615229), taurine (MESH:D013654), silver (MESH:D012834)
- **Species:** Equus asinus (African ass, species) [taxon 9793], Equus caballus (domestic horse, species) [taxon 9796], Mus musculus (house mouse, species) [taxon 10090], Peperomia congesta (species) [taxon 405311], Lepus (hares, genus) [taxon 9980], Haloxylon salicornicum (species) [taxon 454511], Hyaena hyaena (striped hyena, species) [taxon 95912], Procavia capensis (Cape hyrax, species) [taxon 9813], Gallus gallus (bantam, species) [taxon 9031], Sparisoma cretense (species) [taxon 59664], Labriformes (wrasses, order) [taxon 1489928], Camelus dromedarius (Arabian camel, species) [taxon 9838], Bos taurus (bovine, species) [taxon 9913], Gazella subgutturosa (black-tailed gazelle, species) [taxon 59529], Carangidae (jacks, family) [taxon 8157], Neophron percnopterus (Egyptian vulture, species) [taxon 33608], Sorghum bicolor (broomcorn, species) [taxon 4558], Puma concolor (puma, species) [taxon 9696], Gazella dorcas (Dorcas gazelle, species) [taxon 37751], Panicum miliaceum (broomcorn millet, species) [taxon 4540], Odocoileus virginianus (white-tailed deer, species) [taxon 9874], Lynx rufus (bobcat, species) [taxon 61384], Lepus capensis (brown hare, species) [taxon 9981], Bos sp. (species) [taxon 29061], Rattus norvegicus (brown rat, species) [taxon 10116], Cavia porcellus (domestic guinea pig, species) [taxon 10141], Anabasis articulata (species) [taxon 996498], Mugiliformes (mullets, order) [taxon 41872], Felis catus (cat, species) [taxon 9685], Oryctolagus cuniculus (domestic rabbit, species) [taxon 9986], Canis lupus familiaris (dog, subspecies) [taxon 9615], Corvus corax (Common raven, species) [taxon 56781], Homo sapiens (human, species) [taxon 9606], Alectoris chukar (chukar partridge, species) [taxon 9078], Capra hircus (domestic goat, species) [taxon 9925], S. floridanus [taxon 925642], Argyrops filamentosus (soldierbream, species) [taxon 990556], Cetoscarus bicolor (bicolor parrotfish, species) [taxon 188107], Lutjanidae (fusiliers, family) [taxon 30850], Camelus sp. (species) [taxon 30536], Gazella gazella (mountain gazelle, species) [taxon 69302], Ovis aries (domestic sheep, species) [taxon 9940], PX clade (clade) [taxon 569578], Setaria italica (foxtail millet, species) [taxon 4555], Sylvilagus floridanus (eastern cottontail, species) [taxon 9988], Gazella (gazelle, genus) [taxon 9933]

## Full text

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## References

108 references — full list in the complete paper: https://tomesphere.com/paper/PMC12316241/full.md

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Source: https://tomesphere.com/paper/PMC12316241