Paul Szpak

Complexities of nitrogen isotope biogeochemistry in plant-soil systems: implications for the study of ancient agricultural and animal management practices.

Nitrogen isotopic studies have the potential to shed light on the structure of ancient ecosystems, agropastoral regimes, and human-environment interactions. Until relatively recently, however, little attention was paid to the complexities of nitrogen transformations in ancient plant-soil systems and their potential impact on plant and animal tissue nitrogen isotopic compositions. This paper discusses the importance of understanding nitrogen dynamics in ancient contexts, and highlights several key areas of archaeology where a more detailed understanding of these processes may enable us to answer some fundamental questions. This paper explores two larger themes that are prominent in archaeological studies using stable nitrogen isotope analysis: (1) agricultural practices (use of animal fertilizers, burning of vegetation or shifting cultivation, and tillage) and (2) animal domestication and husbandry (grazing intensity/stocking rate and the foddering of domestic animals with cultigens). The paucity of plant material in ancient deposits necessitates that these issues are addressed primarily through the isotopic analysis of skeletal material rather than the plants themselves, but the interpretation of these data hinges on a thorough understanding of the underlying biogeochemical processes in plant-soil systems. Building on studies conducted in modern ecosystems and under controlled conditions, these processes are reviewed, and their relevance discussed for ancient contexts.

Carbon and Nitrogen Isotopic Survey of Northern Peruvian Plants: Baselines for Paleodietary and Paleoecological Studies

The development of isotopic baselines for comparison with paleodietary data is crucial, but often overlooked. We review the factors affecting the carbon (δ13C) and nitrogen (δ15N) isotopic compositions of plants, with a special focus on the carbon and nitrogen isotopic compositions of twelve different species of cultivated plants (n = 91) and 139 wild plant species collected in northern Peru. The cultivated plants were collected from nineteen local markets. The mean δ13C value for maize (grain) was −11.8±0.4 ‰ (n = 27). Leguminous cultigens (beans, Andean lupin) were characterized by significantly lower δ15N values and significantly higher %N than non-leguminous cultigens. Wild plants from thirteen sites were collected in the Moche River Valley area between sea level and ∼4,000 meters above sea level (masl). These sites were associated with mean annual precipitation ranging from 0 to 710 mm. Plants growing at low altitude sites receiving low amounts of precipitation were characterized by higher δ15N values than plants growing at higher altitudes and receiving higher amounts of precipitation, although this trend dissipated when altitude was >2,000 masl and MAP was >400 mm. For C3 plants, foliar δ13C was positively correlated with altitude and precipitation. This suggests that the influence of altitude may overshadow the influence of water availability on foliar δ13C values at this scale.

Stable Isotope Biogeochemistry of Seabird Guano Fertilization: Results from Growth Chamber Studies with Maize (Zea Mays)

Background Stable isotope analysis is being utilized with increasing regularity to examine a wide range of issues (diet, habitat use, migration) in ecology, geology, archaeology, and related disciplines. A crucial component to these studies is a thorough understanding of the range and causes of baseline isotopic variation, which is relatively poorly understood for nitrogen (δ15N). Animal excrement is known to impact plant δ15N values, but the effects of seabird guano have not been systematically studied from an agricultural or horticultural standpoint. Methodology/Principal Findings This paper presents isotopic (δ13C and δ15N) and vital data for maize (Zea mays) fertilized with Peruvian seabird guano under controlled conditions. The level of 15N enrichment in fertilized plants is very large, with δ15N values ranging between 25.5 and 44.7‰ depending on the tissue and amount of fertilizer applied; comparatively, control plant δ15N values ranged between −0.3 and 5.7‰. Intraplant and temporal variability in δ15N values were large, particularly for the guano-fertilized plants, which can be attributed to changes in the availability of guano-derived N over time, and the reliance of stored vs. absorbed N. Plant δ13C values were not significantly impacted by guano fertilization. High concentrations of seabird guano inhibited maize germination and maize growth. Moreover, high levels of seabird guano greatly impacted the N metabolism of the plants, resulting in significantly higher tissue N content, particularly in the stalk. Conclusions/Significance The results presented in this study demonstrate the very large impact of seabird guano on maize δ15N values. The use of seabird guano as a fertilizer can thus be traced using stable isotope analysis in food chemistry applications (certification of organic inputs). Furthermore, the fertilization of maize with seabird guano creates an isotopic signature very similar to a high-trophic level marine resource, which must be considered when interpreting isotopic data from archaeological material.

Regional ecological variability and impact of the maritime fur trade on nearshore ecosystems in southern Haida Gwaii (British Columbia, Canada): evidence from stable isotope analysis of rockfish (Sebastes spp.) bone collagen

The maritime fur trade (1785–1840s) led to the local extinction of sea otters (Enhydra lutris) in many parts of the northeast Pacific. On the basis of studies of extant sea otter populations, it has been established that they have a disproportionate effect on nearshore ecosystems by limiting sea urchin abundance and facilitating the establishment of nearshore kelp forests; in the absence of sea otters, a local reduction in kelp-derived carbon is therefore expected. We measured the isotopic composition (δ13C and δ15N) of rockfish (Sebastes spp.) bone collagen from late Holocene archaeological sites in southern Haida Gwaii, BC, Canada, using δ13C as a proxy for kelp-derived carbon in the diet and δ15N as a proxy for trophic position. We observed significant spatial variability in rockfish kelp-derived carbon (δ13C), but not trophic level (δ15N). Kelp-derived carbon varied largely as a function of site characteristics (wave exposure), suggesting that local oceanographic conditions are important factors with respect to consumer tissue isotopic compositions. Kelp-derived carbon decreased in post-European contact rockfish relative to pre-European contact rockfish, likely as a result of the reduction of kelp forests associated with the local extirpation of sea otters. Although we detected a reduction in kelp-derived carbon in rockfish diets, we found no shift in trophic level at sites occupied following the maritime fur trade. This implies a shift in local ecosystems, and particularly in carbon sources, following the maritime fur trade, likely due to a trophic cascade resulting from the local extirpation of sea otters. Stability in rockfish trophic levels, however, implies that rockfish continued to feed at similar trophic levels, consuming prey with similar nitrogen isotopic compositions.

Mammuthus Population Dynamics in Late Pleistocene North America: Divergence, Phylogeography, and Introgression

After evolving in Africa at the close of the Miocene, mammoths (Mammuthus sp.) spread through much of the northern hemisphere, diversifying morphologically as they entered various habitats. Paleontologically, these morphs are conventionally recognized as species. In Pleistocene North America alone, several mammoth species have been recognized, inhabiting environments as different as cold tundra-steppe in the north and the arid grasslands or temperate savanna-parklands of the south. Yet mammoth phylogeographic studies have overwhelmingly focused on permafrost-preserved remains of only one of these species, Mammuthus primigenius (woolly mammoth). Here we challenge this bias by performing a geographically and taxonomically wide survey of mammoth genetic diversity across North America. Using a targeted enrichment technique, we sequenced 67 complete mitochondrial genomes from non-primigenius specimens representing M. columbi (Columbian mammoth), M. jeffersonii (Jeffersonian mammoth), and M. exilis (pygmy mammoth), including specimens from contexts not generally associated with good DNA preservation. While we uncovered clear phylogeographic structure in mammoth matrilines, their phylogeny as recovered from mitochondrial DNA is not compatible with existing systematic interpretations of their paleontological record. Instead, our results strongly suggest that various nominal mammoth species interbred, perhaps extensively. We hypothesize that at least two distinct stages of interbreeding between conventional paleontological species are likely responsible for this pattern – one between Siberian woolly mammoths and resident American populations that introduced woolly mammoth phenotypes to the continent, and another between ecomorphologically distinct populations of woolly and Columbian mammoths in North America south of the ice.

Zinc Isotope Ratios as Indicators of Diet and Trophic Level in Arctic Marine Mammals.

Carbon and nitrogen stable isotope ratios of bone collagen are an established method for dietary reconstruction, but this method is limited by the protein preservation. Zinc (Zn) is found in bioapatite and the isotopic compositions of this element constitute a very promising dietary indicator. The extent of fractionation of Zn isotopes in marine environments, however, remains unknown. We report here on the measurement of zinc, carbon and nitrogen isotopes in 47 marine mammals from the archaeological site of Arvik in the Canadian Arctic. We undertook this study to test and demonstrate the utility of Zn isotopes in recent mammal bone minerals as a dietary indicator by comparing them to other isotopic dietary tracers. We found a correlation between δ66Zn values and trophic level for most species, with the exception of walruses, which may be caused by their large seasonal movements. δ6Zn values can therefore be used as a dietary indicator in marine ecosystems for both modern and recent mammals.

Effects of lipid extraction and ultrafiltration on stable carbon and nitrogen isotopic compositions of fish bone collagen.

RATIONALE Fish bone collagen isotopic measurements are increasingly important in palaeodietary and paleoenvironmental studies yet differences in the chemical and physical properties of fish relative to other vertebrate bones are rarely considered. Lipid content in fish bone, which can exceed 50%, may underlie the poor collagen integrity criteria typically observed in archaeological studies. METHODS We compare stable carbon and nitrogen isotopic and elemental compositions of bone collagen prepared using four different methods from a wide range of modern fish species to: (1) assess the extent to which lipid content influences bone collagen δ(13) C and δ(15) N values, and (2) evaluate the relative efficacy of chemical (2:1 chloroform/methanol) and physical (30 kDa ultrafilters) methods for removing lipids from bones. RESULTS Lower δ(13) C values were observed when the lipid content exceeded 5% of the initial bone mass. The lipid content did not influence the δ(15) N values. 30 kDa ultrafiltration, a common pretreatment for purifying archaeological collagen, removed fewer lipids and was associated with reduced collagen yields (37% loss) as well as altered amino acid compositions. In contrast, collagen prepared using a 2:1 chloroform/methanol lipid extraction step resulted in significantly improved collagen yields, elemental compositions, and isotopic measurements relative to a control treatment. CONCLUSIONS The chemical lipid extraction method (2:1 chloroform/methanol) performed significantly better than the physical lipid extraction method (30 kDa ultrafilters). Given the high quantities of lipids in fish bones we recommend the inclusion of a chemical lipid extraction step when isolating collagen from modern and archaeological fish bones. Copyright

Early Horizon camelid management practices in the Nepeña Valley, north-central coast of Peru

South American camelids (llamas and alpacas) were of great economic, social and ritual significance in the pre-Hispanic Andes. Although these animals are largely limited to high-altitude (>3500 masl) pastures, it has been hypothesised that camelids were also raised at lower altitudes in the arid coastal river valleys. Previous isotopic studies of Early Intermediate Period (c. 200 BC–AD 600) and Middle Horizon (c. AD 600–1100) camelids support this argument. Here, we utilise carbon and nitrogen isotopic analyses of camelid bone collagen from the Early Horizon (c. 800–200 BC) sites of Caylán and Huambacho on the north-central coast of Peru to examine the management of these animals during the first millennium BC. Most of the camelid isotopic compositions are consistent with the acquisition of animals that were part of caravans, moving between the coast and the highlands. A small number of the animals may have been raised on the coast, suggesting that the practice of coastal camelid husbandry was in the experimental phase during the Early Horizon before growing into a more established practice in the Early Intermediate Period. These results echo zooarchaeological studies from the region that have revealed a paucity of camelid remains in refuse deposits prior to 800 BC, followed by an increase in abundance after 450 BC.

Historical ecology and the conservation of large, hermaphroditic fishes in Pacific Coast kelp forest ecosystems

Zooarchaeological data are essential for rethinking conservation policies of hermaphroditic fish and building sustainable fisheries. The intensive commercial exploitation of California sheephead (Semicossyphus pulcher) has become a complex, multimillion-dollar industry. The fishery is of concern because of high harvest levels and potential indirect impacts of sheephead removals on the structure and function of kelp forest ecosystems. California sheephead are protogynous hermaphrodites that, as predators of sea urchins and other invertebrates, are critical components of kelp forest ecosystems in the northeast Pacific. Overfishing can trigger trophic cascades and widespread ecological dysfunction when other urchin predators are also lost from the system. Little is known about the ecology and abundance of sheephead before commercial exploitation. Lack of a historical perspective creates a gap for evaluating fisheries management measures and marine reserves that seek to rebuild sheephead populations to historical baseline conditions. We use population abundance and size structure data from the zooarchaeological record, in concert with isotopic data, to evaluate the long-term health and viability of sheephead fisheries in southern California. Our results indicate that the importance of sheephead to the diet of native Chumash people varied spatially across the Channel Islands, reflecting modern biogeographic patterns. Comparing ancient (~10,000 calibrated years before the present to 1825 CE) and modern samples, we observed variability and significant declines in the relative abundance of sheephead, reductions in size frequency distributions, and shifts in the dietary niche between ancient and modern collections. These results highlight how size-selective fishing can alter the ecological role of key predators and how zooarchaeological data can inform fisheries management by establishing historical baselines that aid future conservation.

Origins of Prehispanic Camelid Wool Textiles from the North and Central Coasts of Peru Traced by Carbon and Nitrogen Isotopic Analyses

Carbon and nitrogen isotopic compositions of wool textiles from the north (Virú, Early Intermediate Period) and central (Chancay, Late Intermediate Period) coasts of Peru were used to reconstruct the diet and habitat of the camelids (llamas and alpacas) from which they were produced in order to better understand the regional political economies. The Chancay textiles were derived from camelids primarily raised on high-altitude C3 grasslands. Similarly, isotopic data from Virú textiles assembled in north-coast styles are consistent with the importation of highland camelid wool. For both Virú and Chancay, imported raw materials were crafted in local styles, serving as an effective means of materializing corporate power. Stylistically foreign (noncoastal) Virú textiles were characterized by carbon isotopic compositions similar to those for camelids recovered from Early Intermediate Period sites in the Virú Valley and suggest that these textiles originated in the yungas (1,000–2,300 m asl) or the low sierra (2,300–3,500 m asl). Accordingly, although highland camelid wool was imported to the coast, a simple model of exchange involving the movement of wool textiles exclusively from the puna or the altiplano to the coast is untenable.