Parasitism of prehistoric humans and companion animals from Antelope Cave, Mojave County, northwest Arizona

Previously, we reported a tick recovered from Antelope Cave in extreme northwest Arizona. Further analyses of coprolites from Antelope Cave revealed additional parasitological data from coprolites of both human and canid origin. A second tick was found. This site is the only archaeological locality where ticks have been recovered. We also discovered an acanthocephalan in association with Enterobius vermicularis eggs in the same coprolite. This association shows that the coprolite was deposited by a human. This discovery expands our knowledge of the range of prehistoric acanthocephalan infection. In addition, findings from canid coprolites of Trichuris vulpis are reported. This is the first published discovery of T. vulpis from a North American archaeological context. The close association of dogs with humans at Ancestral Puebloan (Anasazi) sites raises the potential that zoonotic parasites were transferred to the human population. The archaeological occupation is associated with the Ancestral Pueblo culture 1,100 yr ago.     

Applying modern analogs to understand the pollen content of coprolites

Most scientists working with coprolites from archaeological contexts assume that human fecal specimens reflect the mixing of the pollen ingested during the period in which the contribution to the coprolite, 19 to 37.5 h, was ingested, that the amount of pollen in a fecal sample directly reflects the amount of pollen originally ingested during that interval, and that differences between the amounts of pollen in different fecal specimens reflect differences in the quantities of pollen ingested at different times. These assumptions were tested and found wanting in an experiment in which two persons sequentially ate separate quantities of 15 pollen types in meals over a four-day interval. The pollen was retrieved and analyzed from feces produced during those four days and five days of subsequent fecal production. Pollen ingested first appeared in relatively small amounts, usually the day after it was ingested. Its concentration per gram of sample then increased rapidly and remained high over a one to three day interval relative to the amounts in previous and subsequent fecal specimens deposited. When pollen concentrations declined some pollen was retained in the gastrointestinal system and much lower concentrations per gram of sample of each type continued to appear in fecal samples for several days. These relatively low pollen concentrations appeared in fecal samples approximately twice as often as did higher concentrations. Our results indicate that comparatively high pollen concentrations can be used to determine that a given pollen type was ingested, but comparisons between pollen concentrations of the same pollen type in different fecal specimens or between different pollen types in the same fecal specimen, cannot be used to determine whether different amounts of pollen were ingested, or what was the relative amount of each ingested. Because pollen concentrations per gram of sample varied widely with time since ingestion, percentages of given pollen types did not occur in predictable patterns and could actually increase as the concentration of the pollen type decreases. Hence, percentages should not be used in coprolite pollen analysis. The experimental results also suggest that variations in the pollen content of different portions of a coprolite are meaningful only in terms of the overall pattern of a sequential group of coprolites.     

南京汤山驼子洞鬣狗粪化石的孢粉分析

南京汤山驼子洞洞穴堆积中鬣狗粪化石含有相对丰富的孢粉,推测草食性哺乳动物食用带有孢粉的植物后,经鬣狗的猎杀、摄食、消化和排泄,孢粉较好地保存在鬣狗粪中,形成粪化石,成为洞穴地层的组成部分。虽然鬣狗粪化石中的孢粉含量不及湖沼相地层中丰富,但有几种植物的花粉保存较好且相对集中,如Pinus,Tsuga,Quercus,Carpinus,Artemisia,Cyperaceae,Polygonum,Polypodiaceae出现的频率较高。孢粉组合揭示早更新世南京地区植被主要是草原或森林-草地,气候凉干或半干旱半湿润,可以和地层中脊椎动物群反映的自然环境相互印证。在研究洞穴古环境的代用指标中,堆积物中的鬣狗粪化石中包含的孢粉是比较好的研究对象,对环境有一定的指示意义。     

The science of coprolite analysis: The view from Hinds cave

This paper explores some of the new perspectives resulting from re-examination of the pollen data from my 1978 comprehensive analysis of 100 coprolites from one of the earliest identified prehistoric latrines in North America. Of particular interest are the empirical results of previously unavailable pollen concentration calculations for some of the prehistoric specimens as well as a time-series of 82 modern fecal specimens produced during an experiment yielding data on the rate of elimination of specific pollen grains from the human system. Experimental data of all sorts are needed to extend coprolite analyses and interpretations, particularly from pollen-ingestion studies conducted with more volunteers over long periods of time. Parasitological studies of human coprolites will benefit from experimental data to determine the fate of the constituents of human feces ingested by dogs. The application of specialized techniques at the microscope, such as Intensive Systematic Microscopy ([Dean, G., 1998. Finding a needle in a palynological haystack. In: Bryant, V.M., Wrenn, J.H. (Eds.), New Developments in Palynomorph Sampling, Extraction, and Analysis. Am. Assoc. of Stratigraphic Palynologists Foundation., Contribs Series No. 33, pp. 53-59]), to locate and quantify rare pollen types needs to be explored. Easy and useful ways to express the abundance of macroremains in a coprolite are also needed.     

A tick from a prehistoric Arizona coprolite

Ticks have never been reported in archaeological analyses. Here, we present the discovery of a tick from a coprolite excavated from Antelope Cave in extreme northwest Arizona. Dietary analysis indicates that the coprolite has a human origin. This archaeological occupation is associated with the Ancestral Pueblo culture (Anasazi). This discovery supports previous hypotheses that ticks were a potential source of disease and that ectoparasites were eaten by ancient people.     

A taphonomic study of modern pollen assemblages from dung and surface sediments in arid environments of Spain

Coastal plant communities in arid southeastern Spain are characterized by insect-pollinated scrub species, which fail to occur in Quaternary pollen sequences from valleys, marshlands and marine cores. We investigate pollen–vegetation relationships in samples from soil surfaces, animal dung, and sediments in depressions or basins that, in theory, should have pollen spectra that are comparable to those from sedimentary basins elsewhere. Pollen spectra from basins or depressions are very susceptible to long-distance wind and water transport. This can mask representation of pollen from the surrounding insect-pollinated vegetation, as can over-representation of basin-margin halophilous and hydrophilous pollen. Pollen spectra from biogenic materials of animal origin are the best analogues of local and regional vegetation, and show the best analytical potential in terms of pollen concentration and taxon diversity. Pollination properties of the species studied indicate they will rarely be found in most conventional pollen records. It cannot be stressed too strongly that insight into Quaternary vegetation of arid regions demands complementary pollen analysis of coprolites, urine-cemented deposits, and cave sediments with preserved biotic remains, in addition to water-lain sediments.     

Multi‐proxy analyses of Late Cretaceous coprolites from Germany

A total of 462 coprolites from three localities exposing Upper Cretaceous deposits in the Münster Basin, northwestern Germany, have been subjected to an array of analytical techniques, with the aim of elucidating ancient trophic structures and predator–prey interactions. The phosphatic composition, frequent bone inclusions, size and morphology collectively suggest that most, if not all, coprolites were produced by carnivorous (predatory or scavenging) vertebrates. The bone inclusions further indicate that the coprolite producers preyed principally upon fish. Putative host animals include bony fish, sharks and marine reptiles – all of which have been previously recorded from the Münster Basin. The presence of borings and other traces on several coprolites implies handling by coprophagous organisms. Remains of epibionts are also common, most of which have been identified as the encrusting bivalve Atreta. Palynological analyses of both the coprolites and host rocks reveal a sparse assemblage dominated by typical Late Cretaceous dinoflagellates, and with sub‐ordinate fern spores, conifer pollen grains and angiosperm pollen grains. The dinoflagellate key taxon Exochosphaeridium cenomaniense corroborates a Cenomanian age for the Plenus Marl, from which most studied coprolites derive. The findings of this study highlight the potential of a multi‐proxy approach when it comes to unravelling the origin, composition and importance of coprolites in palaeoecosystem analyses.     

家养食草动物粪便中的花粉及菌孢子类型及其对人类活动的指示意义

考古遗址中动物粪化石中的微体植物遗存是重建过去生态环境以及人类活动的重要材料,然而对于能够指示人类活动的粪化石孢粉组合类型及其特征仍然缺乏清楚的认识。本文研究了山羊(Capra aegagrus)、绵羊(Ovis aries)、牛(Bos taurus)、骆驼(Camelus)、牦牛(Bos grunniens)和马(Equus caballus)等几种中国常见的家养食草类动物粪便中的主要孢粉和真菌孢子类型。通过分析这些动物粪便的花粉组合特征,探讨了花粉和菌孢子等微体植物遗存对生态环境及人类活动的指示意义。花粉种类丰度低、一些特征种属花粉(如禾本科、藜科)含量高是家养食草类动物粪便花粉组合的主要特征,可以用于指示人类的饲养行为。家养食草动物粪便中的粪生真菌孢子类型主要有Sporormiella,Sodaria,Pleospora,Coniochaeta,Thecaphora and Dictyosporium。研究表明真菌孢子的分布主要受到动物活动范围的影响,这使得利用粪生菌孢子(如Sporormiella)并结合其他记录来反映人类的饲养或游牧活动成为可能。     

Vegetation reconstruction on the basis of pollen in Late Pleistocene hyena coprolites from San Teodoro Cave (Sicily, Italy)

Vegetation reconstruction based on pollen from coprolites of extinct spotted hyena (Crocuta crocuta spelaea, Goldfuss 1832) recovered from excavations carried out in 1998 at San Teodoro Cave (Sicily, Italy) supports previous indications of pre-Late Glacial conditions. Eight of the twelve coprolites analysed contained well-preserved pollen grains. There is a general similarity between the pollen contents from the coprolites but they show variability. They suggest a main vegetation type dominated by steppic taxa (Poaceae, Artemisia, Ephedra, Chenopodiaceae and Asteraceae) but also including arboreal taxa (Pinus and Cupressaceae). Low percentages of pollen of mesophilous woody taxa (Quercus, Betula, Abies, Alnus, Pistacia, among others) are noticeable, suggesting the existence of nearby refugia for temperate and Mediterranean vegetation. A reconstruction of the landscape, using the coprolite pollen record and other pollen records from Sicily and south Italy, shows the predominance, during the pre-Late Glacial, of a wooded steppe biome, with elements representing a variety of local environmental conditions.     

High-resolution coproecology: using coprolites to reconstruct the habits and habitats of New Zealand's extinct upland moa (Megalapteryx didinus)

Knowledge about the diet and ecology of extinct herbivores has important implications for understanding the evolution of plant defence structures, establishing the influences of herbivory on past plant community structure and composition, and identifying pollination and seed dispersal syndromes. The flightless ratite moa (Aves: Dinornithiformes) were New Zealand's largest herbivores prior to their extinction soon after initial human settlement. Here we contribute to the knowledge of moa diet and ecology by reporting the results of a multidisciplinary study of 35 coprolites from a subalpine cave (Euphrates Cave) on the South Island of New Zealand. Ancient DNA analysis and radiocarbon dating revealed the coprolites were deposited by the extinct upland moa (Megalapteryx didinus), and span from at least 6,368±31 until 694±30 (14)C years BP; the approximate time of their extinction. Using pollen, plant macrofossil, and ancient DNA analyses, we identified at least 67 plant taxa from the coprolites, including the first evidence that moa fed on the nectar-rich flowers of New Zealand flax (Phormium) and tree fuchsia (Fuchsia excorticata). The plant assemblage from the coprolites reflects a highly-generalist feeding ecology for upland moa, including browsing and grazing across the full range of locally available habitats (spanning southern beech (Nothofagus) forest to tussock (Chionochloa) grassland). Intact seeds in the coprolites indicate that upland moa may have been important dispersal agents for several plant taxa. Plant taxa with putative anti-browse adaptations were also identified in the coprolites. Clusters of coprolites (based on pollen assemblages, moa haplotypes, and radiocarbon dates), probably reflect specimens deposited at the same time by individual birds, and reveal the necessity of suitably large sample sizes in coprolite studies to overcome potential biases in diet interpretation.     

The importance of interactions between meteorological conditions when interpreting their effect on the dispersal of pollen from homogeneously distributed sources

The interaction between different meteorological conditions recorded in Badajoz (SW Spain) over a 13-year period has been studied. The results were used for analysis of the effect of these conditions on atmospheric concentrations of Plantago and Brassicaceae pollen grain. For both types of pollen there was a positive effect of NE winds and a negative effect of SW winds which could not be explained by the location of the sources, because these species are very abundant all around the sampling station. The two effects were interpreted on the basis of the action of these winds on temperatures and relative humidity, because these conditions physically and biologically directly determine the concentrations of these types of pollen. Wind from the west is associated with an increase of wind speed and a decline in calm periods; wind from the south is associated with increased rainfall and humidity; wind from the opposite directions is associated with a decrease in these meteorological conditions. These main associations can explain the variation in pollen concentration, although there are also variations in seasons and years. These types of pollen also behave differently, so wind speed reduces the concentration of pollen from Plantago but increases it from Brassicaceae. Beyond the immediate application of the analysis to this specific case, there is a clear interest in observing the interaction between meteorological factors to gain a broader perspective for interpretation of the results of aerobiological studies aimed at determining the effect of these factors on the release and transport of particles into the air.     

Use of plots to define pollen-vegetation relationships in densely forested ecosystems of Tropical Africa

Modern soil samples from South Congo were analyzed for pollen content and compared to forest inventories to define modern pollen-vegetation relationships. A correspondence analysis (CA) was applied independently to botanical and pollen data and a hierarchical cluster analysis to pollen data only. Subsequently, a CA using a presence-absence approach has been made to directly compare the two types of data. Results show that the pollen rain and floristic composition of the sampled sites are not directly linked to altitudinal or precipitation gradients, but clear evidence of variation in relation to hygromorphy and soil type is detected. The forests occurring in swampy environments are well differentiated from the forests developed on well-drained soils by pollen and floristic data. Among forests on well-drained soils, a good distinction can be made between those growing on sandy soils and those growing on ferralitic soils. The comparison between pollen spectra and vegetation shows site-to-site variations in pollen assemblages in relation to the floristic heterogeneity of forests, and it appears that few taxa show a good correlation between plant cover and pollen abundance.     

DISC ELECTROPHORETIC STUDY OF POLLEN PROTEINS FROM NATURAL POPULATIONS OF BETULA POPULIFOLIA IN NEW JERSEY

Pollen proteins from individual trees of eight natural populations of B. populifolia Marsh. located in diverse physiographic provinces of New Jersey were separated by disc electrophoresis with 7 % acid and basic acrylamide gels. Specific stains and substrates for detecting general proteins, esterase, and leucine aminopeptidase isoenzymes were utilized. Average Rp values, 95 % confidence intervals, and percentage similarity values were calculated. Dendrograms were constructed to illustrate protein similarity among populations. The effects of increasing the concentration of protein samples subjected to electrophoresis and storage of pollen (4 C over 16 months) were investigated. After statistical analysis we concluded that: each protein system investigated revealed different amounts of intra- and interpopulation variation, different population ranking (determined by amount of variation), and different population similarity groupings; conclusions based on the comparisons of numbers of protein bands were not always valid; disc electrophoretic data revealed information about natural selection and gene flow; the effect of increasing pollen protein concentration was quantitative; with storage pollen proteins increased in overall positive charge.     

Forecasting airborne Platanus pollen in the Madrid region

This study analyses the atmospheric concentration of Platanus pollen in four stations in the Madrid region over a period of 10 years (1994–2003). Various statistical analyses (regression analysis and decision tree) were used to prepare a forecasting model for possible application as a preventive measure in pollinosis. The data comes from the PALINOCAM network and the samplers used were Hirst type (Burkard pollen trap). Platanus pollen is present in the atmosphere during a short period of time in spring, and the maximum concentrations are detected during the last two weeks of March and the first week of April. Regression analysis shows that the pollen concentration of the two previous days is the best predictive variable. The models obtained for the four stations analysed account for between 37 and 61% of the variation in pollen levels in the air. The decision trees show how the introduction of meteorological variables improves prediction for this pollen type.     

An aeropalynological study of metropolitan Toronto

This study uses 6 years of atmospheric pollen data to examine temporal variability of airborne pollen concentrations at various scales. Airborne pollen was collected from 1985 to 1990 with a Burkard trap, located 18 m above ground at Scarborough College, Toronto, Canada. Pollen season parameters are defined and summarized for all taxa in preparation for developing forecasting models. Annual totals of pollen concentration show great interannual variability. The highest coefficient of variation occurs inTsuga, Fraxinus, Betula andFagus, while the lowest inQuercus andAmbrosia. Some taxa show periodic cycles consistent with mast reproductive behaviour. In many studies, the start of the pollen season is defined as an arbitrary percentage of the annual sum. As a result, the start of the season cannot be identified until the season has passed. As well, due to large fluctuations in annual sum, start dates are more variable. This is not practical for the purposes of forecasting. In this study, the start of the pollen season is defined by a critical concentration threshold which signals the onset of the main pollen season in all years. These critical levels ranged from 2 to 60 grains/m³ for the abundant taxa. Interannual variation in the start of the season is approximately 20 days for tree taxa, 5 days for Poaceae, and 2 days forAmbrosia. For many plants, dehiscence is triggered at a critical level of accumulated degree-days. Since annual rates of temperature increase show great variation, there is also interannual variability in the onset of pollen release. Multi-year average pollen curves incorporate these differences in onset and may give an inaccurate representation of the pollen season in a typical year. This paper presents a method of aligning yearly pollen curves to reduce seasonal variation and more accurately represent both the average timing and magnitude of the pollen season. For some types, such asBetula and Poaceae, the resulting curves are positively skewed. Tree taxa, in general, exhibit a more symmetric pollen concentration curve. Aligned average pollen concentration curves are presented for Toronto in the form of a pollen calendar. In addition, phenological data for all common taxa are summarized.     

Intradiurnal fluctuation of pollen in La Plata, Argentina. Part I, herbaceous pollen types

A pollen analysis was carried out on six herbaceous pollen types whose annual concentration represents a percentage higher than 0.3% in the atmosphere of the city of La Plata, Argentina, from July 1998 to June 2001. They are: Ambrosia, Cyperaceae, Chenopodiaceae-Amaranthaceae, Plantago, Poaceae and Urticaceae. Hourly patterns of the pollen types analysed with the intradiurnal daily index (IDI) could be grouped into three categories account the percentage that represent of pollen concentration registered during five hourly periods of the day. Group I includes pollen types that show a sharp peak in abundance gust one at hour of the day. Group II is formed by pollen types that show high concentrations in more than one hourly band during the daylight. Group III is characterized by the pollen types that show lower concentrations over a broad hourly band during the day.     

The diet and ecology of Hawaii's extinct flightless waterfowl: evidence from coprolites

At least five species of large flightless waterfowl have become extinct in the Hawaiian Islands in recent millennia. These birds are thought to have occupied the role of large herbivores in a wide range of terrestrial habitats. A collection of coprolites from one of the species ( Thambetochen chauliodous ) was obtained during excavations in Holocene cave sediments on the island of Maui. The chemical composition and pollen and spore content of the coprolites are analysed and compared with pollen/spore spectra from the cave sediments and from recent goose scats. The results support the contention that these birds were primarily folivorous, and further suggest that ferns were important in the diet. The coprolites have a very fine texture that may result from efficient hindgut fermentation and digestion of plant fibre. Our data are discussed in the light of a recent hypothesis of plant/herbivore coevolution between extinct avian herbivores and native Hawaiian lobelias. The loss of large native herbivores, as well as other changes in vertebrate trophic structure due to extinctions over the past few thousand years, may still be affecting ecological processes in areas of the Hawaiian islands with native vegetation.     

Plant microfossil analysis of coprolites of the critically endangered kakapo (Strigops habroptilus) parrot from New Zealand

We present here the results of the analysis of pollen, spores, starch and other microscopic plant material in 52 coprolites of the critically endangered kakapo (Strigops habroptilus), a New Zealand endemic parrot. The six sampling sites (caves, rock overhangs) are in widely separate geographic regions in the South Island, across an altitudinal range encompassing lowland, montane and sub-alpine environments. Radiometric testing gives average ages of discrete coprolite groups ranging from modern to 2514 ± 43 ¹⁴C yr BP (Wk-19164). A wide variety of plant taxa is identified in the coprolites, comprising cones, flowers, leaves and fronds of ~ 30 podocarps, dicotyledonous trees, shrubs and lianas, grasses, ferns, lycopods, mosses and liverworts. This supports previous observations that kakapo are versatile feeders, using a broad spectrum of foods that may only be available for short periods and intermittent years. Many of the food taxa we identify are consistent with the studies of modern kakapo diet. We also identify several new foods, potentially of value in developing food supplements to increase breeding frequency of the current kakapo population. The most commonly identified plant parts in coprolites were fern fronds (monoletes, Hymenophyllum, Cyathea smithii), followed by podocarp cones and leaves. Lack of starch in the coprolites suggests that the lycopod and fern rhizomes reported in modern kakapo diet were not commonly eaten. The data provide initial botanical evidence of diet from coprolites of an extant bird.     

Critical analysis of coprolite evidence of medicinal plant use, Piauí, Brazil

Human coprolites, dating from 8500 to 7000 years BP, were collected from the rock-shelter of Boqueirão da Pedra Furada, in Piauí, Brazil. These dates fall within the “Serra Talhada” cultural traditions I and II of the Tradição Nordeste, a paleohuman culture of northeastern Brazil. Archaeoparasitological analysis of the coprolites revealed whipworm eggs and hookworm eggs. Analysis of Tradição Nordeste skeletons and hair shows a variety of diseases including dental problems, osteoarthritis, and head louse infestation. Palynological analysis of the coprolites revealed 12 genera that were potentially medicinal. The pollen data were critically analyzed to assess the potentiality that the pollen represented medicinal use. Strong cases could be made that three of the genera, Anacardium, Borreria, and Terminalia, were actually used for medicinal purposes, probably to treat symptoms caused by the intestinal parasites.     

The relationship between pollen and extant vegetation across an arid urban ecosystem and surrounding desert in Southwest USA

Aim In this paper, we explore the relationship between pollen concentration in surface soil and extant perennial vegetation across a rapidly urbanizing arid ecosystem along with its surrounding agricultural and native Sonoran Desert land. We also investigate whether taxa behave as expected from known pollination characteristics (e.g. anemophilous vs. zoophilous) and whether the pollen–plant relationship differs between the undeveloped desert and agro‐urban regions. Location We used a probability‐based sampling scheme at 200, 900‐m² sites across the Central Arizona–Phoenix region, a 6400‐km² area of the south‐western USA. Methods Pollen–plant abundances were mapped across the study area using interpolation techniques, summary bar charts and scatterplots, then two statistical approaches were applied to examine the data quantitatively. Firstly, we used regression analysis of paired pollen concentration and plant cover data; secondly, pc‐ord was used on a cross‐matrix (contingency table) containing the presence/absence scores of both pollen and plants for each taxon at each site. Results Mapping and quantitative analysis revealed that pollen–plant relationships do vary both between anemophilous and zoophilous groupings and by individual taxon. They also revealed that distribution patterns of both pollen and plant abundance vary consistently across the three main landscape types (desert, urban, agricultural) and that pollen taxa groupings are consistent with pollination method and source plant distribution regardless of landscape type. Main conclusions These findings provide a broad range of comparative data to facilitate palynological reconstruction of past vegetation and aid in assessing types of prehistoric impacts on this vegetation. They also document that significant variation can occur in pollen rain across a city, even for taxa with widely distributed pollen, leading to the conclusion that forecasting pollen distribution patterns for allergy‐related pollen types using only one or a few pollen traps is likely to entail substantial error.