Sunday, April 21, 2013

"Do not date" list: radiocarbon dating of macroscopic plant remains


by Jimmy Marty

Establishing accurate sediment core chronologies is of great importance for the interpretation of paleolimnological records. Radiocarbon dating is the primary method used to create chronologies in studies focusing on the past 40-50,000 years. Described here are various macroscopic plant materials, categorized according to their suitability for radiocarbon dating. Materials were evaluated based on the characteristics of good dating material: a) all fixed CO2 is in equilibrium with atmospheric CO2 and b) the material accurately represents the time of sediment deposition.

Sources of error

A common age-altering error encountered in paleolimnology is the old-carbon error, which occurs when dating material has fixed carbon out of equilibrium with the atmosphere. Old carbon reservoirs are commonly associated with hard-water lakes with high concentrations of HCO3-, but sediment and aquatic CO2 are often old-carbon enriched due to isotopic exchange between carbon species (Mook 1980), infrequent mixing (in lakes with high surface area-depth ratios) (Hakkanson 1979; Olsson 2009), and respired CO2 derived from HCO3- fixing organisms (MacDonald et al 1987). This list does not include processes in arctic and volcanic regions, which may be old-CO2 enriched for numerous other reasons (see Bjorck and Wohlfarth 2002).
           
Physical processes such as reworking of old sediment by means of erosion or bioturbation can redistribute plant remains up or down the sediment profile (Turney et al 2000; Bjorck and Wohlfarth 2002). Sediment disturbance is often clear through visual examination of the sediment core face, in which slumps and/or turbation may be evident. Further examination of microscopic composition using smear slides can reveal less obvious terrestrial material indicative of erosional input. Describing these macroscopic and microscopic features of the sediment is of great value to the selection of suitable emergent and floating leaved macrofossils and can help avoid erroneous dates.

Appropriate sample storage techniques should always be performed in order to avoid contamination errors. Sediment cores and macrofossil samples stored in cool and wet conditions for several months or more are susceptible to modern fungal and microbial contamination (Wohlfarth et al 1998). After isolating the macrofossil sample from the sediment, it should be stored in distilled water in a cold room for no longer than one week before submission for dating (Bjorck and Wohlfarth 2002).


Dateable: (click on heading for more comprehensive list of taxa in this category)

Terrestrial plant macrofossils

Terrestrial plants fix only atmospheric CO2 and are considered ideal for radiocarbon dating. Care should still be taken to ensure the material has not been reworked through sediments.
 
 
Pine needle (terrestrial) of Abies balsamea. Dateable.


Seeds of terrestrial tree Betula alleghaniensis. Dateable.

Seeds of terrestrial herb Rudbeckia triloba. Dateable.

Leaf of terrestrial bog shrub Chamaedaphne calyculata. Dateable.

Bud scales of terrestrial tree Populus tremuloides. Dateable.


Charcoal (from grassland regions)

Grassland vegetation fixes only atmospheric CO2. Grassland vegetation has a short terrestrial residence time; it is without the in-built age present in long-lived, slow-decomposing tree species that may remain on the landscape for significant lengths of time (Oswald et al 2005; Grimm 2011).


Emergent-aquatic plant macrofossils



Emergent macrophytes have a strong history of providing good dates (Deevey et al 1954; Heikkinen et al 1977; Hakansson 1982; Tornqvist et al 1992; Lowe et al 2004; Grimm 2011). Most are unable to fix old carbon in the form of HCO3- (Spence and Maberly 1985; Sand-Jensen et al 1992; Maberly and Madsen 2002), and are only able to utilize a very small portion of CO2 absorbed from the sediment and water. Yet there are some cases of emergent vegetation returning dates that are too old (Damon et al 1964; Haynes et al 1966; Turney et al 2000; Wasylikowa and Walanus 2004; Edwards et al 2011). Furthermore, Olsson (1983) demonstrated that emergent and floating leaved vegetation have lower 14C activities than terrestrial vegetation, though not as low as submerged aquatics.

The physiological literature shows that in some situations it is possible for old-carbon to be incorporated into emergent vegetation if there is an old-CO2 reservoir present. Amphibious isoetids such as Lobelia dortmanna and Juncus bulbosus (the seeds of which can be easily confused with other Juncus species) acquire the majority of their CO2 from the sediment via the roots (Wium-Anderson 1971; Winkel and Borum 2009). Recent evidence shows that root uptake of CO2 may not only be limited to isoetids. In a hard water lake, Sparganium angustifolium utilized sediment CO2 in amounts that significantly affected productivity (Lucassen et al 2009). Stratoides aloides is able to utilize HCO3- in significant amounts, the only known amphibious plant that can do so (Prins and De Guia 1986). Additionally, new studies continue to suggest that emergent plants may be able to fix aquatic CO2 when submerged, albeit in unknown quantities. Gas film layers on leaves of emergents like Typha, Phalaris, and Phragmites allow the utilization of aquatic CO2 (Colmer and Pedersen 2008). Morphological plasticity of leaves and roots permit other amphibious macrophytes to adjust CO2 fixation depending on the environment (Mommer et al 2007; Rich et al 2012).

So, are the macrofossils of emergent plants dateable? While there is a short list of clearly unsuitable taxa that includes isoetids and Stratoides aloides, it appears that most other emergent macrophytes only utilize old carbon under very particular circumstances and/or in very small or unknown quantities. When determining if an emergent macrofossil is suitable for dating, it is extremely important to carefully consider the study environment and the ecophysiological characteristics of the plant material. For example, if there is old CO2 present in a lake, the suitability of Sparganium fruits for dating is most likely dependent on whether the lake of study is hard or soft water.


Seeds of emergent aquatic macrophyte Lobelia dortmanna (water lobelia). This species absorbs the majority of  photosynthetic CO2 from the sediment (Wium-Anderson 1971). Do not date.

Seeds of emergent/submerged aquatic macrophyte Juncus pelocarpus  (brown-fruited rush).  This species of rush has both emergent and submerged forms, and a relative, Juncus bulbosus, fixes mostly sediment CO2 (Wetzel et al 1985). Date with caution.


Seeds of emergent aquatic macrophyte Schoenoplectus acutus (hard stem bulrush). A European member of this genus (Schoenoplectus tabernaemontani) internalizes large amounts of sediment CO2, but only fixes very small quantities (Singer et al 1994). Date with caution.

Fruits of emergent aquatic macrophyte Sparganium angustifolium (narrow leaf bur-reed). This species can utilize significant amounts of sediment CO2 in hard water conditions. Date with caution.

Spikelets of emergent aquatic macrophyte Zizania palustris (northern wild rice). Species of Asian rice, of the same tribe as Z. palustris, are adept at the uptake of sediment CO2 (Higuchi et al 1984). Date with caution.


Seeds of amphibious macrophyte Eleocharis acicularis (needle spikerush). E. acicularis can grow under submerged or emergent conditions. A similar amphibious spikerush, Eleocharis vivipara uses aquatic CO2 when submerged (Ueno et al 1988). Date with caution.

Seeds of emergent aquatic herb Polygonum amphibium (water smartweed). P. amphibium is able to produce both floating and aerial leaves based on water level conditions. Related species of the genus Rumex can produce submerged leaves that utilize aquatic CO2 (Mommer et al 2007). Date with caution. 

Seeds of emergent wetland plant Carex comosa (bristly sedge). There is flow of CO2 from root to shoot in some species of Carex, although it may not be fixed and appears to be respiratory (Koncalova et al 1988). Date with caution.

 Floating-leaved-aquatic plant macrofossils

Floating-leaved macrophytes comprise a much smaller number of taxa than emergent macrophytes, with the majority of research primarily involving nymphaeids (e.g. Nymphaea, Nuphar). Much like emergent macrofossils, macrofossils of floating leaved macrophytes have provided good dates in the past (Heikkinen et al 1977; Tornqvist et al 1992) and are unable to fix old carbon in the form of HCO3- (Spence and Maberly 1985; Sand-Jensen et al 1992; Maberly and Madsen 2002). This is not the only similarity nymphaeids share with emergent macrophytes; the suitability of nymphaeids for dating has also been questioned (Olsson 1983; Tornqvist et al 1992; Olsson 2009). Physiological evidence suggests that Nuphar may take up CO2 from the sediment, which is potentially problematic if the CO2 is derived from an old-CO2 reservoir. In Nuphar lutea, relatively large concentrations of CO2 move from the rhizome to the upper parts of the plant where it is then fixed (Dacey and Klug 1982). The source of the CO2 is unknown and could be from either the sediment or a product of respiration in the rhizome, the latter of which is a product of atmospheric O2 (Dacey and Klug 1982). Because the partial pressure of CO2 exceeds that of the partial pressure in the rhizome, it is suggested that the CO2 is acquired from the sediment (Dacey 1979). However, the amount of CO2 fixed through this pathway is unknown, and may be insignificant in the context of radiocarbon dating. Like with emergent macrofossils, one must evaluate the study environment carefully to gauge the suitability of nymphaeid remains for dating. If there is an old CO2 reservoir available, proceed dating nymphaeids with caution.

Seeds of floating-leaved aquatic macrophyte Nuphar lutea (yellow water lily, spatterdock). Date with caution.

 
Seeds of floating-leaved macrophyte Nymphaea odorata (white water lily). Date with caution.


Seeds of floating-leaved macrophyte Brasenia schreberi (watershield). Date with caution.
 
 
 
Wood charcoal

Woody plants fix only atmospheric CO2. However, in some cases wood charcoal may provide inaccurate dates due to long terrestrial residence times (Oswald et al 2005; Grimm 2011). Woody species may take many years to decay and therefore can remain on the landscape for long after atmospheric CO2 was originally fixed. Interpret dates from wood charcoal with care.

Wood fragments

Like wood charcoal, wood may experience long terrestrial residence times and provide inaccurate dates despite use of atmospheric CO2 (Oswald et al 2005; Grimm 2011). Furthermore, old carbon enriched material may adhere to the rough surface texture of wood if the sample is not processes correctly (Oswald et al 2005). For the best dates of woody material, look for identifiable twigs of lakeshore species to ensure near-site deposition.



Submerged-aquatic plant macrofossils

It is well documented that macrofossils from submerged macrophytes and aquatic mosses return radiocarbon dates that are spuriously old (Deevey et al 1954; Hakkanson 1979; Olsson 1983; Birks 2002). Most submerged aquatic macrophytes are able to take up aquatic HCO3- derived from carbonate bedrock that results in radiocarbon dates many thousands of years too old (Spence and Maberly 1985; Sand-Jensen et al 1992; Maberly and Madsen 2002). Aquatic mosses and some submerged macrophytes do not utilize HCO3-; rather they fix CO2 from the sediment (e.g. isoetids) and/or water (e.g. Vallisneria americana), which may still carry an old CO2 reservoir effect (Sand-Jensen et al 1992; Maberly and Madsen 2002). In very well mixed, soft water lakes, it may be possible to date both submerged macrophytes and aquatic mosses (Hakansson 1979; Miller et al 1999; Oswald et al 2005). However, complete absence of a reservoir effect is rare, and dating macrofossils of submerged macrophytes and aquatic mosses is generally unadvisable (Olsson 2009).


 

Seeds of submerged aquatic macrophyte Potamogeton epihydrus (ribbonleaf pondweed). Do not date.

Seeds of submerged aquatic macrophyte Vallisneria americana (water celery). Do not date.

 
Seeds of submerged aquatic macrophyte Najas flexilis (nodding waternymph). Do not date.

Seeds of submerged aquatic macrophyte Callitriche palustris (water starwort). Do not date.

 
Oogonia of submerged aquatic macroalgae Chara sp. Do not date.  

Seeds of submerged aquatic macrophyte Ranunculus aquatilus (white water crowfoot). Do not date.

Aquatic moss fragments (unknown species). Do not date.

References cited



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