1. GEOL 553 LECTURE 21 Biological Evidence Microfossils Pollen Diatom
Macrofossils
Plants
Insects
Mollusca & Ostracoda & Foraminifera & Coccolithophores
Mammalia
Lab: NSF Grant Writing
Discussion: NSF Grant Proposal Guide

3. Micropaleontology deep sea Vertebrate remains
Biological Records of Climate Change
Nature of Record:
Macrofossils
Microfossils
Biomolecular (DNA)
Taphonomy
In-situ
Selective transport
Differential degradation
Interpretation of Assemblages
Modern Analogue
Ecosystems
Paleobiology: single vs. multi proxy
Pollen
Diatom
Plant
Insect
Molusca
Foraminifera
Micropaleontology deep sea
Vertebrate remains
Biological evidence, in the form of plant and animal remains, has always been a cornerstone in the recon struction of Quaternary environments. The analysis of fossil evidence employs uniformitarian principles, namely that knowledge of the factors that influence the abundance and distribution of contemporary organisms enables inferences to be made about the dominant environmental controls on plant and animal populations in the past. Applying this approach to the interpretation of Quaternary fossil assemblages, therefore, the majority of which have living counterparts, it should be possible to reconstruct former environmental conditions with a reasonable degree of confidence. The use of modern ecological information in this way is an essential element of palaeoecology, the study of the interrelationships of organisms in the past, both with their physical environment and with other plants and animals (Battarbee, 2000; Birks & Birks, 1980, 2006). Variations in the type and diversity of plant and animal remains preserved within sedimentary sequences are also used to subdivide the geological record, a field of study known as biostratigraphy. This chapter is concerned almost entirely with the palaeoecological aspects of biological evidence; the principles and practices of biostratigraphy are considered more fully in Chapter 6.

4. Biological Records of Climate Change: Diatom
procedures employed to derive quantitative lake-water pH reconstructions from fossil diatom records
Figure 4.9 Schematic representation of the procedures employed to derive quantitative lake-water pH reconstructions from fossil diatom records (from Birks & Seppa, 2004, but based on an unpublished diagram by Steve Juggins, Newcastle University, UK). Although this schematic relates to diatoms, the procedure can be adapted to quantify the relationships between other biological proxies and environmental variables, for example pollen assemblages and climate.

5. BIDI = Brackish Intertidal Diatom Index
0 – 1 0 = lower elevation
1 = higher elevation

6. Figure 4.17 Arboreal plant macrofossils (shown as dots) in a Lateglacial sediment sequence in Latvia, plotted against the pollen record of the same taxa (from Veski et al., 2012).
The results of plant macrofossil analysis can be presented in a number of different ways. At many sites, particularly where archaeological investigations are being carried out or where a single stratum is being investigated, a simple species list is compiled of all taxa discovered. Where several levels are being examined, the presence or absence of particular plant remains may be indicated by simple dot symbols (Figure 4.17) or the results may be shown in tabular format. Alternatively, the data may be expressed as estimates of abundance using such descriptive terms as rare, occasional or frequent, and thus an impression can be gained of changes in frequency of taxa through time.

7. Biological Records of Climate Change: Insect Remains
Subfossil Coleopteran sclerites recovered by flotation from peat deposits overlying a Bronze Age occupation site near Ballyarnet Lake, Co. Derry, Northern Ireland.
Figure 4.21 Subfossil coleopteran sclerites (see Figure 4.22) recovered by flotation from peat deposits overlying a Bronze Age occupation site near Ballyarnet Lake, Co. Derry, Northern Ireland. Elytra (wing-cases) dominate in this view with an occasional pronotum (p). 1. Pterostichus nigrita – a predatory ground beetle (10–15 mm) widespread in marshes and water margins. 2. Cercyon atomarius – a small (1.5–4.5 mm) dung beetle. 3. Helophorus spp. – a water scavenger genus, with most species occupying emergent vegetation in standing or slow-moving water. 4. Trechus rubens – a small (5–6.5 mm) subterranean ground beetle, under stones and leaf piles on riverbanks and lake shores. 5. Anobium punctatum – a beetle that bores into and feeds upon wood (3–4.5 mm). 6. Hydraena britteni elytrum (6e) and pronotum (6p) – a water beetle common in shallow lake margins (photograph by Nicki Whitehouse, Plymouth University, UK).

8. Late Glacial South Wales, UK Warmest = FU-2 Cooler = FU-3
Coldest = FU-4
(Younger Dryas)
Thermophiles FU-5
Figure 4.23 A coleopteran record from the Lateglacial (c. 14.7–11.5 ka) site of Llanilid, south Wales, UK. The warmest part of the sequence (early Interstadial) is represented by faunal unit FU-2, cooler conditions are recorded in FU-3 (later Interstadial) and marked by the arrival of cold-adapted species, while the cold Younger Dryas/Loch Lomond Stadial is characterized by an increased number of cold-adapted species. Note how these disappear in the early Holocene (FU-5) to be replaced by thermophiles (from Walker et al., 2003).

9. Augment Palynological Analyses
Beetles v Trees
Central England
Augment Palynological Analyses
Figure 4.24 Number of obligate beetle species (key indicates tree affinities) recorded in Holocene sediment sequences from southern and central England. The site names are listed along the base; archaeological sites are indicated by closed circles; the remainder are considered ‘palaeoecological’ (not disturbed by archaeological activities). For further explanation see text (from Whitehouse & Smith, 2010).

10. Present-day European distributions of four coleopteran species found in Lateglacial deposits.
Figure 4.25 Present-day European distributions of four coleopteran species found in Lateglacial deposits (c. 14.7–11.5 ka) at the site of Glanllynnau, north Wales (from Coope & Brophy, 1972
One of the most important factors that has governed the distribution of most insect species during the Quaternary has been climate, particularly thermal conditions (Coope, 1990, 2004). Distribution maps of modern beetles show that the geographical range of many species corresponds with well-defined climatic zones (Figure 4.25) and especially with summer temperature thresholds. Those insect species whose distributions are narrowly restricted are termed stenotherms, while those that can tolerate a broader range of climatic conditions are termed eurytherms. The former are much more important in palaeoclimatic research, since they enable more precise inferences to be made about
former temperature regimes. The acute sensitivity of beetles to temperature variations is reflected in the growing body of evidence that reveals how beetles have adjusted their modern geographical ranges in response to global climate change within the last century or so (e.g. Parmesan, 1999; Hill et al., 2011).
As with most other lines of fossil evidence, however, there are problems in utilizing fossil beetles as climatic proxies.
It can never be established for certain that an insect species has colonized the entire climatic range to which it is suited, nor
that past distributions were entirely in equilibrium with the prevailing climatic conditions.

11. Mutual Climate Range method MCR Test
Figure 4.26 Schematic representation of the mutual climatic range (MCR) method of quantitative temperature reconstructions (courtesy of Adrian Walkling).
Figure 4.27 Test of the MCR method on assemblages of species found living today at thirty-five localities in North America. The reconstructed mean July temperatures (TMAX) are shown on the vertical axis of graph a); the reconstructed mean January temperatures (TMIN) on the vertical axis of graph b). Vertical bars represent the mutual climatic range of the beetles found living at the localities studied. The horizontal axes of the two graphs show the observed TMAX and TMIN values for the thirty-five sites. The slopes and positions of the gradient lines are linear regressions of predicted against observed values for the test sites (from Elias, 1997).
In an attempt to avoid the errors that may arise from the use of the indicator species approach, the mutual climatic range (MCR) method was developed to obtain more representative palaeotemperature estimates from beetle records (Atkinson et al., 1987). This is an extension of the range overlap method, but it employs the ranges of all of the taxa included. Modern distribution maps are first obtained for as many as possible of the species in the fossil assemblage, and the climatic range of each beetle type is then established using contemporary meteorological data. The two most important variables governing beetle distributions appear to be the temperature of the warmest month (TMAX) and the temperature range between the warm est and coldest months (TRANGE), the latter providing an index of seasonality. By knowing the distribution in terms of TMAX and TRANGE, the geographical range of each species may be plotted in ‘climate-space’, and for each species a ‘climatic envelope’ is thus produced (Figure 4.26). For any fossil assemblage, therefore, the mutual climatic range can be determined from a computer-generated plot of the climatic parameters relating to each beetle in the assemblage. From these plots, the values of TMAX, TRANGE and TMIN (temperature of the coldest month) can be obtained, and these constitute the ‘best estimates’ of the mutual climatic conditions within which the particular mix of fossils formerly coexisted. The method is most successful (i.e. produces the narrowest range estimates) where an assemblage contains a large number of species, and where a number of these are obvious stenotherms. The method can be tested by deriving MCR values for modern beetle assemblages and measuring their statistical relationship with modern climate data (Figure 4.27). The results usually show a stronger linear relationship for
summer than for winter temperatures, mainly because many beetle species can survive a wide range of winter temperatures (Elias, 1997). The advantages of the MCR method over the indicator species approach are that it avoids subjective inter pretations and possible bias, as well as over-generalization from the use of geographical overlays. Moreover, geo graphical range limits are often too broad, and cannot take into account such factors as altitude, oceanicity, micro climatic variations and so on. The MCR approach ignores geographical location, and focuses entirely on climatic parameters governing species distributions. Hence a complex geographical distribution may be reduced to a narrow climatic range, reflecting the fact that the often diverse geographical locations in which a species occurs may, in fact, have common characteristics when plotted in climate–space. Also, and most importantly, it does not really matter if the species does not occupy its full (potential) geographical range, so long as it reaches potential climatic boundaries in a sufficient number of places.

12. Biological Records of Climate Change: Nonmarine Mollusca
<<< Molusca exposed on an abandoned beach of Pluvial Lake Lahontan on the eastern side of Pyramid Lake, Nevada
Figure 4.31 a) Fossil shells of freshwater molluscs (principally gastropods) exposed on an abandoned beach of Pluvial Lake Lahontan (section 2.7.1) on the eastern side of Pyramid Lake, Nevada, USA (photograph by Allan Ashworth, North Dakota State University, USA). b) Valvata piscinalis, a small gastropod (up to 5 mm height and width) that inhabits streams, rivers and lakes, preferring running water and tolerant of low calcium levels. Such small specimens can be readily identified, when magnified, by examination of morphological and ornamental details, some of which are illustrated (photograph by Jenni Sherriff, Royal Holloway, University of London, UK).

13. Biological Records of Climate Change: Nonmarine Mollusca
Figure 4.32 Variations in relative abundance of molluscan species from the Lateglacial to mid-Holocene sequence at Holywell Coombe, southern England. Several taxa have been combined to produce Terrestrial Group A (‘catholic’ species, of wide environmental tolerance, mainly open ground), and Terrestrial Group B (more narrow in environmental tolerance, mainly deciduous woodland). Note how Group B taxa expand in molluscan zone b, which coincides with woodland expansion (Corylus avellana) as reflected in the pollen record (right) (from Preece & Bridgland, 1999).

14. Figure 4.33 Variations in absolute abundance (number of individuals per 15 kg of sediment) of terrestrial molluscan species through a Late Quaternary loess and palaeosol succession at Weinan, China. Note that species abundance and diversity are dependent on weathering intensity (pedogenic alteration). 1: Holocene soil; 2: Loess; 3: Weakly-weathered soil (from Wu et al., 2002).

15. Figure 4.34 Variations in species richness, abundance, diversity and habitat types in Holocene tufa sequences at Courteenhall, near Northampton, UK (from Meyrick & Preece, 2001).

16. Examples of some common marine bivalves of the North Atlantic
Biological Records of Climate Change: Marine Mollusca
Examples of some common marine bivalves of the North Atlantic
Figure 4.35 Examples of some common marine bivalves of the North Atlantic and their water depth preferences. 1. Spisula elliptica – open water, on various substrates, to 100 m depth. 2. Callista chione – continental shelf to 200 m depth. 3. Tapes decussatus – lower shore and shallow sublittoral. 4. Mya arenaria – inter-tidal mudflats. 5. Mya truncata – widely distributed on sandy substrates to 70 m depth. 6. Arctica islandica – widespread on muddy sand substrates from very shallow tidal to (exceptionally) 200 m depth; noted for its longevity. 7. Aequipecten opercularis – typically found on hard surfaces in depths of 20–45 m in shallow subtidal areas, but can extend to 180 m depth. 8. Macoma balthica – open sea in inter-tidal zone, and to 100 m depth in the brackish Baltic Sea (photographs by David Roberts and James Scourse, School of Ocean Sciences, Bangor University, UK).

17. Distribution of marine zoogeographical provinces in the Northeast Atlantic: a) present time; b) during the Eemian (Last) Interglacial.
Figure 4.36 Distribution of marine zoogeographical provinces in the Northeast Atlantic: a) present time; b) during the Eemian (Last) Interglacial. NAC – North Atlantic Current; EGC – East Greenland Current (based on Funder et al., 2002 – province boundaries; and Knudsen et al., 2001 – surface current flows).

18. Figure 4.37 Variations in the dynamics of the winter North Atlantic Oscillation (dotted curve: section ) and of annual growth increments of the bivalve Arctica islandica obtained from the central North Sea and the Norwegian Shelf: all values shown as deviations from long-term means (from Schone et al., 2003).

19. https://en.wikipedia.org/wiki/North_Atlantic_oscillation

20. Biological Records of Climate Change: Foraminifera
Figure 4.39 Examples of planktonic foraminiferal species widely employed in Quaternary palaeoceanographical studies. The use of sinistral (1) and dextral (2) forms of Neogloboquadrina pachyderma as palaeoclimatic proxies in high-latitude surface waters is referred to in section Globigerinoides ruber (3) and G. sacculifer (4) are abundant today in subtropical waters (Figure 4.43) and hence employed as proxy indicators of warm SSTs. Globigerina bulloides (5) can tolerate a wide range of temperatures but is most abundant in cool upwelling ocean waters. (SEM images provided by Alessandra Asioli, CNR Institute of Geoscience and Geo-resources, Padua, Italy.)

21. Figure 4.40 Relative sea-level (ERSL) curves for the last 12 ka for the Western Mediterranean based on benthic Foraminifera based transfer functions. Data from three shelves (core 342-1: Alboran Platform; core 367-1: Oran Bight; and core 401-1: Mallorcan Shelf) show broadly similar trends to independent reconstructions of sea-level change over the same period (from Milker et al., 2011).

22. Planktonic foraminiferal provinces in the modern ocean showing the close relationship between sea-surface temperature gradients and species abundances.
Figure 4.43 Planktonic foraminiferal provinces in the modern ocean showing the close relationship between sea-surface temperature gradients and species abundances. The species abundance plots (top) are averaged at 1°C intervals (from Kucera, 2007, reprinted with permission from Elsevier).

23. Figure 4.44 Changes in coiling direction of tests of the highlatitude species Neogloboquadrina. The proportion of rightcoiling specimens increases markedly in surface waters with mean temperature of between 6 and 10°C, reflecting the
replacement of N. pachyderma, which produces mainly sinistral tests, by the dextral-coiling N. incompta (modified from Kucera, 2007).

24. Figure 4.47 Mg/Ca calibration results for several species of planktonic Foraminifera. Temperatures shown are the isotopically derived calcification temperatures; the equation defines the correspondence between temperature and calcification (r = 0.93) (based on Anand et al., 2003).

25. Figure 4.48 Rapid changes in SST in the Alboran Sea (Mediter ranean Sea) over the past 50 ka, inferred from variations in Uk 37 abundance, and correlation with Greenland icecore events; note that marked declines in temperature in the Alboran Sea coincide with Heinrich Events and the Younger Dryas cold stage (from Cacho et al., 1999).

26. Biological Records of Climate Change: Megafauna
Figure 4.49 a) Nigel Larkin cleaning the fossil right femur (length 145 cm) of a mammoth of Middle Pleistocene age discovered at the site of West Runton, Norfolk, UK (photograph by Nigel Larkin, Norfolk Museums & Archaeology Service, Norfolk, UK; from Larkin, 2010, reprinted with permission from Elsevier). b) Occlusal surface (top) and roots of the first lower molar of the ancestral water vole (Mimomys savini) recovered from sediments of early Middle Pleistocene age at Pakefield, Suffolk, UK. Variations in dentition enable fossil vole teeth to be assigned to species and the relative age of temperate stages to be inferred (section 5.5.4). Scale bar represents 1.0 mm (photograph by Harry Taylor & Simon Parfitt, Natural History Museum, London, UK; from Maul & Parfitt, 2010, reprinted with permission from Elsevier). c) Reconstruction of woolly mammoth (Mammuthus primigenius) based on soft tissue, skin, hair, parts of the intestines and delineation of toes recovered from one of the best preserved mammoth carcasses ever discovered, in Yakutia, arctic Siberia (reconstruction and photograph by Remie Bakker of Mammal Works & Dick Mol of the Natural History Museum, Rotterdam, Netherlands; from van Geel et al., 2008, reprinted with permission of Elsevier; Lister & Bahn, 2009).

30. Biological Records of Climate Change: Micropaleontology Deep Sea
Figure 4.41 a) Examples of siliceous skeletons (tests) of the radiolarian groups Spumellaria (S) and Nassellaria (N), important biomarkers commonly preserved in Late Quaternary deep marine sediment; their tests range in size from c. 30–300 μm. 1. Callimitra carolotae (N). 2. Euchitonia elegans (N). 3. Lamprocyclas maritalis (S). 4. Mitrocalpis araneafera (N). 5. Nephrospyris knutheieri (N). 6. Rhizoplegma boreale (S), recently renamed Cleveiplegma (from Dumitrica, 2013; SEM images provided by Kjell Rasmus Bjorklund, Natural History Museum, University of Oslo, Norway). b) SEM scans of coccolithophore specimens from the central Adriatic Sea. 1. Coccosphere of Emiliania huxleyi TYPE A comprising an outer (single) layer of calcitic platelets (coccoliths) which enclose the living organism. This form of the species is typical of nutrient-rich environments and is characterized by rapid growth during bloom conditions (scale bar: 1 μm). 2. Emiliania huxleyi TYPE A with multiple layers of coccoliths, a larger form that is typical of nutrientpoor environments and which grows more slowly (scale bar 1 μm). 3. Complete coccosphere of Calcidiscus leptoporus (ssp. quadriperforatus), an important carbon storage species that thrives in tropical and temperate latitudes (scale bar: 10 μm). 4. Closeup of 3 showing coccolith detail (scale bar: 1 μm) (images provided by Luka Supraha, Uppsala University, Sweden).

31. Figure 4.46 Reconstructions of surface conditions in the Northeast Atlantic during the Last Glacial Maximum (LGM) based on marine microfossil records. a) Summer SSTs based on CLIMAP (1981). b) Summer SSTs based on SIMMAX. c) and d) More detailed reconstructions for LGM salinities and SSTs based on a larger number of palaeo-data sites and calibrated using SIMMAX (from Meland et al., 2005).