47 published journal articles, 14 Ph.D Theses, 10 Masters Theses, 5 technical reports and 3 chapters in published books have been written using Saginaw Forest (SF) as the basis for their research.  The first research conducted on the area was by Reed (1902 after encouragement from Professor Spalding) who examined the successional processes of Third Sister Lake (TSL) with specific emphasis on the plant communities in and around the lake.  Around the same time, Weld (1904) did a similar study on the plant zones of First Sister Lake.  Weld noted that although First Sister Lake and Second Sister Lake used to be connected, TSL was not connected with these two and that all these lakes are fed by springs.  Even at the early date of this research, both authors recognized the successional processes that were occurring as the lakes slowly fill in. 

In some cases, the research was not necessarily directly related to Saginaw Forest and material taken from the forest was the only aspect of the property used in the study.  For example, Berg (1949, 1950, 1950b) and McGaha (1951, 1952) took plants and the insects associated with them from Third Sister Lake and observed these insectsí¢â‚¬â„¢ life histories, while Hatchett (1947) collected isopods from the forest and both described their life cycles and conducted experiments with them and Laing (1939) took macrophytes from TSL and examined their respiration and gas exchange properties.  Old (1931) described many species of sponges collected from locations around the state including TSL.  Berven and Boltz (2001) took leech and wood frog tadpole specimens to determine the effect of leeches on tadpole fitness traits, and Cornell, Berven and Gamboa (1988) collected larval and young wood frogs to examine kin recognition in frogs.  Velasquez (1940) collected cyanobacteria from lakes and ponds around the state including TSL.  Hopper and Elliot (1953) took mud samples containing protozoa from the lake bottom and found that the organisms released phosphorus from organic compounds.  Schultz (1996) and Schultz and Yurista (1998) took Daphnia pulicaria from TSL for use in predation experiments of Bythotrephes cederstroemi in an attempt to understand the degree that Bythotrephes was responsible for changes in the zooplankton assemblage of the Great Lakes while Branstrator (1995) also studied Bythotrephes using Leptodora kindtii and Daphnia galeotae mendotae collected from TSL.  In some cases, collected material aided in the discovery or first published descriptions of new species such as trematode parasites of frogs and other animals (Bosma, 1931, Krull, 1931).   

Other research has been more intimately connected with the forest property.  Studies have been conducted on terrestrial ecology such as tree growth and forest ecosystems, soil ecology, remote sensing, wildlife ecology as well as aquatic studies on fish ecology, plankton communities, water chemistry and sediment analyses.

Terrestrial Ecology

Tree Growth/Forest Ecosystems
Young (1919, 1928) published the first comprehensive work on the state of research on the plantations of Saginaw Forest.  He detailed the early results of experiments such as the effect of varying cultivation methods, planting density, thinning experiments, proximity to black locusts and soil conditions as well as determining annual growth rates and detailing damage from insects, mammals, diseases and weather.  The thinning experiments continued until the 1950s and Hughes (1957) and Spurr et al. (1957) found that growth rates of large trees in thinned plots increased after the first 15í¢â‚¬”20 years compared to unthinned plots. 

Studies were also conducted on the growth of ponderosa pine (Smithberger, 1957) and red pine (Dreisinger 1954).  Kistler et al. (1979) found that the lowest areas of tree species diversity in Saginaw Forest were in a ponderosa pine plantation (Block III, Lot 7) while the highest areas were in the Picea abies plantation of Block V, Lot 7 and in an oak-hickory forest sampled just outside of SF to the north (an area now owned by the Paul Corporation).  Kistler et al. also found that understory plant communities were dependent on the structure of the forest canopy cover.  Johnson et al. (1986) found that several plantations are overmature and in need of cutting.  They also provide some suggestions for alternative access points to the forest (such as the Aprill property to the North), opportunities for interpretation activities and connectivity of the property with Dolph Park.  

Soil Ecology
Ladrach (1964) did a comprehensive survey of soil type throughout Saginaw Forest.  In the most recent research conducted in Saginaw Forest, Kahan (2008) studied how residential housing densities affected the terrestrial biogeochemical cycling of nitrogen in a number of protected areas.  He found that higher housing densities were positively correlated with soil N content, although this did not appear to be a proxy for plant N content, biomass or species diversity.  His research found that Saginaw Forest had very nitrogen rich soils that were more likely to be carbon limited than nitrogen limited. 

Remote Sensing
Webber (1966) tried to use grey scale densities and multi-band spectral imaging to differentiate between eight species of trees in Saginaw Forest, however he was unable to consistently and accurately tell the difference between them.  Olson (1985) also used a variety of remote sensing techniques in research on Saginaw Forest.

Wildlife Ecology
Much of the wildlife studies in the forest have investigated the frog populations, however research has also been conducted on birds and mammals.  Yeager (1937) studied the behavior of red squirrels piling pine cones in three Norway spruce plantations of Saginaw Forest.  He determined that locations of their caches were a result of food availability rather than other aspects of habitat type.  Yambert (1951) developed a wildlife program for Saginaw Forest.  Howard and Kluge (1985) studied wood frogs in SF and found that non-random mating occurred in wood frog populations and that there were size selective advantages for both male and female frogs.  TSL was one of two ponds in Michigan that Rihi and Berven (1991) used to examine size differences of Michigan wood frog tadpole populations compared to populations in the southeastern U.S.  They found that Michigan wood frog larvae grew faster, were larger and were more susceptible to density dependent pressures than their counterparts in Maryland and Virginia although these differences did not persist in laboratory specimens.  Johnson (1969) gives a historical analyses of the effects of insects on Saginaw Forest.  Hann (1939) concluded that castration of Slate-colored Juncos, Red-eyed Towhees, and White throated Sparrows at a banding station in SF did not affect their migratory patterns. 

Aquatic Ecology

Insect and Fish Communities
Ball, Brown and Hayne conducted a series of experiments on the fish population of the lake culminating in the removal of all fish from TSL after treatment with rotenone in 1941.  Their findings indicated that Libelluline dragonfly larvae were more available to fish than Gomphinae nymphs (Ball, 1943).  Ball (1947) observed that a large proportion of bluegills and yellow bullheads tended to remain in one location (within 30m) for long periods of time while largemouth bass roved throughout the lake.  Ball (1948) found the invertebrate community to be dominated by caddisfly, midge and dragonfly larvae as well as leeches.  Ball and Hayne (1952) found that the rotenone treatment did not have an adverse affect on most of the insect populations, although leeches, Chaoborus and some dragonflies were adversely impacted.  Brown and Ball (1942b) detail the effects of rotenone treatment on fish and other creatures in TSL and specifically examined the relationship between depth, temperature and toxicity of treatment.  14 fish species (plus a green sunfish hybrid) were recovered from the poison treatment (Brown and Ball, 1942), of which the majority of the biomass was bluegill while more than 50% of the fish were either blacknose or black chinned shiners.  TSL supported 86.7 fish/acre.  A year after poisoning, long-ear sunfish (previously not observed) colonized TSL.  Since these experiments, no research has been conducted on the fish population of TSL. 

Eggleton (1931) found 27 invertebrate species within 18 genera in the deeper waters of TSL and examined the effect of anoxic and hypoxic conditions on several aquatic invertebrate larvae.  In a separate study (Eggleton, 1931b), he examined the distribution, migration and abundance of Chaoborus larvae.  McNaught (1993) examined the relationship between Chaoborus and invading Leptodora kindtii larvae and accidently introduced the latter into TSL.  Bridgeman et al. (2000) found that decreases in diversity and abundance of the invertebrate community had occurred between Eggletoní¢â‚¬â„¢s study and their sampling in the late 1990s.  They also found that the more profundal invertebrates had shifted their spatial distribution to shallower waters and suggest that these changes are due to decreasing vertical mixing caused by the use of road salt. 

Macrophyte and Plankton Communities

Reed (1902) determined that Potamogeton zosteraefolius  was the most common macrophyte in TSL, however Ball (1948) found 28 macrophyte species of which Potamogeton amplifolius was the most prevalent. 

Cotner and Wetzel (1992) found that algae have the potential to outcompete bacteria phytoplankton if phosphorus inputs increase and the lake is not carbon limited.  Lehman and Sandberg (1982) found that phytoplankton were limited by phosphorus and not by nitrogen, however Lehman and Naumoski (1986) found that phosphorus and nitrogen inputs to TSL yielded an order of magnitude and a 25% increase in chl a production respectively.  In April-May, the dominant phytoplankton are chrysophytes and cryptophytes, but by late May, chlorophytes and cyanobacteria come to dominate the community (Kiesling, 1990). Bridgeman et al. (2000) found that the winter/spring phytoplankton community was dominated by Oscillatoria prolifica and O. limnothrix although diatoms and dinoflagellates were also found. 

Dorazio (1986) found that a cyclopoid copepod, Mesocyclops edax predated on three of nine rotifer species as well as copepod nauplii in TSL.   Hammer (1995) used sediment cores and plankton samples to determine that some diatom species that had previously been present in TSL no longer occur here.  Diatoms dominated the winter phytoplankton community, while the zooplankton community was dominated by cyclopoid copepods with small cladocerans (Bosmina) also being important (Judd et al., 2005). 

Korstad (1980) found that grazing zooplankton can be beneficial to zooplankton due to their high rates of nutrient release while Lehman and Naumoski (1985) determined that Daphnia can vary their turnover rates of phosphorus depending on the nutrient status of their food.  Lee (1982, 1984) looked at genotypic plasticity in Daphnia populations of TSL and a small pond at the UM Botanical Gardens and found that clones in traditionally unpredictable areas had more genetic plasticity than those that exist in more stable environments.  He also noted that Daphnia populations above the thermocline in TSL were smaller and had longer tail spines than those populations that persisted below the thermocline.  Kiesling (1990) studied the seasonal succession of the zooplankton community and also found that minnows and larval bluegill were more responsible for the structure of the zooplankton community than invertebrate predators.  McNaught et al. (2004) found changes in the zooplankton community following colonization by Leptodora kindtii including increases in species diversity (suspected to be temporary), decreases in Bosmina and Ceriodaphnia populations, increases in the proportion of largebodied Daphnia and increases in Diaptomus populations.

Lake Chemistry
Experiments with nutrient additions and lake enclosures were conducted by Bender and Jordan (1970), Jordan (1970), and Jordan and Bender (1973).  Although they found that enclosures did not accurately mimic actual lake conditions, they found that algal response varied depending on season, concentration, species, combination and form of nutrient addition.  Cotner and Wetzel (1991, 1991b) investigated the ability of bacteria to hydrolyze phosphorus and found that bacteria in lower productive areas were more likely to contain enzymes to release phosphorus nutrients and that in TSL, most of the phosphorus regeneration in bacteria was through alkaline phosphatase metabolic pathways.

The history of dioxane contamination of TSL and its East Inlet was reviewed by Englebert et al (1988).  Bridgeman et al. (2000) found that chloride and conductivity have continued to increase and contribute to increased stability of the water column which results in longer periods of anoxic conditions and summer stagnation.  They attribute these chemical changes to the increasing use of road salts as urbanization has increased in the watershed.  Nutrient supply to the epilimnion is dominated by allocthonous processes (mostly from the East Inlet) rather than internal mixing (Judd et al., 2005) and lake stability has continued to increase since Bridgemaní¢â‚¬â„¢s study. 

Sediment Analyses
Dunette (1973, 1989) and Dunnette et al. (1985, 1990) conducted a series of experiments examining sulfate reduction in TSL.  Their work found that TSL had relatively low rates of sulfate reduction and putrefaction/cysteine decomposition and that sulfate reduction was responsible for 4-5x total hydrogen sulfide production compared to putrefaction.

Kelly and Chynoweth (1980, 1981) and Robertson (1978, 1978b) and Robertson and Chynoweth (1978) studied methane production in TSL.  They found that laboratory analyses accurately predicted field measurements for surface sediments, but not for deeper sediments. They also discovered that organic matter deposition and methanogenesis are linearly related and that sedimentation is a larger driver for methane release than temperature in freshwater lakes. 

Eggleton (1931) was the first to describe the alternating bands of dark organic matter and lighter gray clay layers that make up the sediment of TSL.  Potzger and Wilson (1941) investigated pollen grains in the sediment of TSL to determine forest succession during the post-pleistocene period.  Hammer (1995) and Hammer and Stoemer (1997) found an annual rate of sediment deposition of 2.5 cm/year and described this pattern as a result of storm events bringing in allocthonous clay material while the lakeí¢â‚¬â„¢s organic matter decomposes between events.  They also found a difference in the diatom assemblages of the two different types of strata.  Sediment analyses by Judd et al. (2005) found that Asterionella diatoms have increased since Hammer and Stoemerí¢â‚¬â„¢s study.