Journal of Archaeological Science (1994) 21, 667-673 Bone Weathering in the Tropical Rain Forest Martha Tappen Department of Anthropology, Harvard University, Cambridge, MA 02138, US.A (Received 24 May 1993, revised manuscript accepted 30 October 1993) Bones deposited on the rain forest floor exhibit distinctly different surface modifications from those found in savannas. ‘The development of weathering cracks is considerably slower than in more open environments, suggesting that the dearth of rain forest fossil assemblages is not caused by increased weathering in these environments. Ubiquitous surface ‘modifications include green plant growth on the bone’s exposed surfaces, termite-caused pitting, and extensive rodent ‘nawing. If preserved on fossil bone these modifications may serve as palacoenvironmental indicators, Keywords: TAPHONOMY, RAIN FOREST PALAEOECOLOGY Introduction etween the time of an animal's death and the burial of its bones (leading (o their preservation as fossils) many processes can directly modify Skeletal remains. These modifications form an impor- tant part of the palaeontological and archaeological record and provide clues for reconstructing tapho- nomic history. Bone weathering is @ “perthota process (Clark & Kietzke, 1967) brought on by en- vironmental conditions, and as such varies between environments. Actualistic studies have demonstrated that microenvironmental conditions, such as soil pH and amount of shade, strongly influence the rate of bone weathering (Behrensmeyer, 1978). To apply actu- alistic observations of bone weathering rates to fossil siles requires that either the rate and type of weather- ing is uniform in all environments, or that the palaeo- environment from which the fossil sample was derived was roughly equivalent to that from which the actual~ istic observations were made. In this paper | compare bones from the Central African rain forest with those from @ Central African savanna and East African savanna (the latter based on published descriptions). ‘The bone modifications found in the rain forest differ from those found in savannas, and thus if observed on fossil bone could assist in reconstructing the local palaeoenvironment from which they were originally deposited. Understanding weathering patterns of modern-day bone can help to gamer significant palaeobiological and palacoenvironmental information. For example, weathering cracks have been shown to follow bone structure (Collagen fibres), providing information about biomechanics of living and extinct species (N. C. Tappen, 1969, 1976; N.C. Tappen & Peske, 1970) They also act as an indicator of the length of time bones were exposed on the surface prior to burial (030-440394050667 +07 $08,000 Behrensmeyer (1978) defined six progressive stages of subaerial bone weathering, from Stage 0 (no cracks) to Stage 5 (falling apart), as part of her study of attri tional bone deposition in Amboscli Park, Kenya. These time-successive stages of increasing weathering suggest that the degree of weathering is directly related to amount of time of subaerial exposure. Taphono- ‘mists and zooarchaeologists have widely adopted this weathering stage scheme in their analyses of faunal assemblages. Attempts have also been made to esti- mate the duration over which a fossil bone assemblage accumulated, but not without problems (Potts, 1986; Bunn & Kroll, 1986; Lyman & Fox, 1989). This paper adds another dimension to the study of bone weathering and surface damage. I report on bone surface modification in Parc National des Virunga, Secteur Nord (hereafter PNV), an equatorial savanna, and in the Ituri Rain Forest, both in Zaire (Figure 1). Bone weathering in the savanna environment at PNV is similar to that documented by Bebrensmeyer in Amboseli Park. However, bone modification in the rain forest follows an entirely different pattern. Bone weathering in the classic sense of progressive eracking is delayed and sometimes absent. Other surface modifications occur, and are potentially preservable on fossil bone, Pare National des nga (savanna) ‘The study area in PNY is a medium and tall grass savanna with scattered Acacia and Euphorbia trees, located on the floor of the Western Rift at the north end of Lake Rutanzige (ex-Lake Amin, ex-Lake Edward). It receives more than twice as much annual rainfall as Amboseli Park, and is notably less seasonal and without frequent droughts (930mm of rain per annum (Verschuren, 1986); compared with 350:mm at (© 1994 Academic Press Limited(668 M. Tappen Central Africa Sutvey X ee as ? Me Figure 1. Location of the rain forest and savanna study areas in Central AV, Amboseli (Coe, Cumming & Phillipson, 1976)). Soils are formed from the volcanic ash of the Katwe ‘Volcano (Uganda) and for the most part have a neutral pu. AS part of an investigation of natural bone deposi- tion rates in the park, weathering patterns were noted for about 6000 bones found across the park in 1986 and 1988 (M. Tappen, 1992, 1994 in press). Weathering in PNV shows the same basic patterns as at Amboseli Park; it begins with linear cracking and progresses with ‘more numerous deeper cracks and forming a fibrous texture (Behrensmeyer, 1978). It is not surprising that weathering patterns are similar, as weathering cracks have to do with the structure of the bone itself (N.C, Tappen, 1969) and actualistic observations on weathering cracks have been noted in a variety of habitats, including temperate ones [e.g. Wisconsin (Tappen & Peske, 1976), England (Andrews & Cook, 1985), Nebraska (Fiorillo, 1989)] However, the savanna study is not yet of sufficient duration to ascertain that weathering raves are pre- cisely the same as at Amboseli. The longest observa- tions were over four years of a buffalo skeleton found recently dead, still ereasy, with tendons and without cracks in the summer of 1986, The weathering of this skeleton was recorded two years later (in 1988, Figure 2) and photographed four years later (1990, Figures 3 & 4). AL two years the bones were in weathering stage 1 with a few approaching stage 2, while the bones exposed for four years were in weathering stage 2. The rate of progression of weathering is within the range of Amboseli, though at the slower end of the distribution (Behrensmeyer, 1978: 157, table 1). The sample size is at this time too small to assess the rate of weathering im this environment, but the characteristics of the Weathering observed on the surface assemblage are similar to those recorded by Behrensmeyer (1978). While cracking may sometimes be less deep, further study is required to verify this. Figure 2 Weathering of a bullslo skelton after (wo years of exporure in PNV.Figure 3, Weathering ofthe same baffle skeleton in PNY after 4 yedtsof exposure. (Photo by Leith Smith). Figure 4. Weathering ofthe same buffalo skeleton in PNV after $ years of exposure. Ths bone was also broken by teampling. (Photo by Leith Smit, Tturi Rain Forest Bone weathering in the Tturi Forest was studied in ‘conjunction with Laden’s study of Efe land-use ecology (Laden, 1992) in 1986 near the Harvard Ituri Project’s Ngodingodi research station and again in 1990 in the forest 35 km north of Ngodingodi, near the village of Nepoko. Annual rainfall averages about 1900 mm (Wilkie, 1987), with notable wet and dry seasons. Natural bone deposition is generally light in the forest and bones are dificult to find because of poor visibility Table 1 Elephont sites examined in the lar Forest Bone Weathering in the Tropical Rain Forest 669 caused by dense vegetation and leaf litter, and because bbones are not bleached white by the sun. Weathering was studied for eight elephant skeletons (total number of bones=118) (Table 1). All the elephants were killed and eaten by inhabitants of the Ituri, and were located with the assistance of Efe guides ‘The date of death is known for only some of the elephants, based on Efe and Lese informants who had dlirect knowledge of the elephant’s death. None of the skeletons observed were greasy or had any flesh or ligaments left, nor were there any remains left of the sapling barazas (small shelters) or smoking racks that were built when the animals were processed. Given these observations these skeletons are believed to have been deposited at least several years prior to observa- tion. Soil pH was taken with a LaMotte Soil Test Kit at Sites 1 (pH 40) and Site IV (pH 425), indicating their generally acidic nature. At most of the sites only a portion of the original skeleton remained. Several factors likely contributed to the loss of bones from the sites: (1) Sites T and IV were periodically submerged and were in very soft mud and itis likely some elements were buried; (2) some bones may have been missed despite intensive survey of the area; (3) some bones may have been entirely consumed by rodents (see below); (4) some bones may have been transported away from the site by the people who processed the meat; (5) some bones may have been burnt in fires, since the Efe often throw bones in fires when they are finished eating; (6) some bones may have been so severely chopped up that they were essentially destroyed. These last three processes have been observed at elephant butcheries (Fisher, 1992). Surface damage had four main causes, producing distinct types of damage: 1. Green plant growth on the upper surface of the bone (788% of the bones) (Figure 5). It is not clear that moss or algae leave recognizable microscopic traces on fossil bone. Often fungi are also found on the upper and lower surfaces, sometimes etching a net-like (s0- called “root”) pattern into the bone. In contrast, in the savanna at PNY, moss or fungi were observed only in very rare instances (only 14 bones in 9 different ppaiches), and all were found in localized dense patches of vegetation. ‘Number of Number ef "Number of bones Site ets ‘bones | Weathering stage with racks Sewing 1 unknown 9 2 u primary forest—Marantaceae grove 0 tinknovwn 8 5 0 Feerine forest tt 37 2 ° 0 primary forest Vv vnknown a ° 0 feed plain Sverne forest y 125 2 1 B ranite dome gap (haba) v 16 4 ° ° ‘ld secondary forest vir 10 3 1 1 ‘ld secondary forest vin ois 6 3 fold sacondary forest under Kongo nut tee670M. rappen Figure S. Elephant Site VII humeras. The ubiquitous green plant srowih on the upper sie of the banes inthe It is shewa bere by the dark colour. Lighter afeas are where rodent. gnawing bas removed the plant growth Figure 6, The underside of mandible from Flephant Site VII, Showing extensive pitting de to termites, 2. Termite activity which creates pits (Figure 6). While also occurring in savanna environments, the frequency with which termite modification occurs in the rain forest is remarkably high (59% of the bones). The termites live almost exclusively on the lower, un- exposed surface of the bone, building up walls of soil around their colony. Inside’these homes, they create small round pits that, while usually shallow, can be quite deep, incising through several millimetres of cortical bone, and sometimes occurring in linear trails (Figures 7 & 8). By contrast, at PNV, termite or other insect pits were found on only 14 bones (again less than 1% of the bones), even though termites are abundant in the park (although they may be different species). However, Behrensmeyer (1978) noted termite pits were “not uneommon”’ at Amboseli Park, and they are also common on bones near Lake Baringo (J. Kingston, pers. comm.) 3. Extensive rodent gnawing. This was found on 559% of the Ituri elephant bones (Figures 5 & 9). Many of, the bones were in the process of being completely Figure 7, Termite piting in linear tail, from Elephant Site VII ‘Cut maris are vsile inthe upper left comer. Figure & Scanning electron micrograph of termite pitting. The smooth dark area near the edges ofthe photo represents the original bone surface, Figure 9. Covtical bone probably removed by some sort of chemi ching on a hone fragment from Elephant Site VIIL. Al edges ofthis bbone have been gnawed by rodent.Figure 10. An elephant hone ina state of advanced weathering from Pate National det Vieungs consumed by rodents, Efe informants and matching tooth size concur that the most common species con- suming the elephant bones is the giant forest squirrel, Protoxerus stangeri. Rodent gnawing is also common in savanna environments (e.g. Braia, 1981); however, aside from within porcupine lairs, to’ my knowledge it does not approach the frequencies seen in the Tturi, In sharp contrast, in PNV only $4, or about 1%, of the bones studied exhibited rodent gnawing, 4, Weathering cracks rare (15% of the bones). Only two skeletons had bones beyond stage 1 weathering, (Elephant Site 1 had one bone with stage 2 weathering and Elephant Site VIII had bones in stages 1-3) In contrast, in the savanna, bones from the older skeletons exposed for 10 years or more would usually be in advanced weathering stages. Weathering cracks are extremely delayed in the rain forest. This delay is beyond what would be expected just from the effect of the large size of elephant bones. Elephant Sites II, IT (greater than 7 years old), IV and VE (about 16 years ld) had no bones with weathering cracks, site VII had fone bone with weathering cracks, site VIII had three, Meanwhile, the elephant bones in PN savanna exhibited classic weathering cracks as described by Behrensmeyer (Figure 10) ‘Additionally, when weathering cracks do occur on forest bones, they are just as likely to begin on the underside as the upper side of the bone. In contrast, in savannas, weathering cracks usually oceur first on the upper surface [except when found on very alkaline soils where salt crystals form on the underside of the bone and cause eracking (Behrensmeyer, 1978)]. In the rain forest, where soils are often acidic or neutral, the cracks that are observed on the underside of the bones often appear to be more like grooves than cracks, possibly due to chemical etching. ‘One elephant skeleton (I year 3 months), from Site V, was exposed in an upland gap on a granitic dome. ‘This skeleton was exposed to much more sunlight than the others in the forest, and weathering cracks (stage 1) similar to those found in the savanna were present on Bone Weathering in the Tropical Rain Forest 671 13 (41%) of the bones. Weathering cracks at the other forest sites combined were present on only 6% of the bones. Thus, the youngest skeleton of known age was weathering more like it would in savanna conditions (although moss, rodent gnawing and termite pits were also abundant). Another feature occasionally observed on elephant bones in the rain forest is etching into the outer layer of bone, especially on the lower surface. Large areas of ‘outer cortical bone can be removed in some cases (Figure 9). This appears to be chemical in nature, and may involve the relatively acidic soils (¢.g. pH 4'5) of the forest. Another possibilty is that it could be caused by organisms which consume the green plant material growing on the surfaces of bone (Behrensmeyer, pers. comm,), This removal of the outer layer of bone is relatively uncommon. Very few naturally occurring bones were found in the Ituri, despite intensive search efforts (M. Tappen, 1992). Based on observations of culturally deposited bones (most from middens around Efe camps), it is clear that there are some differences between modifica- tion of elephant bones and those of smaller animals (such as of monkeys and duikers). Like the large bones, small bones had green plant growth on their upper surfaces and delayed or nearly halted cracking caused bby weathering. Termite pits and rodent gnawing were not found on the small bones, and it appears that these bone consumers select mostly (only?) large bones. Animals, including rodents, avoid even unoccupied Efe ‘camps, for fear of being hunted (Laden, 1992). This ‘may be one reason there is litle rodent gnawing on the sample I studied. Bones of intermediate sized animals, such as forest bulfalo or okapi, were not found outside of hearths and so itis not known if they are frequently inhabited by termites or gnawed on by rodents, Discussion 1t appears that weathering erack formation is slower in the rain forest environment than in open savannas, Behrensmeyer noted that microenvironmental con ditions and especially whether or not @ bone is in the shade are critical factors affecting weathering rates. She argued that exposure to sunlightshade, soil conditions and wetting-drying have a large effect on the weather- ing rate of an individual bone, and that extremes in temperature and moisture increase weathering rates, ‘The slower rates of weathering she found in the swamp environment at Amboseli and the slower rates in the rain forest suggest that drying of bones increases weathering cracks. Table 2 presents a qualitative summary of the typical conditions in the immediate environment of a bone lying on the ground’s surface in the Tturi, PNV, and Amboseli. In the forest, shelter from sunlight is the main cause of all of these prevalent conditions except the low soil pH. Sunlight is blocked by the overlying:672 M. Tappen Table 2. Characteristics of the oumediar environment ofa typical bone lying on the suface i the Iu Forest, Pare National des Ving wetter sevanna and Ainboselt dryer savarma ‘Variation in humidity and eur almost none igh UY tight Humidity NV high imermedtite Amboseli very high ‘en ey eniperature Soit pit acide nevtrat sometiocs high alkaline forest canopy, creating a situation with little diurnal variation in temperature or humidity. The lack of warming sunlight along with dense vegetation and its ubiquitous transpiration insures that evaporation rates are low, that it is always humid, and bones never dry ‘out. The air temperature remains cool and relatively constant in contrast with out in the open, Even just metres away in a cleared garden, temperature can be dramatically higher. The temperature on the surface of bones therefore always remains cool, contrasting with bones at Amboseli which are hotter midday than in the morning or evening (Behrensmeyer, 1978: 154). The ultraviolet component of sunlight is significantly re- duced in shady environments, so any deteriorating effects of uitraviolet radiation on bone structure are reduced. In a temperate environment (England), bones out side in a sheltered spot have been observed to exhibit no weathering cracks after seven and a half years (Andrews & Cook, 1985). This, along with the slow weathering rates in the Tturi, suggests that ultraviolet light is important in bone weathering (an idea long held by Behrensmeyer, pers. comm.). In the rain forest, bones receive very little direct light, and so do not become bleached white, dry out, or crack. Extreme variations in temperature do not occur, and while there is variation in moisture (two of the elephant sites I have seen periodically flood), they never become “bone dry”. Controlled laboratory experiments are needed to tease apart the relative importance of exposure to ultraviolet light, temperature variation and humidity in weathering, If ultraviolet light, extreme drying and/or wetting and drying are the main catalysis for bone weathering, continued study may show that weathering rates are slightly slower in the relatively moist savanna at PNY than the extremely dry arca at Amboseli. It would be interesting to study weathering rates in a true desert environment, where weathering may be much more rapid, None of the modifications seen on rain forest bone are unique to that environment, but the high frequen- cies in which they occur together may be. If these features (termite pitting on the lower side, cracking, grooving only on the lower side, and extensive rodent gnawing) are found together at high frequencies in a fossil assemblage, it may supplement other palaeo- environmental indicators from the site, However, the attractiveness of bones to consumers needing calcium or other nutrients may vary between different rain forests ‘A common assumption is that bones are rapidly destroyed by acidic soils in the rain forest. While this may ultimately be their fate, rapid subacrial destruc- tion by chemical processes ‘was not evident in my investigations. Large elephant bones 10-15 years old were still present and often without weathering cracks on the surface, Progressive cracking due to subaerial weathering is actually slower than in open environ- ments, The sparseness of rain forest representation in the fossil record may be instead due to other biological and taphonomic processes. These include the smailer body size of animals, smaller group size, possibly less frequent catastrophic mass deaths (e.g. by drought), and the consumption of bones by rodents, insects, plants and fungi. Such factors may be much more important for the sparseness of the rain forest palac- ontological record than bone weathering. Additionally, sedimentation regimes may play a role; for example, in the Ituri thee is litte sedimentation, and the erosional nature of the topography does not promote burial. Acknowledgements I would like to thank the Government of Zaire for permission to work there, and LZ.C.N. park guards and Zairois citizens who assisted me in my bone surveys. I thank J. W. K. Harris for inviting me to join the Semliki Research Expedition, and Leith Smith for taking photos of a weathering skeleton for me in the summer of 1990. 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