
25th Annual Meeting and Symposium of the
Desert Tortoise Council, April 21-24, 2000 Abstracts

Trace Element Systematics of Desert Tortoise Shell
Gordon B. Haxel1,3, Roy J. Knight1,4,
Kristin H. Berry1,5, Bruce L. Homer2, and Maurice
A. Chaffee1,4
1U.S. Geological Survey, 2University
of Florida, Gainesville, Florida; 3Flagstaff, Arizona; 4Denver,
Colorado; 5Riverside, California

Among the suspected causes of accelerated morbidity and mortality of the
desert tortoise (Gopherus agassizii), a threatened species, is elemental
toxicity related to several heavy metals and metalloids. We and several
colleagues are evaluating this hypothesis by investigating the biogeochemical
pathways that link rock, soil, vegetation, and tortoises; comparing trace
element concentrations in the shells and internal organs of healthy and diseased
tortoises; and studying the chemical effects of geological environment and
various human activities and products on tortoise health.
Here we report data that establish for the first time the basic trace element
patterns that characterize the two major components of desert tortoise shell -
bone and scute. Shell materials from six apparently healthy tortoises
(accidentally traumatically injured or killed by vehicles) and from 14 diseased
tortoises were analyzed by instrumental neutron activation analysis (INAA).
Powdered samples were irradiated by thermal neutrons in the USGS research
reactor in Denver. Five gamma ray counts were performed: < 1 hour; and 5, 7,
14, and 60 days after irradiation. Approximately 18 elements were determined
with satisfactory accuracy and precision in tortoise bone, scute, or both; and
six additional elements could be determined at moderate to high concentrations.
As the measured abundances span more than seven orders of magnitude, from major
(structural) elements such as Ca in bone (about 23 percent) to trace elements such as
Sc or Sb (10 - 40 ng/g (ppb)), presentation of the data is facilitated by
normalization to the average composition of upper continental crust (UCC) (Fig.
1).
Despite the contrast in bulk composition between bone (apatite-like) and
scute (composed of keratin, an insoluble protein), the normalized trace element
patterns of bone and scute from healthy tortoises are rather similar, differing
by no more than one-half an order of magnitude for most elements (Fig. 1). The
higher Ca and Na content of bone is to be expected. These basic trace element
patterns evidently are determined primarily by biologic processes rather than by
geologic factors, as the analyzed tortoises lived on diverse geologic substrates
in widely separated areas of the Mojave and Colorado Deserts, southern
California. Local geologic factors may influence details of the patterns.
The trace element spectra (Fig. 1) comprise three segments: alkali and alkali
earth elements on the left side, transition metals (Sc-Ni) in the middle, and
chalcophile ("sulfur-loving") metals and metalloids (Zn-Se) on the
right. The transition metals plotted either are trace nutrients (Cr, Fe, Co, Ni)
or are of little or no biological significance at parts-per-billion levels (Sc,
La - Lu, Th, Ta). Zinc, Se, and probably As are essential trace nutrients,
although all are toxic at high concentrations; Sb and Au have no known natural
biological role. In both bone and scute, the transition metals, except Cr and
Ni, are effectively "diluted" to ~10-2 times their upper-crustal
abundances. In contrast, the chalcophile elements Zn, As, Sb, and Se, along with
Au, have markedly higher normalized abundances. This relative enrichment of the
chalcophile elements in both bone and scute suggests that sulfur may be
important in controlling these trace element patterns.
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| Figure 1. The normalized trace element
patterns of bone and scute from healthy tortoises. The trace element spectra is
comprised of three segments: alkali and alkali earth elements on the left side
(Na-Ca), transition metals (Sc-Ni) in the middle, and chalcophile
("sulfur-loving") metals and metalloids (Zn-Se) on the right. |
Desert tortoises spend six to nine months of the year underground in their
burrows, in close contact with soil. Scute (the outer layer of the shell) is not
appreciably enriched relative to bone in lithophile trace elements such as Le,
Ce, Th, or Ta nor in potentially labile elements such as As and Se, suggesting
minimal elemental input to the shell through contact with soil, at least in
healthy tortoises.
Our trace element data also reveal several interesting and possibly important
differences between healthy and diseased tortoises. Scute from diseased
tortoises is strongly enriched, relative to scute from healthy tortoises, in
several trace metals, most notably Au and W (!). The significance of this
anomaly is unclear. Three diseased tortoises have much higher scute As and
slightly lower scute Se than the other healthy and diseased tortoises. In
tortoise bone, As and Sb are particularly informative. These two trace elements
evidently are incorporated into tortoise bone in approximately the same ratio as
they occur in UCC: As/Sb about 7.5. Two diseased tortoises have highly elevated
bone As and Sb, with somewhat low As/Sb ratio. These pronounced anomalies in the
chalcophile trace elements As, Se, and Sb in several diseased tortoises suggest
some perturbation in the concentration, siting, chemical state, or utilization
of sulfur within the organism. Whether such a disturbance is more likely a cause
of disease or a result of disease remains to be determined. We shall continue to
evaluate these ideas as we acquire data for more tortoises and for additional
potentially toxic chalcophile elements such as Cd, Hg, and Pb.
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