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THE

TEXAS JOURNAL

OF

SCIENCE

GENERAL INFORMATION

MEMBERSHIP. — Any person or members of any group engaged in scientific work or interested in the promotion of science are eligible for membership in The Texas Academy of Science. Dues for members are $20.00 annually; student members, $12.00 annually; sustaining members, at least $30.00 in addition to annual dues; life members, at least $400.00 in one payment; patrons, at least $500.00 in one payment; corporate members, $250.00 annually; corporate life members, $2000.00 in one payment. Library subscription rate is $45.00 annually. Payments should be sent to the Dr. Michael J. Carlo, P.O. Box 10986, Angelo State University, San Angelo, Texas 76909.

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The Texas Journal of Science is published quarterly at Lubbock, Texas U.S.A. Second class postage paid at Post Office, Lubbock, Texas 79401. Please send form 3579 and returned copies to PrinTech, Box 4240, Lubbock, Texas 79409-3151.

ISSN 0040-4403

THE TEXAS JOURNAL OF SCIENCE

Volume 42, No. 1 February 1990

CONTENTS

Feasibility of penaeid shrimp culture in inland saline groundwater-fed ponds.

By L. L. Smith and A. L. Lawrence . 3

Bats from the coastal region of southern Texas.

By S. S. Chapman and B. R. Chapman . 13

A nutritional analysis of diet as revealed in prehistoric human coprolites.

By K. D. Sobolik . 23

Abnormal terminal Cretaceous Foraminifera of east-central Texas.

By H. Montgomery . 37

Syntheses of 2-(2-pyridyl)cyclohexanone and related cyclohexanones.

By E. H. Sund, P. D. Koplyay, and S. D. Wood . 45

An efficient retractable mobile antenna tower for radio-telemetry studies.

By M. T. Pollock, S. Demarais, and R. E. Zaiglin . 49

Origin of color of American Indian black and red ceramics.

By M. Hyman and M. W. Rowe . 55

Observations on obtaining white-tailed deer fawns for experimental purposes.

By I. M. Ortega, L. D. Perry, D. L. Drawe, and F C. Bryant . 69

A note on some aspects of Pitman nearness.

By J. W. Seaman, Jr., and D. M. Young . 73

Phyllodont (Paralbulinae) fish toothplates from the Lower Cretaceous

Glen Rose Formation of central Texas. By H. H. Huggins . 77

Importance of canopy position for growth of Celtis laevigata seedlings.

By O. W. Van Auken and R. J. Lohstroh . 83

Solar pond feasibility for low cost energy and water production.

By M. A. K. Lodhi . . . 91

General Notes

Distributional records of the yellow-nosed cotton rat, Sigmodon ochrognathus

Bailey, in Texas. By R. R. Hollander, B. N. Hicks, and J. F Scudday . 101

Manatee stranding on the coast of Texas. By S. Fernandez and S. C. Jones . 103

Occurrence of house dust mites in central Texas. By S. L. Outhouse and

S. D. Castro . 104

A bighorn sheep, Ovis canadensis , from the late Pleistocene of Mesa Del Oro,

Cibola County, New Mexico. By R. A. Smartt, D. J. Hafner, and S. G. Lucas . 107

Instructions to authors

111

THE TEXAS JOURNAL OF SCIENCE EDITORIAL STAFF

Editor:

J. Knox Jones, Jr., Texas Tech University Assistant to the Editor:

Marijane R. Davis, Texas Tech University Associate Editor for Botany:

Chester M. Rowell, Marfa, Texas Associate Editor for Chemistry:

Marvin W. Rowe, Texas A&M University Associate Editor for Mathematics and Statistics:

Patrick L. Odell, Baylor University Associate Editor for Physics:

Charles W. Myles, Texas Tech University Editorial Assistants:

Larry L. Choate, Texas Tech University

Jim R. Goetze, Texas Tech University

Richard W. Manning, Texas Tech University

Scholarly papers in any field of science, technology, or science education will be considered for publication in The Texas Journal of Science. Instructions to authors are published one or more times each year in the Journal on a space-available basis, and also are available from the Editor (The Museum, Box 4499, Texas Tech University, Lubbock, Texas 79409, 806/742-2487, Tex-an 862-2487).

The Texas Journal of Science is published quarterly in February, May, August, and November for $20 per year (regular membership) by The Texas Academy of Science, Drawer H, College Station, Texas 77844. Second-class postage rates (USPS 003804) paid at Lubbock, Texas. Postmaster: Send address changes and returned copies to The Texas Journal of Science, P.O. Box 4240, Texas Tech University, Lubbock, Texas 79409-3151, U.S.A.

FEASIBILITY OF PENAEID SHRIMP CULTURE IN INLAND SALINE GROUNDWATER-FED PONDS

Linda L. Smith and Addison L. Lawrence Texas A&M University Shrimp Mariculture Project,

Texas Agricultural Experiment Station, P O. Drawer Q, Port Aransas, Texas 78373

Abstract. — Saline groundwater in Willacy County of southern Texas was used to polyculture juvenile (1.2 + 0.18 grams) Penaeus vannamei and five-day-old postlarval P. setiferus in earthen ponds. Initial stocking densities of 50,700 juvenile P. vannamei and 247,000 P. setiferus per hectare produced 871.7 + 14.28 kilograms per hectare for a 120-day grow-out period. Survival of P. setiferus was less than one percent. Final size and survival of P. vannamei were 19.9 ± 0.71 grams and 86.7 + 4.53 percent, respectively. Despite the poor performance of P setiferus, P. vannamei production data compared favorably with that of a bay-fed system stocked with P. vannamei from the same population, indicating that this particular source of saline groundwater can be used for culture of at least one penaeid species. Use of chemically similar bodies of saline groundwater for mariculture over a more widespread area of Texas is encouraged. Key words: Penaeus sp.; saline groundwater; aquifer; mariculture.

World-wide, salt adversely affects more than 30 percent of the rangeland and 27 percent of the cropland by reducing the potential of these areas to produce traditional animal and plant agricultural crops (Rains, 1979; Kelley, 1982). About two-thirds of the continental United States are underlain by saline water (Feth et al., 1965; Feth, 1970). Excluding those coastal areas directly inundated by ocean or bay waters, the major salt-affected areas in the U. S. are associated with the Great Salt Lake basin, interior valleys of California, and with the Colorado and Rio Grande drainage basins (Chapman, 1960). Vast areas of Texas are underlain by saline groundwater in the form of subsurface and near surface aquifers and outcrops. The geology of 29 principal saline aquifers (that is, those containing 3000 parts per million or more of total dissolved solids and that are capable of producing a minimum of 378 liters per minute) have been described in Texas (Winslow and Kister, 1956; Texas Water Development Board, 1972). Mariculture, rather than traditional agriculture, has been suggested as a means of improving productivity of these salt-affected lands (Stickney and Davis, 1981; Issar et al., 1983; Payne, 1983; Sarig, 1984).

The most familiar mariculture-related use of inland saline waters has been the harvesting of Artemia. Within a more strict mariculture context, microalgae have been cultured for both pharmacological and aquacultural purposes in Israel (Goldman, 1979; Pasternak, 1982) and the U. S. (Walsh et al., 1985; Goldstein, 1986a). The culture of higher species in inland saline groundwater is limited mainly to studies on tilapia (Granoth and Porath, 1983; Hamza and Zaki, 1987) and gilthead sea

4

THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

bream (Pasternak, 1982) in Israel and Egypt, and, to a lesser extent, oyster (Goldstein, 1986b), rainbow trout, and red drum (J. Davis, Texas Agricultural Extension Service, Texas A&M University, personal communication) in the U. S.

In the 1981 draft of the Texas Aquaculture Plan, saline groundwater was identified as an untapped resource and it was suggested that high priority be given to determining the suitability of such water for mariculture (Stickney and Davis, 1981). Since that report, no formal research investigations have been carried out on the use of Texas saline groundwater for shrimp mariculture. Results of unpublished studies conducted since the early 1970s indicate that saline groundwater in some parts of western Texas may be suitable for crustacean culture on a commercial scale (J. Davis, Texas Agricultural Extension Service, Texas A&M University, personal communication). We examined the growth of penaeids in saline water originating from an inland groundwater source in southern Texas. For the purposes of documentation, the same population of P. vannamei was stocked into ponds near Corpus Christi, which received saline water from the Laguna Madre Bay. The bay system acted as a control, not in the true sense of duplication, but rather as a means of assessing the potential of the population to adapt to different conditions.

Materials and Methods

Groundwater System

The 0.2-hectare ponds were located west of Raymondville, Texas, in Willacy County, approximately 40 kilometers inland from the Laguna Madre (98°57' W, 26°28' N). Willacy County is underlain by the Gulf Coast Tertiary Aquifers, which are uniquely characterized by an abundance of sand and shale, minor amounts of limestone, and lack of carbonates. Salinity varies with depth (zero to 3048 meters) and from less than five to 120 parts per thousand (Texas Water Development Board, 1972). The source of saline water used in this study was an aquifer that flowed over a layer of caliche clay within four meters of the land surface. The ponds were dug directly into the aquifer layer, resulting in a continuous flow of water through all walls and the bottom, which prohibited complete draining and drying of the pond bottom between stockings. Water depth was a constant 4.5 meters.

The ionic composition of this saline (28.3 parts per thousand) groundwater (Table 1) was similar to that of both natural seawater (Sverdrup et al., 1942) and ponds fed from the Laguna Madre Bay system. The saline groundwater was of the sodium chloride type, which typifies groundwater samples in the U. S. having 20,000 parts per million or more of total dissolved solids (Feth et al., 1965). The main differences between the ionic compositions of the saline groundwater and natural seawater were the amounts of calcium and sodium.

Each of the two ponds was stocked with both juvenile P. vannamei (50,700 shrimp per hectare) and five-day-old postlarval P. setiferus (247,000 shrimp per hectare). P. setiferus (less than 0.01 gram) had been reared at the Texas A&M University Larviculture facility in Galveston from wild-caught broodstock that originated from the Gulf of Mexico near Port Aransas. P vannamei (1.2 ± 0.18 grams), spawned from Mexican maturation broodstock, had been reared in a 0.15-hectare nursery pond at the Texas A&M University Pond facility in Corpus Christi for 45 days at a density of 250,000 postlarvae per hectare. Both 0.2- hectare ponds were harvested 120 days after stocking.

PENAEID SHRIMP CULTURE

5

Table 1. Chemical composition of saline groundwater, Laguna Madre water and natural seawater.

Ion or	Groundwater		Laguna Madre		Natural SWa
element	mg/1	%	mg/1	%	mg/1	%
Calcium	615	6.6b	425	3.4b	400	3.2b
Magnesium	1275	13. 7b	1148	9.0b	1272	10.1b
Potassium	152	1.6b	371	2.9b	380	3.0b
Sodium	7291	78. lb	10741	84. 7b	10561	83. 7b
Bicarbonates	468	2.5C	781	3.6C	140	0.6°
Chlorides	16208	85. 6C	18425	85. lc	18980	87. 2C
Sulfates	2253	11. 9C	2441	11. 3C	2648	12.2C
Copper	<0.05		<0.05		0.001-0.01
Manganese	0.05		0.06		0.001-0.01
Zinc	<0.05		<0.05		0.005
pH	8.2		8.0		7. 5-8.4
Salinity (ppt)	28.3		34.3		34.3

a After Sverdrup et al., 1942: 173, 176-177, 194.

bPercent of each ion out of the total amount of calcium, magnesium, potassium and sodium in each sample.

'Percent of each ion out of the total amount of bicarbonates, chlorides and sulfates in the sample.

The 0.2-hectare ponds were not fertilized prior to, or during, the experiment. Because this was the first stocking into these ponds, any nutrient productivity was related to that occurring naturally in the water, was transferred with shipping water at stocking, or was influenced by artificial feed or proximity to cropland runoff. Salinity (refractometer) and transparency (Secchi disc) were recorded daily; temperature and dissolved oxygen (YSI model 58) were recorded daily at 0600 and 1700 hours. An aeration system consisting of two three-meter-long sections of diffuser hoses connected to an air compressor was installed by the middle of the experimental period and used daily thereafter.

A commercially prepared, pelleted feed (24 percent protein, Texas Farm Products) was fed twice daily. Feeding was suspended whenever the dissolved oxygen level fell below 3.0 parts per million. During the first four weeks of the experiment, the feeding level was determined according to the schedule in Table 2. Thereafter, the rate was determined according to the schedule of Chamberlain et al. (1981), based on weekly samples of 30 individuals. For feeding purposes, survival was estimated to decrease to 93 percent ( P. vannamei ) and 72 percent ( P. setiferus) by week five, with an additional one and three percent decrease, respectively, each week thereafter.

Bay System

The 0.10 hectare ponds were located at the Texas A&M University Shrimp Mariculture Research facility (described by Conte, 1975) near Corpus Christi, Nueces County, Texas. All ponds were dried and tilled prior to filling. Seawater used to fill and maintain pond levels was pumped through 500-micron filter bags directly from the Laguna Madre Bay. Approximately two weeks prior to stocking, each pond was fertilized with 77 kilograms per hectare of urea, 14 liters per hectare of 54 percent phosphoric acid, and 1360 kilograms per hectare of dried cow manure. Applications of inorganic fertilizers were applied as needed to maintain a Secchi disc depth of 30 centimeters. Freshwater was added as needed to

6

THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Table 2. Schedule used to determine feeding levels in groundwater-fed ponds during the first four weeks of culture, prior to initiation of weekly sampling.

Species	Week	Assumed % survival	Estimated weight (g)	% body weight
P. vannamei	0	100.00	1.0	—
	1	97.00	1.5	19.0
	2	96.00	2.5	12.4
	3	95.00	3.5	9.4
	4	94.00	4.5	7.6
P setiferus	0	100.00	0.01	—
	1	93.75	0.10	19.0
	2	87.50	0.20	19.0
	3	81.25	0.35	19.0
	4	75.00	0.60	19.0

Table 3. Stocking density of P. vannamei and hydrological variables (mean ± SD) in the groundwater-fed ponds and the bay-fed ponds.

Groundwater system Bay system

	Pond A	Pond B	n=3	n=3
Stocking density (number per hectare)	50,000	50,000	25,000	75,000
Salinity (ppt)	32+2	23+1	25+2	25+2
Temperature AM (C) PM	24.8+2.5 27.8+2.2	24.7+2.4 28.1 + 1.6	25.9+2.4 29.3+3.0	25.9+2.5 29.4+3.0
Dissolved oxygen AM (ppm) PM	3.4+1. 6 7.0+2. 7	3. 5+1.2 9.4+1. 9	4.9+0. 9 0.4+1. 2	5. 0+0. 8 9.2+1. 1
Transparency (cm)	54.3+16.3	38.3+7.8	40.2+12.5	43.6+13.0

maintain salinity near 25 parts per thousand. Hydrological data (temperature, salinity, dissolved oxygen and transparency) were monitored as described for the groundwater system.

Juvenile P. vannamei (1.60 ± 0.28 grams), from the same population used to stock the groundwater system, were stocked into ponds, in triplicate, at 25,000 and 75,000 shrimp per hectare (Table 3). Feed was distributed at a rate of eight kilograms per hectare per day for the first two weeks. Thereafter, feed was distributed according to the schedule of Chamberlain et al. (1981), based on weekly subsamples of about 0.3 percent of the initial stocking density and assuming a survival of 70 percent. Feeding was suspended when dissolved oxygen levels fell below 3.0 parts per million. Ponds were harvested 115 days after stocking.

Differences in mean values between systems were analyzed by general linear regression using the 1988 version of PC-SAS (Statistical Analysis System, Cary, Indiana) and considered significant at the five percent level.

PENAEID SHRIMP CULTURE

7

Results

Hydrology

Hydrological data from both the groundwater and bay systems are summarized in Table 3. Generally, hydrological conditions in groundwater-fed pond B more closely resembled conditions in the bay-fed ponds than did those in pond A. The almost 10 parts per thousand difference in salinity between the groundwater-fed pond A (32 ± 2 parts per thousand) and pond B (23 ± 1 parts per thousand) was due to the proximity of pond A to an area of saline surface deposits. No attempt was made during the experiment to adjust salinity in the groundwater-fed ponds. Mean water temperatures were similar between the two systems. During weeks 10, 13, and 15, water temperatures in the groundwater-fed ponds dropped an average 10, 5, and 9° C, respectively, in response to cold fronts. Up to 10 days were required for temperatures to return to pre-cold front levels. Corresponding drops in the bay-fed ponds were less severe (3 to 6° C) and recovery occurred within three to five days. Dissolved oxygen levels were similar between the two systems. Low (less than 3.0 parts per million) dissolved oxygen levels occurred on 44 mornings in groundwater-fed pond A and on 33 mornings in pond B, with nearly half of those levels between 1.0 and 2.0 parts per million. Dissolved oxygen levels in the bay-fed ponds were more stable than those in the groundwater-fed ponds, partly due to the more efficient aeration system. Lowest measured dissolved oxygen level in the bay-fed ponds was 2.3 parts per million. Levels between 2.0 and 3.0 parts per million occurred in isolated ponds on two occasions. Natural productivity, estimated using Secchi values, was substantially lower in pond A than in pond B. Transparency was more stable in the bay system than in the groundwater system, due to the strict fertilization regime.

Production

Despite hydrological differences, mean growth and final weight of P. vannamei and P. setiferus did not differ significantly between the two groundwater-fed ponds (Fig. 1). Mean final weight of P. setiferus (8.0 ± 0.50 grams, n=2) was significantly lower than that of P. vannamei (19.9 ± 0.71 grdms, n=2). Survival of P setiferus in the groundwater system, however, was less than one percent and the contribution of this species to the final production was negligible (about 2.0 kilograms per hectare).

Mean survival and final size of P vannamei grown in the groundwater- fed ponds were not significantly different from those of P. vannamei cultured in the bay-fed ponds (Table 4), although growth rate of shrimp stocked in the groundwater system more closely resembled that of shrimp stocked at the higher density in the bay system (Fig. 2). Production of P vannamei from the groundwater system stocked at 50,000 shrimp per

THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Figure 1. Apparent growth of P. vannamei and P. setiferus in saline groundwater-fed pond.

A (solid line) and B (dotted line).

hectare was within the expected range of the bay system stocked at lower (25,000 shrimp per hectare) and higher (75,000 shrimp per hectare) densities. Food conversion ratio was better in the groundwater-fed system than in the bay-fed system.

Discussion

Inasmuch as this was the first attempt to culture shrimp in saline groundwater from this particular source, the primary concern was to assess the potential of the system by documenting growth and survival of a penaeid species. Juvenile P. vannamei proved to be a successful species for culture in this saline groundwater system. Despite differences in

PENAEID SHRIMP CULTURE

9

Table 4. Stocking and production data for P vannamei in the groundwater and bay systems.

		Saline water source
	Groundwater	Bay
Density (number per hectare)	50,000	25,000	75,000
Percent survival, mean ± SD	86.7+4.53	84.8+18.42	91.0+8.73
Final size, mean ± SD (g)	19.9+0.71	20.8+2.14	17.6+1.90
Production, mean ± SD (kg/ ha)	869.7+14.3*	461.5+89.9	1191.3+45.5
Count/ pound, range	46-56	46-56	57-66
FCR	2.15:1	3.5:1	2.4:1
n	2	3	3

*Does not include contribution from P. setiferus.

hydrology, nutrient input, and pond design between the two systems, it is important to note that performance of P. vannamei stocked at 50,700 shrimp per hectare in the groundwater-fed ponds was consistent with that of shrimp from the same population stocked into Laguna Madre Bay-fed ponds near Corpus Christi at 25,000 and 75,000 shrimp per hectare. The fact that the groundwater-fed ponds were not fertilized makes their performance all the more noteworthy. Although the bay-fed pond system could not act as a control in the true sense of system and design replication, it did serve as a means of documenting the relative growth potential of P. vannamei in the groundwater-fed system. Improvements in pond design and aeration, as well as the initiation of a fertilization regime, may increase future yields from this system.

Postlarval P. setiferus did not prove to be a successful species under these experimental conditions. The larger size of P vannamei at stocking plus its ability to tolerate high density culture conditions (Chamberlain et al., 1981) are possible explanations for the much greater production of P. vannamei compared to P. setiferus. Although salinity and temperature ranges recorded during this study were within those reported for wild P. setiferus , Lindner and Cook (1970) proposed that the rapidity of temperature change is more critical to survival of P. setiferus than actual temperature extremes. The rapid drop in water temperature due to cold fronts coupled with the low morning dissolved oxygen levels observed for several days may have influenced mortality losses of P. setiferus from the groundwater-fed ponds. Oxygen levels between 1.0 and 2.0 parts per million occurred on 20 days in the groundwater-fed experimental system

10 THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Figure 2. Apparent growth of P. vannamei stocked into the saline groundwater system at 50,000 shrimp per hectare (dashed line), and into the bay system at 25,000 (dotted line) and 75,000 (solid line) shrimp per hectare.

and fell below 1.0 parts per million on four days. Seidman and Lawrence (1985) suggested a conservative estimate for critical dissolved oxygen level for growth to be 1.91 parts per million for juvenile P. vannamei and 2.22 for P. monodon. Levels above 2.0 parts per million have been recommended for pond culture of P aztecus (Kraner, 1975). The suitability of P. setiferus in this system should be readdressed under different stocking conditions.

Sarig (1984) credited the use of marginal saline waters unsuitable for agricultural irrigation as one reason for the general success of aquaculture in Israel. Although there are reports of experimental aquarium systems relying on saline water derived from wells or aquifers (Clark and Eister, 1971; Nakamura, 1971; Strasburg, 1971), U. S. researchers, in general, have been slow in recognizing the potential of saline groundwater for aquaculture. Goldstein (1986a, 1986b) reported successful culturing of Spirulina sp. and Crassostrea sp. in New Mexico saline groundwater. The Aquaculture Research Corporation, located at the mouth of the Barnstable Harbor in Cape Cod, Massachusetts, uses saline water pumped from an aquifer located about 20 meters below the complex to culture microalgae required to feed Mercenaria (Walsh et al., 1985).

Since 1973, members of the Texas Agricultural Extension Service and county extension agents, working with private individuals, have been investigating the commercial potential of stocking penaeids into western

PENAEID SHRIMP CULTURE

11

Texas ponds using saline groundwater sources (J. Davis, Texas Agricultural Extension Service, Texas A&M University, personal communication). Small farms presently are located in seven Texas counties (Dawson, Fisher, Howard, Kent, Martin, Pecos, and Ward) and areas near the Salt Fork of the Red and Brazos rivers also are being examined. Reasonably good success has been obtained in recent years with the availability of good seedstock. For example, one farm in Howard County produced approximately 1345 kilograms of live heads-on P. vannamei per hectare in 1988 (27 count, tails). Results from our research strongly suggest that saline groundwater in southern Texas also can be used to culture at least one penaeid species. The formal examination of other chemically similar bodies of saline groundwater in Texas for penaeid mariculture is encouraged.

Acknowledgments

The authors gratefully acknowledge Mr. B. W. Kirsch, the Kirsch family, and their employees for their dedication and determination to this project. Technical assistance from Michael A. Johns and Scott A. Davis also is appreciated. This work was sponsored in part by Texas A&M University Sea Grant College Program, supported by the National Oceanic and Atmospheric Administration, Office of Sea Grant, U.S. Department of Commerce, under Institutional Grant no. NA83AA-0-0061 , and by a grant from the Caesar Kleberg Foundation for Wildlife Conservation, A. L. Lawrence, principal investigator. Ms. Ginny Mitchell is acknowledged for preparation of initial drafts of this manuscript.

Literature Cited

Chamberlain, G. W., D. L. Hutchins, and A. L. Lawrence. 1981. Mono- and polyculture of Penaeus vannamei and Penaeus stylirostris. J. World Maricult. Soc., 12:251-270.

Chapman, V. J. 1960. Salt marshes and salt deserts of the world. Interscience Publishers, Inc., New York, 392 pp.

Clark, J. R. and R. Eister. 1971. Sea water from ground sources. Pp. 173-184, in Sea-water systems for experimental aquariums. A collection of papers (J. R. Clark and R. L. Clark, eds.), T. F. H. Pubis., Inc., Jersey City, New Jersey, 192 pp.

Conte, F. S. 1975. Penaeid shrimp culture and the utilization of waste heat effluent. Proc. Waste Heat Aquaculture Workshop, 1:23-47.

Feth, J. H. 1970. Saline groundwater resources of the conterminous United States. Water Resources Res., 6:1454-1457.

Feth, J. J. et al. 1965. Preliminary map of the conterminous U. S. showing depth to and quality of shallowest ground water containing more than 1000 parts per million dissolved solids. U.S. Geol. Surv. Hydrol. Invest. Atlas HA-199, Washington, D. C., 31 pp, 2 maps.

Goldman, J. C. 1979. Outdoor algal mass cultures — I. Applications. Water Res., 13:1-19. Goldstein B. 1986a. Saline groundwater aquaculture I. The growth of Spirulina spp. in the saline groundwaters of New Mexico. Pp. 64, in Proc. 17th Annual Meeting, World Mariculture Soc., Reno, Nevada.

- . 1986b. Saline groundwater aquaculture II. The growth of Crassostrea spp. in the

saline groundwaters of New Mexico. Pp. 12, in Proc. 1 7th Annual Meeting, World Aquaculture Soc., Reno, Nevada.

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Granoth, G., and D. Porath. 1983. An attempt to optimize feed utilization by tilapia in a flow-through aquaculture. Pp. 550-558, in Internat. Symp. on Tilapia in Aquaculture (L. Fishelson and Z. Yaron, compilers), Tel Aviv Univ., Tel Aviv, Israel, xi+624 pp.

Hamza, A. K., and M. I. Zaki. 1987. Experimental rearing of some marine fishes in brackishwater systems in Egypt. Bamidgeh, 39:39-48.

Issar, A., G. Oron, and D. Porath. 1983. Warm brackish groundwater as a source of supply for integrated projects of root zone warming, aquaculture, irrigation and recreation projects in arid regions. Pp. 106-115, in Internat. Symp. on Tilapia in Aquaculture (L. Fishelson and Z. Yaron, compilers), Tel Aviv Univ., Tel Aviv, Israel, xi+624 pp.

Kelley, D. B. 1982. Salt-affected rangelands: potential for productivity and management. Pp. 507-510, in Biosaline research: a look to the future (A. San Pietro, ed.), Plenum Press, New York, xiii+578 pp.

Kramer, G. L. 1975. Studies on the lethal dissolved oxygen levels for young brown shrimp, Penaeus aztecus Ives. Proc. World Maricult. Soc., 6:157-167.

Lindner, M. J., and A. L. Cook. 1970. Synopsis of biological data on the white shrimp Penaeus setiferus (Linnaeus) 1767. FAO Fisheries Rep., 4:1439-1469.

Nakamura, E. L. 1971. Salt well water facilities at the Bureau of Commercial Fisheries Biological Laboratory, Honolulu. Pp. 169-172, in Sea-water systems for experimental aquariums. A collection of papers (J. R. Clark and R. L. Clark, eds.), T. F. H. Pubis., Inc., Jersey City, New Jersey, 192 pp.

Pasternak, D. 1982. Biosaline research in Israel: alternative solutions to a limited fresh water supply. Pp. 39-57, in Biosaline research: a look to the future (A. San Pietro, ed.), Plenum Press, New York, xiii+578 pp.

Payne, A. 1. 1983. Estuarine and salt tolerant tilapias. Pp. 534-543, in Internat. Symp. on Tilapia in Aquaculture (L. Fishelson and Z. Yaron, compilers), Tel Aviv Univ., Tel Aviv, Israel, xi+624 pp.

Rains, D. W. 1979. Salt tolerance of plants: strategies of biological systems. Pp. 47-68, in The biosaline concept: an approach to the utilization of underexploited resources (A. Hollaender, J. C. Aller, E. Epstein, A. San Pietro, and O. R. Zaborsky, eds.), Plenum Press, New York, viii+391 pp.

Sarig, S. 1984. The integration of fish culture into general farm irrigation systems in Israel. Bamidgeh, 36: 16-20.

Seidman, E. R., and A. L. Lawrence. 1985. Growth, feed digestibility, and proximate body composition of juvenile Penaeus vannamei and Penaeus monodon grown at different dissolved oxygen levels. J. World Maricult. Soc., 16:333-346.

Stickney, R. R., and J. T. Davis. 1981. Aquaculture in Texas. A status report and development plan. Texas A&M Univ. Sea Grant Publ., TAMU-SG-81-1 19, 103 pp.

Strasburg, D. W. 1971. An aerating device for salt well water. Pp. 161-167, in Sea-water systems for experimental aquariums. A collection of papers (J. R. Clark and R. L. Clark, eds.), T. F. H. Publ., Inc., Jersey City, New Jersey, 192 pp.

Sverdrup, H. U., M. W. Johnson, and R. H. Fleming. 1942. The oceans: their physics, chemistry, and general biology. Prentice-Hall, Inc., Englewood Cliffs, New Jersey, x+1087 pp.

Texas Water Development Board. 1972. A survey of the subsurface saline water of Texas. Rept. Texas Water Development Board, 157 ( 1 ):vi+ 1 - 1 13 pp.

Walsh, D. T., R. A. Kraus, C. A. Withstandley, S. M. Talin and E. J. Petrovits. 1985. Dimensioning of a mass algal facility for the temperate zone nursery culture of bivalve molluscs. J. World Maricult. Soc., 16:451-463.

Winslow, A. G. and L. R. Kister. 1956. Saline-water resources of Texas. U.S. Geol. Surv. Water-Supply Paper, 1365:1-105.

BATS FROM THE COASTAL REGION OF SOUTHERN TEXAS

Sandra S. Chapman and Brian R. Chapman

Department of Biology, Corpus Christi State University, Corpus Christi, Texas 78412

Abstract. — A complete literature review revealed a paucity of information regarding chiropteran distribution and habitat along the southern coast of Texas. This survey describes distributional and ecological observations for 13 species of bats from that region. Key words: Chiroptera; distribution; natural history; Texas.

The coastal region of southern Texas offers a diversity of habitats suitable for bats, yet few surveys of their distribution in that region have been undertaken. In an unpublished survey of Texas mammals and birds, Lloyd (1891) reported several observations of bats in southern Texas, but identified only two species. Bailey (1905) recorded four species from Cameron and Hidalgo counties, only a small part of coastal southern Texas as here defined. Mulaik (1943) subsequently found five species in Hidalgo County. In his survey of Texas biotic provinces, Blair (1950) included only two bats from coastal southern Texas. Little information has been published on the distribution of bats in the region since Blair’s (1952) survey. Davis (1974) reported 11 species from the coastal region of southern Texas, but as Schmidly et al. (1977) noted, this reference lacks exact collecting localities and a listing of institutions where specimens are deposited.

To provide a more complete picture of distribution and seasonal abundance of the bat fauna of the coastal region of southern Texas, 438 specimens from 12 counties were examined. These included collections from field work in Cameron, Nueces, San Patricio, and Willacy counties from April 1987 until October 1988. Wherever possible, pertinent life history observations also are recorded.

Region Studied

The study area defined herein as “southern coastal Texas” includes those counties along the Gulf of Mexico from Refugio and Bee counties south to Mexico (Fig. 1). The western boundry includes Live Oak, Jim Wells, Brooks, and Hidalgo counties.

Coastal southern Texas is situated in the Tamaulipan biotic province of Texas (Blair, 1950). Oberholser et al. (1974) referred to the habitats in the Tamaulipan province from San Antonio to the Rio Grande as South Texas brush country and coastal prairie. A part of this province, including Cameron, Willacy, Hidalgo, and Starr counties is designated as the Matamoran district (Blair, 1950), from which Blair (1952) reported seven species of bats. Included in the Matamoran district are approximately 14 hectares of Rio Grande palmetto palms ( Sabal texana ), which occur naturally in a grove southeast of Brownsville, Cameron County. Resident populations of Lasiurus ega and L. intermedius may occur in this palm grove. The remainder of the Tamaulipan province is referred to as the Nuecian district (Blair, 1952).

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Bats were collected in Cameron County at the Fort Brown Hotel in Brownsville. Vegetation surrounding the hotel included Washington fan palms ( Washingtonia robusta ) and Rio Grande palmetto palms. Specimens were taken from the decorative columns on the second story of the hotel. These columns were composed of pumice rocks held in place by mortar. Bats were removed from gaps in the mortar using forceps.

One collection site in Nueces County was located 1.6 km. S Driscoll. A discription of the site is provided in Spencer et al. (1988). Bats were removed with insect nets from Washington fan palms. Bats also were collected from the Corpus Christi Public Compress cotton warehouses in Nueces County. Using forceps, bats were removed from gaps between wooden beams and concrete walls inside the warehouses.

Rob and Bessie Welder Wildlife Foundation was used as the collection site in San Patricio County. This refuge is located in a transitional zone between the Gulf Prairies and Marshes and the South Texas Plains (Thomas, 1975). The area where bats were taken was described by Drawe et al. (1978) as a Woodland-Spiny Aster complex. Specimens were taken over Moody Creek using mist nets.

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The study site for Willacy County was located on the Yturria Ranch, La Chata Division. Specimens were collected by setting mist nets and trip wires over an earthen cattle tank. The vegetation surrounding this site was dominated by large honey mesquite trees ( Prosopis glandulosa) and mixed brush.

Accounts of Species

Specimens examined were preserved either as skins accompanied by skulls or in alcohol and deposited in the following collections (museum abbreviations given in parentheses): Corpus Christi State University Vertebrate Collection (CCSU); Texas A&I University Vertebrate Collection (A&I); Pan American Univeristy Vertebrate Collection (PAU); Texas Cooperative Wildlife Collection, A&M University (TCWC); Texas Natural History Collection, University of Texas at Austin (TNHC); The Museum, Texas Tech University (TTU); U. S. National Museum of Natural History (USNM); American Museum of Natural History (AMNH); Welder Wildlife Foundation Collection (WWF). The arrangement of species and use of vernacular names follows Jones et al. (1988).

Mormoops megalophylla Peters, Ghost-faced Bat

The ghost-faced bat, referable (Smith, 1972) to the subspecies M. m. megalophylla , is known from Cameron and Hidalgo counties in the coastal region of southern Texas (Mulaik, 1943; Constantine, 1961; Davis, 1974; Hall, 1981; Jones et al., 1988). No additional specimens were collected or examined during this survey although this species may range along the Texas coast (Barbour and Davis, 1969).

This bat occupies a range of diverse habitats from desert scrub to tropical forests (Barbour and Davis, 1969). In Edinburg, Texas, four specimens were captured in January and February while hanging from a rough plaster ceiling in a junior high school (Davis, 1974). Apparently, ghost-faced bats do not hibernate, but no information is available on the subject (Jameson, 1959; Barbour and Davis, 1969).

Choeronycteris mexicana Tschudi, Mexican Long-tongued Bat

This species is known by a single record from Hidalgo County, represented by photographs only (LaVal and Shifflett, 1972). Jones et al. (1988) reported that this record probably represents an accidental northward occurrence. Choeronycteris mexicana is a monotypic species (Hall, 1981).

Mexican long-tongued bats occupy a range from tropical lowlands to montane habitats (Villa-R., 1967; Matson and Patten, 1975). The specimen from Hidalgo County was taken in December 1970 from a garage in the Santa Ana National Wildlife Refuge. The refuge was

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described by LaVal and Shifflett (1972) as mostly subtropical riparian forest.

Myotis velifer (J. A. Allen), Cave Myotis The cave myotis is known only from Hidalgo County in the southern coastal study region (Hayward, 1970; Jones et ah, 1988). The subspecies is M. v. incautus according to Hall, 1981.

These bats are usually cave dwellers, although they also roost in old, abandoned buildings and mine tunnels (Hayward, 1970; Davis, 1974; Schmidly, 1983). The species occupies a variety of habitats (Blair, 1952; Davis and Russell, 1952; Baker, 1956; Hayward, 1970; Davis, 1974; Schmidly, 1983). Some populations of cave myotis are known to hibernate (Hayward, 1970; Schmidly, 1977), whereas others probably migrate (Davis et ah, 1962; Hayward, 1970).

Pipistrellus subflavus (F. Cuvier), Eastern Pipistrelle Reports of this bat, referable to the subspecies P. s. subflavus , from the coastal counties are rare. It has been reported previously from Kleberg, Bee, and Cameron counties (Bailey, 1905; Davis, 1959; Zehner, 1985). Zehner (1985) recorded a specimen found in August in a building on Padre Island National Seashore. Additional specimens reported from the study region were collected in April. This bat hibernates (Davis, 1974), but no information is available on seasonal movements.

Specimens examined (2).— Bee Co.: 2 mi. N Beeville on U. S. Hwy. 181, 1 (CCSU). Ki.ebf.rg Co.: 2.4 mi. S Padre Island National Seashore, 1 (CCSU).

Lasiurus borealis (Muller), Eastern Red Bat

Red bats have been collected from Aransas, Refugio, Bee, Nueces, Hidalgo, and Cameron counties in the coastal region of southern Texas (Miller, 1897; Mulaik, 1943; Blair, 1952; Raun, 1966; Davis, 1974). This species also occurs in San Patricio and Kleberg counties. Lasiurus borealis is a monotypic species according to Baker et ah (1988).

According to Schmidly et ah (1977), red bats occur in all major vegetation regions. Museum specimens examined from coastal Texas were collected from May through October. Texas populations of this species may be migratory (Davis, 1974; Hall, 1981), although Schmidly et ah (1977) reported that this bat has been collected in the eastern part of the state in all seasons of the year.

Specimens examined (7). — Bee Co.: Beeville, 1 (TNHC). Aransas Co.: Rockport, 2 (AMNH). San Patricio Co.: Sinton, 2 (WWF). Kleberg Co.: Kingsville, A&I campus, 2 (A&I).

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Lasiurus cinereus (Palisot de Beauvois), Hoary Bat

The hoary bat, referable to the subspecies L. c. cinereus , has been reported from Cameron and Hidalgo counties (Bailey, 1905; Mulaik, 1943; Davis, 1974; Hall, 1981). Examination of museum collections also revealed a specimen of this species from Kleberg County.

Hoary bats have been found roosting in wooded areas hanging in the open from a branch or twig (Davis, 1974). One specimen (A&I 172) collected in Kleberg County was found roosting approximately one meter above the ground in a grapefruit tree. Specimens have been collected in the study region in January, April, May, September, and October. This species migrates southward in winter (Stones and Wieber, 1965; Davis, 1974), but probably does not hibernate (Findley and Jones, 1964; Stones and Wieber, 1965).

Specimens examined (11). — Kleberg Co.: Kingsville, 1 (A&I); 7.5 mi. S Kingsville in South Pasture, 1 (A&I). Hidalgo Co.: 1.25 mi. SE Edinburg, 1 (PAU); 5 mi. S Pharr, 1 (PAU); Weslaco, 1 (PAU). Cameron Co.: Brownsville, 6 (USNM).

Lasiurus ega (Gervais), Southern Yellow Bat

This species, previously believed to be rare in southern Texas, had been reported only from Cameron County (Baker et al., 1971). Recent collections show that southern yellow bats are common in Nueces County (Spencer et al., 1988). The species also has been collected in Kleberg and Hidalgo counties.

Southern yellow bats, referred to the subspecies L. e. panamensis by Baker et al., 1971, have been collected in the coastal region of southern Texas in most months of the year. In Brownsville, this species roosts in natural palm groves where it is considered to be a permanent resident (Baker et al., 1971; Davis, 1974). All specimens collected in Nueces County were found roosting in Washington fan palms (Spencer et al., 1988). Although most bats in the genus Lasiurus are solitary, many of the specimens from Nueces County were found roosting with Lasiurus intermedius. Most specimens from Nueces County were collected in March; however, this species has been observed flying and roosting there from February until November. Persons at the Driscoll study site reported having seen these bats there in all months of the year for many years, even though this species is thought to be migratory (Barbour and Davis, 1969).

Specimens examined (100). — Nueces Co.: 1 mi. S Driscoll, 4 (CCSU); Corpus Christi, 2 (CCSU). Kleberg Co.: Kingsville, A&I campus, 1 (A&I). Hidalgo Co.: no precise locality, 1 (PAU). Cameron Co.: 5 mi. SE Brownsville, 91 (1 A&I, 90 TTU); Brownsville, 1 (AMNH).

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Lasiurus intermedius (H. Allen), Northern Yellow Bat

This species may be a common resident in coastal southern Texas. It was known previously from San Patricio, Kleberg, Cameron, and Hidalgo counties (Bailey, 1905; Taylor and Davis, 1947; Davis, 1974; Hall, 1981). During field work, this species, referable to L. i. intermedius according to Hall and Jones (1961), also was collected in Nueces County (Spencer et ah, 1988).

Northern yellow bats have been collected from January to September in the study area. In Cameron County, they have been taken from the dead fronds of tall palms (Barbour and Davis, 1969; Baker et al., 1971). Most specimens from Nueces County were collected from Washington fan palms (Spencer et al., 1988). Although several bats of this species have been found roosting together in dead fronds (Barbour and Davis, 1969), L. intermedius had not previously been reported roosting with other bat species. At the Driscoll Ranch study site, northern yellow bats were found roosting with southern yellow bats (Spencer et al., 1988).

Specimens examined (48). — San Patricio Co.: Sinton, 1 (TNHC). Nueces Co.: 1 mi. S Driscoll, 5 (CCSU); Corpus Christi, 4 (CCSU); Kleberg Co.: Kingsville, Texas A&I campus, 20 (17 A&I, 1 TCWC, 2 TTU). Cameron Co.: no precise locality, 1 (TCWC); 5 mi. SE Brownsville, 5 (TCWC); Brownsville, 12 (4 AMNH, 8 USNM).

Lasiurus seminolus (Rhoads), Seminole Bat

The seminole bat has been reported only from Cameron County in coastal southern Texas (Hall, 1981). Although this species may range along the coast through Texas into Mexico (Barkalow and Funderburg, 1960; Barbour and Davis, 1969), we collected no specimens, and the one record from the region may be an extralimital occurrence or possibly represents a mididentified L. borealis. In any event, Jones et al. (1988) did list southern Texas as within the range of the seminole bat.

Spanish moss and pine-oak and long-leaf pine forests are favored roosting sites (Schmidly, 1983). Specific habitats in southern Texas remain unknown. These bats probably are active throughout the year (Davis, 1974). Although migratory patterns are not known, Barbour and Davis (1969) reported a definite distributional shift southward in autumn. The occurrence of this species in Cameron County is in need of verification.

Nycticeius humeralis (Rafinesque), Evening Bat

Evening bats have been reported from San Patricio, Bee, Kenedy, Refugio, Cameron, and Hidalgo counties in coastal southern Texas (Miller, 1897; Mulaik, 1943; Blair, 1952; Davis, 1974; Hall, 1981). In addition, we have examined museum specimens from Kleberg County and individuals have been collected in Willacy County.

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This bat, referable to the subspecies N. h. humeralis, was observed roosting in gaps under loose bark and cavities in mesquite trees ( Prosopis glandulosa ) in Willacy County. Several specimens also were taken from between decorative rocks at a hotel in Cameron County. Specimens from coastal southern Texas have been taken from January to September. Migration and hibernation abilities of this species are unknown.

Specimens examined (181). — Bee Co.: Mineral, 1 (A&I); 8 mi. N Beeville, 1 (TNHC). Refugio Co.: Woodsboro, 1 (TCWC). San Patricio Co.: 8 mi. N Sinton, 1 (TCWC); Welder Wildlife Foundation, Sinton, 4 (WWF). Kleberg Co.: Kingsville, 4 (A&l). Kenedy Co.: Encino Division, King Ranch, 3 (A&I); Rudolph, Norias Division, King Ranch, 14 (TCWC); 6 mi. ESE Rudolph, Norias Division, King Ranch, 5 (PAU). Hidalgo Co.: Bentson State Park, 5 (TCWC). Willacy Co.: 5.75 mi. N Raymondville, U. S. Hwy. 77, Bakke-Esparanza Ranch, 6 (TCWC); Yturria Ranch, La Chata Division, 80 (CCSU). Cameron Co.: Southmost Nursery, Brownsville, 2 (A&I); Southmost Ranch, Brownsville, 1 (A&I); Brownsville, 1 (AMNH); Brownsville, Fort Brown Hotel, 4 (CCSU); 5 mi. SE Brownsville, 48 (5 A&I, 43 TTU).

Antrozous pallidus (Le Conte), Pallid Bat

Pallid bats apparently are rare in coastal southern Texas and are known only from Cameron County (Davis, 1974; Hall, 1981). Specimens from this area are referable (Martin and Schmidly, 1982) to the subspecies A. p. pallidus.

Pallid bats are primarily cave dwellers, but have been captured roosting in man-made structures such as attics, barns, and abandoned buildings (Davis, 1974). The specimen examined from Cameron County was netted in September in a natural Rio Grande palmetto grove. Possibly this was a migrant, because pallid bats probably undertake seasonal movements (Vaughan and O’Shea, 1976).

Specimens examined (1). — Cameron Co.: 5 mi. SE Brownsville (TTU).

Tadarida brasiliensis (Saussure), Brazilian Free-tailed Bat

This bat, presumably referable to the subspecies T. b. mexicana, is common throughout the study region. It has been reported from Bee, San Patricio, Refugio, Jim Wells, Hidalgo, and Cameron counties (Bailey, 1905; Mulaik, 1943; Blair, 1952; Short et al., 1960; Davis, 1974). Based on specimens collected and examined, this species also occurs in Aransas, Nueces, and Kleberg counties.

This bat utilizes man-made structures as roosting sites in coastal southern Texas (Barbour and Davis, 1969). In Nueces County, specimens were collected and others examined at the cotton warehouse study site every month from April 1987 through August 1988. Bats were taken in Cameron County from the Fort Brown Hotel in the same manner as described for N. humeralis. This species is known to migrate.

Specimens examined (87). — Bee Co.: 6 mi. E Beeville, Hwy. 202, Medio Creek Bridge, 5 (CCSU); Poesta Creek, Beeville, 1 (CCSU). Aransas Co.: Rockport, 1 (CCSU). San

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Patricio Co.: Sinton, 23 (TCWC); Welder Wildlife Foundation, Sinton, 1 (WWF). Nueces Co.: Corpus Christi, 26 (24 CCSU, 2 USNM). Kleberg Co.: Kingsville, Texas A&I campus, 14 (A&I); Kingsville, 2 (TCWC); Padre Island National Seashore, 1 (CCSU). Hidalgo Co.: 13 mi. S Edinburg, 4 (TCWC); 5 mi. S Mission Anzaldouas Dam, 1 (TCWC); Edinburg, Pan American University campus, 5 (PAU); Santa Anna Wildlife Refuge, 1 (USNM). Cameron Co.: Fort Brown Hotel, Brownsville, 2 (CCSU).

Tadarida macrotis (Gray), Big Free-tailed Bat The only record of the big free-tailed bat in the southern coastal region is that of a male found hanging from a screen door in San Patricio County (Raun, 1961). The habitat surrounding the collection site was described by Drawe et al. (1978) as a live oak-chaparral community. Although little information is available, Davis (1978) suspected that this species hibernates in the Big Bend region of Texas.

Specimens examined (1). — San Patricio Co.: Welder Wildlife Foundation, Sinton, (WWF).

Acknowledgments

We wish to thank the following people for allowing us to examine specimens under their care: Drs. J. Knox Jones, Jr., Clyde Jones, and Robert J. Baker, The Museum, Texas Tech University; Dr. David J. Schmidly and Mr. George Baumgardner, Texas Cooperative Wildlife Collection, Texas A&M University; Dr. Allan H. Chaney, Texas A&I University; Dr. Frank W. Judd, Pan American University; Mr. Robert L. Martin, Texas Natural History Collection, University of Texas at Austin; Dr. Don E. Wilson, U. S. National Museum, Dr. Sydney Anderson, American Museum of Natural History; and Dr. James G. Teer and Mr. Gene W. Blacklock, Welder Wildlife Foundation. We are grateful to Mr. Michael McBain, of Corpus Christi Public Compress, Clay Pascal and his family. Dr. Michael E. Tewes, Texas A&I University, and Daniel Butler for giving us opportunities to collect in coastal southern Texas. This manuscript benefited from the constructive criticism by Drs. Genaro Lopez and John W. Tunnell, Jr. We also thank Paul Choucair, Karen Hooten, Cristie Wall, and Walter Kilgus for their assistance in collection of specimens.

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— — . 1966. A heretofore unnoted collection of Texas mammals. Texas J. Sci., 18:225-226. Schmidly, D. J. 1977. The mammals of Trans-Pecos Texas. Texas A&M Univ. Press, College Station, 27-57 pp.

— . 1983. Texas mammals east of the Balcones Fault Zone. Texas A&M Univ. Press, College Station.

Schmidly, D. J., K. T. Wilkins, R. L. Honeycutt, and B. C. Weyhard. 1977. The bats of East Texas. Texas J. Sci., 28: 1 27- 1 44.

Short, H. L., R. B. Davis, and C.F. Herreid, II. 1960. Movements of the Mexican free¬ tailed bat in Texas. Southwestern Nat., 5:208-216.

Smith, J. D. 1972. Systematics of the chiropteran family Mormoopidae. Univ. Kansas Publ., Mus. Nat. Hist., 56:1-132.

Spencer, S. G., P. C. Choucair, and B. R. Chapman. 1988. Northward expansion of the southern yellow bat, Lasiurus ega, in Texas. Southwestern Nat., 33:493.

Stones, R. C., and J. E. Wieber. 1965. A review of temperature regulation in bats (Chiroptera). Amer. Midland Nat., 74:155-167.

Taylor, W. P., and W. B. Davis. 1947. The mammals of Texas. Bull. Texas Game, Fish and Oyster Comm., 27:1-79.

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Thomas, G. W. 1975. Texas plants: an ecological summary. Texas Agric. Exp. Sta. MP- 585:7-14.

Vaughan, T. A., and T. J. O’Shea. 1976. Roosting ecology of the pallid bat, Antrozous pallidus. J. Mamm., 57:19-42.

Villa-R., B. 1967. Los murcielagos de Mexico. Univ. Nat. Auton. Mexico, xvi + 491 pp. Zehner, W. 1985. First record of Pipistrellus subflavus (Chiroptera: Vespertilionidae) on Padre Island, Texas. Southwestern Nat., 30:306-307.

A NUTRITIONAL ANALYSIS OF DIET AS REVEALED IN PREHISTORIC HUMAN COPROLITES

Kristin D. Sobolik

Department of Anthropology, Texas A&M University, College Station, Texas 77843

Abstract. — Nutritional analysis of archaeological material has not been extensively conducted. The main reasons for the lack of such studies include the problems inherent in archaeological preservation of dietary remains, and the lack of information on the nutritional content of prehistoric foods. Coprolites (dessicated human feces), however, are unique resources for determining the prehistoric diet of archaeological populations. This paper presents a nutritional analysis conducted on 38 coprolites approximately 1000 years old that were excavated from a dry, limestone rockshelter in Val Verde County, Texas. The nutritional analysis indicates that the prehistoric diet of these people was relatively nutritious, although other archaeological information from the botanical, faunal, and human remains needs to be assessed before a complete nutritional statement can be made. Key words: nutrition; coprolites; diet.

Nutritional analyses provide insights into the health of a population, supplying answers to previous unanswered questions, pinpointing nutritional deficiencies, and possibly exposing cause and effect relationships of specific dietary deficiences and health problems. Such cause and effect relationships include the lack of vitamin C in the diet and the development of scurvy (Ortner and Putschar, 1985), and the increase in osteoporosis with a calcium-deficient diet (Robson, 1972). The low quality of diet obtained after alkali treatment of corn (which is deficient in niacin and a niacin precursor, tryptophan) also is realized with such work (Wing and Brown, 1979). However, nutritional studies have been conducted mainly on modern populations, and references pertaining to such studies are directed mostly toward modern American diets.

Nutritional analyses in prehistoric archaeological settings have mainly been ignored (Yesner, 1980). One reason for the lack of such studies stems from the problems that researchers face in the initial phases of such analyses and with the subsequent interpretation of data. Archaeological data of any kind are a limiting factor. Samples recovered from an archaeological excavation are most likely the best preserved and most resilient of such samples, and so do not reflect the frequencies with which food items may have been used by the prehistoric population. This is especially true for archaeological food remains, both faunal and floral (Carbone and Keel, 1985). Another major problem with analyzing prehistoric food remains is determing what actually was used as a dietary resource and what was used in some other fashion, or is a contaminant.

This paper will discuss a nutritional analysis based on 38 coprolites

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excavated from Baker Cave, Val Verde Co., Texas. This analysis is unique in that it is an initial step in determining the nutrition of a prehistoric population. Coprolites are an excellent source of information for nutritional analyses because the contents are what was most likely eaten. Coprolites, however, will yield an underestimate of the amount of animal protein consumed by a group of people, as it has been estimated that all animal protein is digested, and larger animal bone remains generally are not ingested. Trace elements and minerals found in the water supply also will be underestimated. A nutritional analysis of coprolite samples thus is only a partial statement of the nutrition of a prehistoric group because coprolites only reflect one portion of the diet. Other aspects that need to be analyzed for a complete nutritional statement include food items represented in faunal and floral archaeological material, and prehistoric human remains.

Sample Location

Baker Cave is located in a cultural area known as the lower Pecos region. This region is situated in southwestern Texas and northern Coahuila, Mexico, specifically centering around the Pecos and Devils rivers and their confluence with the Rio Grande (Fig. 1). The environment in this area is semiarid, leading to increased preservation of prehistoric cultural material found in rockshelters and open sites. Baker Cave is specifically located in an ecotonal environment at the eastern edge of the Chihuahuan Biotic Province, bordered by the mesquite- chaparral zone of southern Texas, the oak-cedar zone of the Edwards Plateau to the east, and the sotol-lechuguilla zone to the west (Chadderdon, 1983).

Prehistoric occupation began in the lower Pecos region at least 9000 years ago (Shafer and Bryant, 1977; Shafer, 1981, 1986). Subsistence patterning was centered around the arid environment from which a conservative foraging adaptation was mainly followed (Williams-Dean, 1978; Dering, 1979; Stock, 1983; Lord, 1984; Sobolik, 1988a). Agriculture was not practiced in this region mainly due to the inhospitable environment, to the time-consuming nature of the practice, and to the relative success of the inhabitants in a hunting-gathering subsistence (Lee and DeVore, 1968).

The nutritional analysis of the food items observed in the Baker Cave coprolites is the first of its kind conducted on an actual sample of coprolites.

The most extensive nutritional analysis of the area (Winkler, 1982) included a review of ethnobotanical and coprolite research in the lower Pecos area as well as some nutritional values of a few of the botanical dietary items recovered.

PREHISTORIC HUMAN COPROLITES

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TERRELL /

CROCKETT

SUTTON

N:

\

V E

ROE

EDWARDS

1 HINOS CAVE

2 MURRAH CAVE

3 MOOREHEAD CAVE

4 FATE BELL SHELTER

5 EAGLE CAVE

6 CENTIPEDE CAVE

7 DAMP CAVE

8 COONTAIL SPIN SHELTER

9 MOSQUITO CAVE

10 ZOPILOTE CAVE

11 BONFIRE SHELTER

12 PARIDA CAVE

13 BAKER CAVE

Figure 1. Location of Baker Cave in Val Verde County, Texas (from Lord, 1984).

Due to the status of the prehistoric occupants of Baker Cave as hunter- gatherers and their longevity in the area, it has been assumed that these people were healthy, nutritionally prosperous, and physically strong. Osteological data analyzed by Marks et al. (1985) and reviewed by Reinhard et al. (1990), however, indicate problems of iron and other nutrient deficiencies, caries, enamel hypoplasia, and extreme dental wear. This information suggests times of stress and malnutrition in the seasonal rounds and overall lifestyle of the lower Pecos population. The nutritional analysis of the coprolite constituents will test these studies.

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Coprolite Constituents

The Baker Cave coprolites were studied using both micro- and macroremains (Sobolik, 1988a). Microremains included pollen and parasites, whereas macroremains included bone, hair, fiber, seeds, charcoal, and insects. Macroremains were divided into those associated with plants and those associated with animals. The major constituents of the coprolites are listed in Table 1. This table shows the percentage of the 38 coprolites that contained each constituent. Animal remains from coprolites were identified using bone, fur, feathers, and scales.

Table 1. The main constituents of the Baker Cave coprolites. Values represent the percent of the total number of coprolites in which a constituent was observed.

Constituent	Percentage of coprolites
Fiber	100
Onion Bulbs	29
Prickly Pear Seeds	16
Mustard Seeds	8
Juniper Seed Hulls	5
Mesquite Pods	3
Goosefoot Seeds	3
Bone	53
Fish	37
Rodent	34
Bird	18
Lizard	5
Rabbit	3

The pollen types most easily recognized as intentionally ingested items (Sobolik, 1988/?) include Brassicaceae (mustard family) pollen, Artemisia (sagebrush) pollen, and Poaceae (grass family) pollen. Eight coprolite samples from the Baker Cave latrine area contained extremely high frequencies of Brassicaceae pollen. Many of the samples contained aggregates of this pollen type, suggesting that anthers were eaten. Artemisia pollen also was observed at high frequencies, and aggregates of this pollen type were observed as well. Poaceae (grass) pollen was found at high frequencies in many of the coprolites analyzed, indicating intentional ingestion of grass inflorescences or seeds.

Human Dietary Requirements

The recommended dietary allowances of the committee on Dietary Allowances (1980) will be used to illustrate the essential dietary constituents needed by humans. Dietary requirements include carbohydrates, protein, and lipids (for energy), and fiber, vitamins,

PREHISTORIC HUMAN COPROLITES

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Table 2. Human nutritional requirements.

Carbohydrates	Lipids
Fiber	Linoleic acid
Protein	Linolenic acid
Amino acids	Fat-soluble vitamins
Alanine	Vitamin A
Histidine	Vitamin D
Isoleucine	Vitamin E
Leucine	Vitamin K
Lysine
Methionine
Phenylalanine
Threonine
Tryptophan
Valine
Water-soluble vitamins	Minerals
Vitamin C	Calcium
Thiamin	Magnesium
Riboflavin	Sodium
Niacin	Potassium
Vitamin B6	Phosphorous
Vitamin B12	Chlorine
Folacin	Sulfur
Biotin
Pantothenic Acid
Trace elements
Manganese
Iron
Copper
Iodine
Zine
Flourine
Selenium
Chromium
Molybdenum

minerals, and water (Table 2). These substances are directly converted to energy in order to maintain or rebuild body structures, or are stored for later use. The requirements that will be discussed reflect the average of the actual needs of a population (FAO/WHO, 1973). Some individuals need more and some less of the specific nutrients; therefore, only the group population is considered.

Recommended nutritional intakes have not been estimated for other populations. However, some studies (Srinivasan, 1981; Seckler, 1982) maintain that recommended dietary allowances should not be used to compare and analyze “populations that have adapted to different physical and socioeconomic environments” (Messer, 1986:57). Populations may

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adapt and survive on lower nutritional intakes than those deemed adequate for western societies (Stini, 1975), mainly by reducing energy expenditure with a decreased level of work and physical activity (Messer, 1986). Individuals also may “adapt to lower levels of energy and protein intake at no function cost” (Messer, 1986:58).

Energy is the essential outcome from use by the body of dietary constituents. Carbohydrates, lipids, and protein are the sources of energy for the human body (Committee on Dietary Allowances, 1980; FAO/ WHO, 1973). Energy needs are dependent on physical activity, body size, age, and climate (Committee on Dietary Allowances, 1980; FAO/ WHO 1973). Additional energy (kilocalories) also is needed for pregnant (+300 kcal) and lactating (+500 kcal) females. If energy intake is deficient, then part of the dietary protein is needed to provide that energy (FAO/ WHO, 1973). Physical activity is the main factor determining differences in energy requirements, and is variable within a population, especially delineated along gender lines and roles. The more active a person the more energy that person expends and needs replaced. Body size is also important in determining how much energy an individual needs. A person of large size requires “more total energy per unit of time for activities” (FAO/ WHO, 1973) than one of smaller size. The energy used in moving a larger body mass is higher (Committee on Dietary Allowances, 1980).

Age is also a significant determining factor in the amount of energy needed and expended by an individual. As a person grows older, basal metabolic rate and activity gradually decline. The resting metabolic rate has been estimated to decline two percent per decade (Durnin and Passmore, 1967), but the decline in physical activity varies for each individual (Committee on Dietary Allowances, 1980). Age also affects energy requirements due to changes in body composition and weight, and a possible increase in disease and disability (FAO/ WHO, 1973). More energy also is required for growth and metabolism. During childhood, proper energy intake is essential to fulfill the needs of an expanding and ever active body.

Climate also may influence the amount of energy expended. When performing activities in a colder temperature, energy requirements are increased, along with overall body temperature and metabolic rate (Johnson, 1963). Johnson (1963) also provided evidence that in temperate environments energy expenditure is greater than in warm environments, with the cold inducing the production of heat through shivering (FAO/ WHO, 1973). Differential human behaviors and adaptations to climate are great, however, and climate has been determined to be an unquantifiable energy correction by the FAO/ WHO (1973) and the Committee on Dietary Allowances (1980).

PREHISTORIC HUMAN COPROLITES

29

Nutritional Content of the Coprolite Food Items

Analysis of the nutritional aspect of a diet that is approximately 1000 years old has many obstacles. The main problem is that what was considered edible and “good eating” at that time is not necessarily viewed in that context today. Many of the Baker Cave dietary items are not recorded in modern day nutrition books and lists. Substitutions of the closest approximate item had to be made for some of the dietary constituents, such as frog ( Rana ) in place of lizard and pigeon in place of small birds.

Energy

In estimating efficiency of the diet of Baker Cave inhabitants, it must be realized that they were shorter in stature and lighter in weight than modern populations, which have been used to estimate recommended energy intake. Consequently, it is probably safe to assume that the Baker Cave occupants required a lower energy intake. However, the activity of the Baker Cave inhabitants would be much higher than that observed in a modern day population, due to their status as nomadic hunter- gatherers. In order to evaluate the amount of energy a population receives, carbohydrate, lipid, and protein intake must be considered. The main energy sources of the prehistoric population under analysis are those foods with the highest kilocalorie (kcal) value per 100 grams of edible portion, and with the highest lipid, carbohydrate, and protein content.

Food items recovered from the coprolites with the highest caloric values (Table 3) are prickly pear seeds (289 kcal), after they have been ground to a pulp in what is called “queso de tuna” (Winkler, 1982), goosefoot seeds (195 kcal), mustard seeds (469 kcal), birds (294 kcal), and rabbits (166 kcal). Foods with high protein and fat contents are similar and include birds, mustard seeds, rabbits, fish, and goosefoot seeds. Protein from meat would be underrepresented in the coprolites because meat is entirely digested, and bone remains from larger animals would not be observed in the coprolite specimens. Carbohydrate content cannot be determined due to the lack of data.

A “meal” of onion bulbs (35 kcal), a small fish (96 kcal), a bird (294 kcal), and mustard seeds (469 kcal) (Table 3), will provide at least 894 kcal and 49.72 grams of fat. With the addition of goosefoot seeds (195 kcal), another bird, and “queso de tuna” (289 kcal) the lowest energy intake is 1672 kcal. This amount of energy is probably sufficient for females and children, with the fat content more than sufficient for the daily requirements for both males and females (15 to 25 grams). With the addition of a few more staples, such as Opuntia fruits and especially nutritious seeds and meat, the daily energy intake would be sufficient to sustain males in the population.

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THE TEXAS JOURNAL OF SCIENCE—VOL. 42, NO. 1, 1990

Thus, mainly due to the higher caloric values, protein content, and fat content of birds, rabbits, mustard seeds, goosefoot seeds, and larger meat products not observed in the coprolites, along with the supplement of other less energy-sufficient dietetic items, the inhabitants of Baker Cave probably received sufficient energy.

Fiber

The main bulk of the diet of the prehistoric occupants of Baker Cave was that of fiber from many different sources. The main effect of fiber is to provide bulk to the diet, increase fecal output (Cummings, 1986), allow for easier food passage, and possible prevention of disease. Hunter- gatherers (prehistorically and historically) typically have had a diet that revolved around the consumption of fiber in bulk (Burkitt and Spiller, 1986). High fiber diets also have been observed for other prehistoric occupants in the Baker Cave area (Williams-Dean, 1978; Stock, 1983). Specific fiber content of foods usually have not been analyzed, but due to the high-fiber diet of Baker Cave occupants, their fiber intake was most likely nutritionally sufficient.

Protein and Amino Acids

Due to the problems in amino acid content conversion, the overall amino acids found in foods cannot be directly compared to the required amount. Infants require much more protein by weight than other age groups due to their continual expansion of tissue and bone. This protein requirement usually is met with the intake of human breast milk. An adult 70-kilogram male requires 56 grams of protein daily and an adult (55 kilograms) female requires 44 grams of protein daily (Committee on Dietary Allowances, 1980).

Foods high in protein are mainly the same foods high in energy content — goosefoot seeds, mustard seeds, lizards, rodents, rabbits, and birds. In my sample “meal,” the approximate protein content is 68.63 grams. This provides the necessary protein for the average adult male and female. Goosefoot seeds, fish, and rabbit are three food items found in the coprolites that have the highest protein content. It must be realized, however, that the protein content of larger game, not represented in the coprolites, would produce a much higher intake than what is represented here in the sample meal. Meat is one of the best and most efficient sources of protein in the diet.

Vitamins

Vitamins are divided into fat soluble (vitamins A, D, E, and K) and water soluble (vitamins C, B6, Bn, thiamin, riboflavin, niacin, folacin, biotin, and pantothenic acid). There is an overall lack of information on

Table 3. Nutritional content of food items from the Baker Cave coprolites (values given in 100 grams of edible portion).

PREHISTORIC HUMAN COPROLITES

31

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fat soluble vitamin requirements, as well as the requirements of vitamins B6, B 12, folacin, biotin, and pantothenic acid. This deficiency must be resolved before the complete nutritional status of the Baker Cave occupants can be determined.

Sixty milligrams of vitamin C daily is the recommended allowance for adults. This requirement could be met by eating two prickly pear fruits with the seeds, onions, or a few goosefoot seeds. Due to the high frequency of prickly pear seeds and onions in the Baker Cave diet, vitamin C probably was not a problem for the inhabitants.

Thiamin, vitamin Bi, is required at an average of 1.0 to 1.5 milligrams per day for adults. This requirement can be easily met with the ingestion of my “meal,” as well as through eating mustard seeds and catfish. Due to the availability of these foods, thiamin probably was present in the diet at the appropriate level to maintain health.

Riboflavin is needed in the diet in slightly higher amounts than thiamin (average of 1.2 to 1.7 milligrams recommended daily), and the same foods that contain high amounts of thiamin also contain high amounts of riboflavin. Thus the same food combinations that provided a nutritionally sufficient supply of thiamin also supplied that of riboflavin.

Niacin is required in amounts averaging 13 to 16 milligrams daily. Foods high in niacin include mustard seeds, chenopod seeds, and rabbit. All of these items were observed in the coprolites, and would provide an individual with a sufficient supply of niacin.

Minerals

The essential mineral content of food is fairly well represented, especially for calcium and phosphorous. The highest calcium content in the food array is goosefoot seeds (1402 milligrams), showing that 100 grams of this food item will provide the daily recommended calcium for all ages. Mustard seeds (521 milligrams) and onions (140) are also high in calcium. The requirements for phosphorus are the same as those for calcium (800 to 1000 milligrams daily). Mustard seeds are also high in phosphorus (841 milligrams) as are chenopod seeds (327) and rabbit (224). These food items thus could have supplied the daily requirements of calcium and phosphorus for the Baker Cave population.

Data on the human requirements of electrolytes — sodium, potassium, and chlorine, as well as magnesium, iron, copper, zinc, and sulfur — are scarce. Sodium, required in amounts from 1100 to 3300 milligrams for adults, is found only in relatively low amounts in the food items studied. Thus, a huge amount of these foods would need to be consumed in order to fulfill the recommended daily allowance for sodium. However, it is quite probable that a food item (or items) not studied contains higher amounts of sodium than that already observed. This is also the case for

PREHISTORIC HUMAN COPROLITES

33

potassium and chloride, although not enough information on chloride has been obtained to make even a generalized statement.

Drinking water contains a high number of these electrolytes, which are obtained from the soil. The content of the drinking water in the Baker Cave area has not been analyzed, but until further research can be conducted it can be assumed tentatively that the occupants of Baker Cave most likely were not deficient in sodium, potassium, or chlorine.

Trace Elements

As with other nutritionally important elements, not enough information is available on trace elements of food items. The trace element requirments for the inhabitants of Baker Cave, however, probably were met with the large amounts of dirt and sand that were consumed, and in drinking water. Soil contains the majority of the trace elements and in quantities that probably would not reach toxicity levels for humans. Dirt and sand were found in 91 percent of the coprolites, indicating that the Baker Cave occupants probably ingested enough trace elements.

Conclusions

Determination of the health of the Baker Cave occupants has been attempted through a nutritional analysis of the contents of latrine coprolites. It must be realized that contents of coprolites do not reflect the entire dietary intake, and other dietary remains must be analyzed before a complete nutritional statement can be made. Remains of coprolites are the parts of foods that have not been digested. Energy and nutrients that have been digested must be estimated from the remaining portion. The nutritional analysis also is biased toward the season in which the coprolites were deposited. In order to determine what the nutrition of the prehistoric population was like year-round, analysis of a larger sample size, as well as other dietary indicators, must be available.

It has been suggested that the lower Pecos populations had a nutritionally sound diet and an overall healthy existence, due to their stable occupation of the lower Pecos area for more than 9000 years (Shafer, 1986). The preliminary nutritional analysis conducted indicates that the Baker Cave occupants at approximately A.D. 900, as reflected through their coprolites, were generally healthy. Skeletal information, summarized by Steele and Olive (1990), and Reinhard et al. (1990) suggests that there was nutritional stress in the lower Pecos population. This “stress,” however, was mainly observed in the dentition, and most likely resulted from the rough, fiberous diet inherent in the lower Pecos area, and to the high sugar and carbohydrate content of many of the food items, such as prickly pear and mesquite.

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The nutritional status of the occupants of Baker Cave, as indicated in this analysis, appears to have been generally good, although a much more rigorous study needs to be conducted. Such a study should include an analysis of the archaeological faunal, floral, and human remains from the area. More information on the nutritional contents of food items in the lower Pecos region is also essential. Additional information is especially needed on trace mineral and electrolyte content, fat soluble vitamin content, and a few water soluble vitamin contents. With the addition of this information, a nutritional analysis would provide an extensive study and would greatly facilitate the knowledge that can be gained archaeologically about the lower Pecos prehistoric population, and about other similar groups.

Acknowledgments

D. Gentry Steele, Harry J. Shafer, and Vaughn M. Bryant, Jr., Texas A&M University, provided discussions and encouragement on all aspects of this analysis. The field work that provided the data for this study was supported by the Witte Museum of San Antonio, Texas, and the Center for Archaeological Research of the University of Texas at San Antonio. Project director was Dr. Thomas R. Hester and field director was Kenneth M. Brown.

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Souci, S. W., W. Fachman, and H. Kraut. 1981. Food composition and nutritional tables 1981/82. Wissenschaftliche Verlagsgeselischaft nbH Stuttgart, 1352 pp.

Srinivasan, T. N. 1981. Malnutrition: some measurement and policy issues. J. Dev. Econ., 8:3-19.

Steele, D. C., and B. W. Olive. 1990. Bioarcheology of the western portion of the Gulf Coastal Plain study area. In A cultural resources overview of Region 3 Southwest Division, Corps of Engineers (T. R. Hester and D. G. Steele, eds.), report submitted to the Arkansas Archeol. Surv. and the Southwest Div., Corps of Engineers, in press.

Stini, W. A. 1975. Adaptive strategies of human populations under nutritional stress. Pp. 19-41, in Biosocial interrelations in population adaptation (E. S. Watts, F. E. Johnston, and C. W. Lasker, eds.), Mouton Publishers, The Hague, 412 pp.

Stock, J. A. 1983. The prehistoric diet of Hinds Cave (41 VV 456), Val Verde County, Texas: the coprolite evidence. Unpublished M.A. thesis, Texas A&M Univ., College Station, 246 pp.

Williams-Dean, G. J. 1978. Ethnobotany and cultural ecology of prehistoric man in southwest Texas. Unpublished Ph.D. dissertation, Texas A&M Univ., College Station, 287 pp.

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Wing, E. S., and A. B. Brown. 1979. Paleonutrition: method and theory in prehistoric foodways. Academic Press, Inc., New York, 202 pp.

Winkler, B. A. 1982. Wild plant foods of the desert gatherers of west Texas, New Mexico and northern Mexico: dome nutritional values. Unpublished M.A. thesis, Univ. Texas, Austin, 85 pp.

Yesner, D. R. 1980. Nutrition and cultural evolution: patterns in prehistory. Pp. 85-115, in Nutritional anthropology: contemporary approaches to diet and culture (N. W. Jerome, R. F. Kandel, and G. H. Pelto, eds.), Redgrave Publishing Co., Pleasantville, New York, 433 pp.

ABNORMAL TERMINAL CRETACEOUS FORAMINIFERA OF EAST-CENTRAL TEXAS

Homer Montgomery

Department of Geology, University of Puerto Rico, Mayaguez, Puerto Rico 00708

Abstract. — Morphologically abnormal planktic foraminifera are present in the uppermost meter of the upper Maestrichtian Kemp Clay in east-central Texas. Morphologically abnormal foraminifera have been well described from Recent sediments, but not from fossil populations. Abnormal specimens are heavily concentrated in biserial genera Heterohelix, Pseudoguembelina, and Planoguembelina. Abnormal specimens are assigned to several categories based upon common abnormal morphological characteristics. The most unusual specimens are “twinned.” The majority of the deformities probably were associated with ecological deterioration or predation. Key words: foraminifera; Cretaceous- Tertiary boundary; Brazos River.

Upper Cretaceous rocks of the Kemp Clay containing abnormal foraminiferal faunas are present at Cretaceous-Tertiary boundary sections along the Brazos River in Falls County, Texas (Fig. 1). Section B2 contains abundant, well-preserved, planktic foraminifera. The planktic populations are neither diverse nor completely morphologically “normal.” Grossly abnormal specimens comprise approximately 0.5 percent of the complete study sample of 20,000, ranging up to five percent in individual samples. With the exception of foraminifera from other Kemp Clay localities and smaller concentrations of abnormal specimens recovered from Texas Turonian, Cenomanian, and Paleocene rocks, abnormal foraminifera from the Brazos Kemp Clay are found in a much greater percentage than I have encountered in other Cretaceous and Tertiary faunas in Texas. The Brazos percentages are low in comparison with other reported, Recent, deformed populations. The highest percentage of “abnormals” I am familiar with is a population of deformed Globorotalia menardii , which constituted up to 67 percent of the total observed specimens (Akttirk, 1976).

Little is known concerning the causes of abnormal morphologies in fossil foraminifera. Abnormal test morphologies in extant species have been attributed to: 1) stressful ecological conditions produced by detrimental water quality (Arnal, 1955; Watkins, 1961; Seiglie, 1974); 2) predation and wound healing (Aktiirk, 1976), and selective mechanical damage imitating predation (Be and Spero, 1981; Be, 1982). Under stressful environmental conditions (low oxygen and overly abundant organic matter), foraminifera were found to be smaller and had thicker tests than normal individuals (Seiglie, 1974). Abnormal test shape and unusual ornamentation was attributed to polluted environments (Watkins, 1961). Aberrant Globorotalia menardii were attributed to high water temperature (Lidz, 1966). Crushed shells resulted in “bizarre”

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specimens, whereas amputated individuals regenerated normally (Be and Spero, 1981). Test distortion and abnormal expansion of ultimate chambers were attributed to the effects of parasites or mechanical, traumatic damage (Akturk, 1976).

Biostratigraphy

The Cretaceous-Tertiary boundary is placed between samples 4 and 5 based on the initial appearance of the Tertiary forms Globoconusa daubjergensis and Globorotalia pseudobulloides (Fig. 2). Guembelitria cretacea persists undiminished at least 50 centimeters above the appearance of G. daubjergensis. Several heterohelicids, especially Heterohelix striata , are present in association with G. daubjergensis. The Paleocene foraminiferan Chiloguembelina sp. increases in abundance inversely to H. striata. Rugoglobigerina rugosa also is found in the basal Paleocene. Globotruncana aegyptiaca and Globotruncana duwi persist into uppermost Cretaceous strata. Planktic to benthic ratios vary considerably from 2.7 in the lowest part of section B2 to 1.0 just below the first occurrence of Paleocene foraminifera (Hansen et al., 1984, 1987). The planktic to benthic ratio in the basal Paleocene is no greater than 0.4. Bioturbation is probably responsible for some of the more obvious reworking of Cretaceous fossils into Paleocene strata. No general homogenation of sediment has occurred as only the foraminifera listed

ABNORMAL FOSSIL FORAMINIFERA

39

Figure 2. Outcrop profile showing samples locations and Cretaceous-Tertiary boundary.

above are found associated with Tertiary foraminifera, whereas coextant Cretaceous taxa are not.

The fauna suggests an incomplete boundary interval in spite of the uninterrupted appearance in outcrop. Abathomphalus mayaroensis and Pseudotextularia intermedia are not present, indicating probable absence of the terminal Cretaceous section. The Paleocene is missing the lowest Danian primitive globigerinids. The possibility of facies or paleogeographic control, or both, of these taxa is not impossible, but remains unproven.

Abnormal Morphologies

Foraminiferal abnormalities at the Brazos locality are concentrated heavily in the biserial Heterohelix group, which underwent a bewildering combination of torsion, extreme chamber shape, and number modifications, as well as growth of various special structures such as

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Figure 3. Abnormal foraminiferans of the Brazos River, Texas. Bar is one millimeter.

bulla or uncharacteristically overlapping chambers. Abnormal Paleocene foraminifera are concentrated in the globular trochospiral group, whereas biserial forms are mostly affected in the middle Cretaceous. Abnormalities can be divided into categories based on easily recognized

ABNORMAL FOSSIL FORAMINIFERA

41

aberrant morphologies. The categories below are presented in relative order of abundance from greatest to least.

Attached twins. — Attached twins most often appear as two foraminifera sharing the last few chambers (Fig. 3, 1-2). A much smaller proportion of twins have a common proloculus. Other specimens are composed of three or four complete but fused tests (Fig. 3, 3). The twin deformity is apparently only present in biserial species such at Heterohelix striata, Pseudoguembelina costulata, P. palpebra, and Gublerina robusta. Twins join one another most commonly at 30 to 60 degree angles, but tests may lie in the same plane or at 90 degrees.

Exaggerated expansion of final chamber. — Exaggerated ultimate chamber expansion is especially extreme in biserial taxa (Fig. 3, 4). Ultimate chamber volumes of two or three times that of normal specimens is not uncommon. Abnormal chamber size frequently is associated with increase in aperture size. An associated decrease in ultimate chamber ornamentation is common, but chamber perforation does not seem to vary.

Kinking. — Other foraminifera, especially Planoglobulina sp., are kinked or bent at an angle reflecting a change in growth direction (Fig. 3, 5). The kink is an offset in direction of chamber growth, not just an offset due to a missing chamber. Kinked individuals frequently posses chambers of radically expanded volume in post-kink position.

Bulla overgrowths. — Various bulla and unusual apertural coverings are present in many taxa severely complicating identification. Globotruncana is the dominant group involved. Bulla frequently partially cover apertures or wrap around the test, usually extending from the apertural area (Fig. 3, 6). Eccentric bulla appear not only in the Kemp Clay, but also upsection in Globorotalia pseudobulloides of the Paleocene Wills Point Formation.

Gaping or multiple apertures. — Racemiguembelina powelli, Pseudo¬ guembelina costulata, P. palpebra, Planoglobulina carseyae and P. acervulinoides possess gaping or unusual multiple apertures (Fig. 3, 7). Apertures extend completely across the test with openings approaching 25 percent of the chamber area.

Multiple ultimate chambers. — Multiple ultimate chambers are present mostly in biserial species, but also are found in Globotruncana sp. Biserial foraminifera have up to six chambers arranged radially around the ultimate chamber (Fig. 3, 8). One specimen of Globotruncana had two complete and identical ultimate chambers (Fig. 3, 9).

General monstrosities. — In the monstrosity category are various Globotruncana, Rugoglobigerina, and Heterohelix (Fig. 3, 10-12). Many monstrosities resemble the experimentally crushed specimens of Be and Spero (1981) in which one or more chambers were broken or collapsed

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MICROSPHERIC FORM

PHENOTYPES (PROBABLE GENOTYPES)

Figure 4. Illustration of proposed origin of twinned specimens through reproductive

complications.

and subsequently repaired. Fragments of crushed chambers have been retained, and the tests appear substantially thickened overall.

Discussion

Attributing the various deformities in planktic specimens to a single cause is not easily accomplished. Rather, specific deformation mechanisms are indicated at least at the generic level. This scenario does not preclude comprehensive pressures such as environmental or predatory stress.

Twined specimens with two proloculi and one ultimate chamber may have resulted from reproductive complications. Upon separation of nuclei of the microspheric form (with each gathering protoplasm) two nuclei (genotypically identical) remained in one mass of protoplasm (Fig. 4). Two proloculi formed. The protoplasm divided normally yielding two megalospheric individuals that remained adjacent each other. The individuals fused after secretion of one or more chambers. This process was apparently repeated with three and four proloculi.

The cause for radical ultimate chamber expansion along with an increase in apertural size or number (or both) remains difficult to

ABNORMAL FOSSIL FORAMINIFERA

43

explain. Olsson (1973) argued that kummerform (diminutive ultimate chamber) foraminifera are the norm, and that normal form (equal or slightly expanded ultimate chamber) may result from environmental stress. If this is the case, then by extrapolation the Brazos foraminifera suffered extreme stress. Hecht and Savin (1972) presented the opposite argument that kummerforms are stress related and associated with colder water temperatures.

Test kinking may represent incipient coiling except that the test remains relatively linear after kinking rather than continuing to coil. The recognized gerontic trend is uncoiling, thus the Brazos forminifera would not be a gerontic population. More probably kinking is a response to stress rather than incipient coiling.

Bulla placement may have functioned as apertural covering, possibly for protection. Bulla covering apertures would be a counter strategy to the biserial foraminifera with gaping apertures. Bulla development appears abnormal, and could attributed to stressful conditions. The normal growth cycle in these specimens appears complete. I believe bulla developed on unusually long-lived specimens. Hecht and Savin (1972) presented limited evidence that bulla are more common in warmer waters.

Foraminifera with two or more ultimate chambers are perhaps some sort of gerontic kummerforms. Multiple ultimate chambers tend to be downsized and of similar shape. If this is a sort of ultimate kummerform, then long life certainly is indicated.

Conclusions

The Brazos River foraminiferal monstrosities probably resulted from different mechanisms stimulated by stressful shifts in their environment. Twinned specimens indicate reproductive difficulties. Crushed and repaired specimens reflect increased predation by organisms such as copepods. Other abnormalities, including incipient coiling, development of bullae, and extreme ornamentation, may be directed toward defense. Rapid test expansion and the addition of oddly placed chambers reduce surface area in relation to greatly increased protoplasm volume — again a possible defense mechanism. Cretaceous planktic foraminifera definitely underwent size increase and complex structural development (keels, bullae, tegilla, and so forth). The Brazos monstrosities mimiced this trend with occasionally extreme results.

The next step in studying abnormal foraminifera is to better document their occurrence in other parts of the Cretaceous and Tertiary as well as in other areas of clearly different paleogeographic setting. Thin sectioning of specimens will better determine causal factors. For example, crushed and repaired foraminifera tend to have retained fragments and

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abnormally thickened tests. Examination of the common sutures in twinned tests should be intriguing.

The geological significance of the bizarre Brazos River foraminiferal populations remains unclear. Essentially instantaneous extinction scenarios at the end of the Cretaceous are prominant in the literature (especially Alvarez et al., 1980), and are not uncommonly inconsistent with much of the boundary data (Keller, 1989) or are based solely on speculation. Morphological monstrosities extant just below the notorious Cretaceous-Tertiary boundary would indicate preboundary stress rather than an instantaneous boundary extinction “event.”

<

Acknowledgments

I would like to thank the University of Texas at Austin for use of their scanning electron microscope as well as Ingrid Carre of the University of Puerto Rico for photographic work. Field assistance and advice from R. Farrand and H. Billman were invaluable.

Literature Cited

Aktiirk, S. E. 1974. Traumatic variation in the Globorotalia menardii (d’Orbigny) group in late Quaternary sediments from the Caribbean. J. Foram. Res., 6:186-192.

Alvarez, L., W. Ivarez, F. Asaro, and H. V. Michel. 1979. Extraterrestrial cause for the Cretaceous-Tertiary extinction. Science, 208:1095-1 108.

Arnal, R. E., 1955. Some occurrences of abnormal Foraminifera. Compass, 32:185-194.

Be, A. W. H. 1983. Gametogenic calcification in a spinose planktonic foraminifer Globigerinoides sacculifer ( Brady). Marine Micropaleo., 5:283-310.

Be, A. W. H. and H. J. Spero. 1981. Shell regeneration and biological recovery of planktonic Foraminifera after physical injury induced in laboratory culture. Micropaleo., 27:305-316.

Hansen, T. A., R. Farrand, H. Montgomery, and H. Billman. 1984. Sedimentology and extinction patterns across the Cretaceous-Tertiary boundary interval, Brazos River Valley, East Texas. Amer. Assoc, of Petrol. Geol., Field Trip Guidebook, pp. 21-36. Hansen, T. A., R. Farrand, H. Montgomery, H. Billman, and G. Blechschmidt. 1987. Sedimentology and extinction patterns across the Cretaceous-Tertiary boundary interval in East Texas. Cretaceous Res., 8: 229-252.

Hecht, A. D. and S. M. Savin. 1972. Phenotypic variation and oxygen isotope ratios in Recent planktonic foraminifera. J. Foram. Res., 2:55-67.

Keller, G. 1989. Extended Cretaceous-Tertiary boundary extinctions and delayed population change in planktonic Foraminifera from Brazos River, Texas. Paleoceanography, 4:287- 332.

Lidz, L. 1966. Deep-Sea Pleistocene biostratigraphy. Science, 154:1448-1452.

Olsson, R. K. 1973. What is a kummerform planktonic foraminifer? J. Paleo., 47: 327-329. Seiglie, G. A. 1974. Foraminifers of Mayaguez and Anasco Bays and its surroundings. Caribbean J. Sci., 14: 1-68.

Watkins, J. G. 1961. Foraminiferal ecology around the Orange County, California, ocean sewer outfall. Micropaleo., 7: 199-206.

SYNTHESES OF 2-(2-PYRIDYL)CYCLOHEXANONE AND RELATED CYCLOHEXANONES

Eldon H. Sund, Peter D. Koplyay, and Scott D. Wood Department of Chemistry, Midwestern State University,

Wichita Falls, Texas 76308

Abstract. — Five 4-alkyl-2-(2-pyridyl)cyclohexanones were synthesized by the condensation of morpholine enamine of the appropriate cyclohexanone with pyridine- 1- oxide in the presence of benzoyl chloride. In addition to cyclohexanone, 4-methyl, 4-ethyl, 4-isopropyl, and 4-/m-butylcyclohexanone were utilized. Hydantoin derivatives were prepared and the enol/keto ratios determined. Key words: enol/keto; tautomerization; hydantoin; cyclohexanone.

Our interest in enolizable ketones (Sund and Strickland, 1988) led us to a paper that discussed the synthesis of 2-(2-pyridyl)cyclohexanone (Fig. 1) and its enolization (Hamana and Noda, 1965). We synthesized a series of 4-alkyl-2-(2-pyridyl)cyclohexanones using their method, which consisted of condensing morpholine enamine of the appropriate 4- alkylcyclohexanone with pyridine-l-oxide in the presence of benzoyl chloride. The enamines were prepared from the requisite 4- alkylcyclohexanones and morpholine by a standard method and used without further characterization (Hiinig et al., 1973). These 4-alkyl-2-(2- pyridyl)cyclohexanones were yellow to orange oils and were characterized by conversion into hydantoin (Fig. 1) derivatives (Henze and Speer, 1942).

Table 1 lists the 4-alkyl-2-(2-pyridyl)cyclohexanones prepared as well as their boiling points, yields, enol/keto ratios and melting points, and yields of the hydantoin derivatives.

Experimental Section

All chemicals were purchased commercially, except 4-ethyl- cyclohexanone, which was prepared by the standard oxidation of 4- ethylcyclohexanol. Elemental analyses were performed by Huffman Microanalytical Laboratories, Golden, Colorado 80403. We determined melting points on a Thomas-Hoover melting point apparatus and they

Figure 1. Enol (left), keto (center), and hydantoin (right).

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THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Table 1. 4-alkyl-2-(2-Pyridyl)cyclohexanones and 8-alkyl-6-pyridyl-l ,3-diaza-2,4-dioxospiro [4.5] decanes (Fig. 1).

R	Yield %	Bp °C/mm	enol/ keto ratio	mp, °C	Yield %
H	46	135/0. 9a	2.3	258-259	45
ch3	32	125/2	1.5	295-297	87
C2H5	46	152/6	1.8	288-290	45
iso-C3H7	38	137/2	1.8	274-276	65
tert-C4H9	48	142/3	1.5	302-304	73

(a) reported bp 99-100/0.01 mm (Funke and Rissi, 1954); bp 138-140/0.13 mm (Hamana and Noda, 1965).

were corrected. Only one preparation was done for each ketone and thus the yields represent single preparations and could in all probability be improved. The following example illustrates the synthesis of 4-alkyl-2-(2- pyridyl)cyclohexanones.

4-ethyl-2-(2-pyridyl)cyclohexanone. — Fifty milliliters of chloroform and 5.7 grams (0.06 mol) pyridine-l-oxide were placed in a stirred flask. The enamine, 2-ethyl- 1-morpholino-l-cyclohexane, 23.4 grams (0.11 mol), was added, the reaction mixture cooled in an ice bath, and 9.9 grams (0.07 mol) benzoyl chloride added dropwise. The reaction mixture stood for three days at room temperature. The Japanese literature suggests that some pyridine- 1 -benzoate forms during this period of time. The benzoate is a facile leaving group, thus facilitating the acylation of the pyridine by the enamine. At the end of this time, the reaction mixture was acidified with 20 percent HC1 to a pH of 1 and the solvents removed under reduced pressure using a water aspirator. The residue was redissolved in five percent HC1 and the solution extracted with a toluene-ether mixture. The aqueous layer was made alkaline with solid K2CO3, extracted with methylene chloride, and after evaporating the methylene chloride, distilled under reduced pressure, collecting the fraction distilling at 152°C/(6 mm Hg). We obtained 5.6 grams (46 percent yield) of 4-ethyl-2- (2-pyridyl)cyclohexanone. A hydantoin was prepared, mp 286-288° C. The NMR spectrum was obtained with a Perkin-Elmer 90 MHz R-32 spectrometer. The NMR spectrum in DCCI3 showed the following characteristics: we assigned a complex series of signals at one to four parts per million to the various aliphatic protons, the number of peaks being doubled because both enol and keto tautomers were present in approximately equal amounts, a multiplet at 6.9-7. 7 ppm for the aromatic protons, with two sharp doublets at 8.5 ppm and 8.2 ppm due to the protons in the 6 position of the pyridine ring of the enol form and the keto form, respectively. Tetramethylsilane was used as an internal standard; the chemical shifts are reported in ppm relative to it all cases.

SYNTHESES OF 2-(2-PYRIDYL)CYCLOHEXANONE

47

Table 2. Analytical data for review. 8-alkyl-6-pyridyl-l,3-diaza-2,4-dioxospiro [4.5] decanes analysis.

		Calculated			Found
%c	%H	%N	%c	%H	%N
H	63.66	6.16	17.13	63.85	6.31	16.99
ch3	64.85	6.61	16.20	64.59	6.69	15.92
c2h5	65.91	7.01	15.37	65.61	7.08	15.07
iso-C3H7	66.88	7.37	14.62	66.85	7.46	14.93
/m-C4H9	67.75	7.69	13.99	67.54	7.75	14.15

The relative area of the enol proton at 8.5 ppm and the keto proton at 8.2 ppm were determined by integrating the peak areas. Several integrations were performed and the enol/ keto ratio was determined to be approximately 1.8. Elemental analysis for C, H, and N in agreement with theoretical values were obtained and submitted for review (Table 2).

Acknowledgement

We gratefully acknowledge financial support by a Robert A. Welch Foundation departmental grant.

Literature Cited

Funke, A., and E. Rissi. 1954. Pyridylcyclohexanones. Compt. Rend., 239:713-715.

Hamana, M., and K. Noda, 1965. Studies on tertiary amine oxides. XXIV. Reactions of aromatic N-oxides with enamines of cyclohexanone in the presence of acylating agents. Bull. Chem. Soc. Japan, 13:912-920.

Henze, H. R., and R. L. Speer. 1942. Identification of carbonyl compounds through conversions into hydantoins. J. Amer. Chem. Soc., 64:522-523.

Hiinig, S., E. Lucke, and W. Brenninger, 1973. 1-morpholino-l-cyclohexene. Pp. 808-809, in Organic syntheses, collective vol. V (H. Baumgarten, ed.), John Wiley and Sons, New York.

Sund, E. H., and S. K. Strickland. 1988. Syntheses of l-phenyl-2-(4-pyrimidinyl)ethanone and related ethanones. J. Chem. Eng. Data, 33:216-217.

AN EFFICIENT RETRACTABEE MOBILE ANTENNA TOWER FOR RADIO-TELEMETRY STUDIES

Matthew T. Pollock, Stephen Demarais, and Robert E. Zaiglin

Department of Range and Wildlife Management, Texas Tech University, Lubbock, Texas 79409, and Harrison Interests, Ltd., 602 Dorothy Jo Circle, Uvalde, Texas 78801 (REZ)

Abstract. — The design and construction of a retractable mobile antenna tower for use with null radio-telemetry receiving systems are described. We discuss the advantages it offers for radio-telemetry studies on large areas, study sites with rough roads, and localities where overhanging vegetation poses a problem for travel with a permanently erected mobile unit. The integrity of the antenna was maintained throughout the study because of the retractable design and its durability. Key words: antenna; mobile; retractable; storage; telemetry.

Various mobile telemetry antennas have been described for tracking radio-collared animals. A retractable, directional tower antenna was used with ruffed grouse, Bonasa umbellus (Marshall and Kupa, 1963). A directional loop antenna, mounted to a vehicle roof-top and rotated from within, was used to track striped skunks, Mephitis mephitis (Verts, 1963). Other vehicle-mounted antennas (Andelt, 1985; Balkenbush and Hallett, 1988) and antenna elevating devices (Kolz and Johnson, 1975) also have been described to aid in the collection of telemetry data.

Mobile antenna towers offer several benefits over land-based antennas. Areas devoid of permanent land-based antenna towers can be “explored” until the proper station (or stations) can be identified. Mobile antennas are particularly beneficial when animals move out of the receiving range for land-based antenna towers. Increased mobility allows the observer to reduce the distance to the animal (Hegdal and Colvin, 1986), thereby decreasing error polygons and overall error potential. Elevated antennas also improve signal strength and minimize antenna interference from motor vehicles (Kenward, 1987).

Mobile antenna towers do have some limitations associated with them. A permanently erected mobile antenna tower cannot be driven in areas with overhanging vegetation or under electric power lines. When using permanently erected mobile units, roads may have to be cleared of overhanging branches (Hegdal and Colvin, 1986), especially for a null system consisting of two yagis spaced one wavelength apart (183 centimeters). Additionally, supports for the antenna array may not protect the elements on rough roads, which may affect accuracy.

A completely retractable mobile tower alleviates problems of overhanging vegetation and rough roads. Furthermore, an antenna capable of total retraction provides a safe means for storage during travel and maintenance of the receiving system. Marshall and Kupa (1963) developed a retractable mobile directional antenna that was designed for

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THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Figure 1. Photograph of the mobile antenna tower in the fully retracted position ready for travel.

storage of a single yagi. Balkenbush and Hallett (1988) used a retractable antenna but did not describe the way in which the single yagi antenna was stored for protection. The present report describes the construction of an efficient, retractable null mobile antenna tower (hereafter referred to as the mobile unit), which includes storage of a dual yagi array.

Study Area and Methods

A study of white-tailed deer ( Odocoileus virginianus ) habitat utilization was conducted on the 42,510-hectare Piloncillo Ranch in Dimmit, Webb, and LaSalle counties, Texas. The ranch has a good network of roads, primarily of sandy loams, which frequently erode during storms. In addition, many of them are found in areas possessing overhanging woody vegetation.

Project objectives required two simultaneous readings upon the radio-collared animals with error polygons less than two hectares. To avoid purchasing and erecting more permanent land-based towers, we developed the mobile unit for use in conjunction with our existing land-based towers. The mobile unit had to withstand travel at moderate speed over rough, sandy roads with overhanging vegetation and electric power lines. Our mobile unit consisted of five parts: a null antenna array, a mast, an antenna holder, a vertical tripod mast support, and a horizontal mast support.

The null antenna array consisted of two, four-element, directional yagi antennas (Advanced Telemetry Systems, Inc., Isanti, Minnesota), separated by a cross boom 183 centimeters wide, and fastened with T-clamps. Coaxial cables extended from the respective yagis, along the cross boom, and down the mast.

The function of the mast was to elevate the antenna 1.5 transmitter wavelengths (2.74 meters) above the pickup truck cab, thereby maximizing signal strength and minimizing

MOBILE ANTENNA FOR RADIO-TELEMETRY STUDIES

51

A)

B)

Figure 2. Line drawing showing the antenna holder (A) in its proper orientation and the vertical tripod mast support (B) for use during data collection periods.

potential interference. A four-meter pipe, composed of one- and three-meter sections, functioned as the antenna mast. The smaller section was connected to the bed of the truck by a pivoting bracket, which allowed the mast to be erected to a vertical orientation and retracted to a near-horizontal orientation (Fig. 1). A steel rod fit securely into the bottom section and extended from it 30 centimeters. This allowed the three-meter section to be slipped over the steel rod and rotated in a full 360° circle. An adjustable compass rosette, attached to the bottom portion, was used to orient the antenna to true-north and to quantify animal locations. Upon erection of the tower, the mast slid securely into the vertical tripod mast support.

The holder held the array in the bed of the truck, protecting the antenna elements during travel. The antenna holder consisted of two 0.64-centimeters (0.25-inch) angle irons measuring 20.3 centimeters in length, a 7.6-centimeters section of 3.2-centimeters (1.25-inch) pipe, a sheet metal plate (eight by 20 centimeters), and a nut-and-bolt assembly. Two pieces of angle iron were welded together (Fig. 2A), and placed into the vehicle’s accessory mount. The sheet metal plate (eight by 20 centimeters) was welded to the angle irons so the plate extended over the inside of the truck bed by 10.2 centimeters. The pipe, welded vertically onto the sheet metal plate, held the antenna array during travel (Fig. 2A). A nut-and-bolt assembly on the side of the pipe allowed for tightening of the antenna array into the holder.

The vertical tripod support served to maintain the mast in an upright position during periods of telemetry data collection. It consisted of three sections of four-sided pipe (one

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THE TEXAS JOURNAL OF SCIENCE VOL. 42, NO. 1, 1990

meter long), an oval-shaped sheet metal plate (25.4 centimeters in diameter), and a latch for securing the mast in a vertical position (Fig. 2B). The tripod legs were secured to the bed of the truck by a bolt that passed through the surface of the bed. The oval-shaped sheet metal plate was welded to the top of the tripod legs. A U-shaped cut was made into the plate which permitted the mast to be placed in an upright position.

During transport of the mobile unit, the antenna mast was positioned horizontally on a support located on the truck bumper, similar to Marshall and Kupa’s (1963) design. The support consisted of a pipe, welded to a steel plate bolted to the bumper. A 13-centimeters section of 0.64-centimeters (0.25-inch) angle iron, welded to the top of the support, prevented the mast from falling sideways and into the bed. A hose clamp was moved down the mast over the angle iron during travel to hold the mast firmly in place.

Results and Discussion

Field tests were conducted to assess the error and utility of the antenna. Known and calculated bearings were compared for 15 beacons placed throughout the study area (Springer, 1979). Observations from 23 stations produced a mean standard deviation of 1.86° (range 0.24 to 5.60°). Travel at speeds up to 40 miles per hour did not affect the integrity of the antenna according to subsequent accuracy tests. Furthermore, set-up of this design required only 20 seconds and was not fatiguing to the operator. The ease of construction, maintenance, and set¬ up made this an efficient antenna design. The design permitted easy travel through areas with dense overhanging brush because antenna elements were shielded by the truck.

Approximately 12 hours were required for the construction of the mobile unit and the cost of the materials was inexpensive. The sheet metal, pipe, and angle iron used in construction, cost approximately $60. The antenna array was purchased commercially for about $257, making the total cost some $317.

In summary, our retractable mobile unit was a cost-effective product that offered flexibility for collecting telemetry data under the conditions described. A thorough coverage of our large study area was obtained without building many permanent land-based towers, clearing overhanging vegetation, or avoiding travel under electric power wires. Furthermore, accuracy tests proved that this design was an efficient one in which the integrity of the antenna was maintained.

Acknowledgments

We thank Thurman McNeil for equipment design and construction, and Fred Bryant, James Bergan, and David Haukos for manuscript review. This is contribution T-9-560 of the College of Agricultural Sciences, Texas Tech University, Lubbock.

Literature Cited

Andelt, W. F. 1985. Behavioral ecology of coyotes in south Texas. Wildlife Monogr., 94:1-45. Balkenbush, J. A., and D. L. Hallett. 1988. An improved vehicle-mounted telemetry system.

Wildlife. Soc. Bull., 16:65-67.

MOBILE ANTENNA FOR RADIO-TELEMETRY STUDIES

53

Hegdal, P. L., and B. A. Colvin. 1986. Radio telemetry. Pp. 679-698, in Inventory and monitoring of wildlife habitat (A. Y. Cooperider, R. J. Boyd, and H. R. Stuart, eds.), U. S. Dept. Interior, Bur. Land Manag., Serv. Center, Denver, Colorado, xviii + 858 pp. Kenward, R. 1987. Radio tracking. Pp. 115-150, in Wildlife radio tagging, Academic Press, Orlando, Florida, x + 222 pp.

Kolz, A. L., and R. E. Johnson. 1975. An elevating mechanism for mobile receiving antennas. J. Wildlife Manag., 39:819-820.

Marshall, W. H., and J. J. Kupa. 1963. Development of radio-telemetry techniques for ruffed grouse studies. Trans. N. Amer. Wildlife. Conf., 28:443-456.

Springer, J. T. 1979. Some sources of bias and sampling error in radio-triangulation. J. Wildlife Manag., 43:926-935.

Verts, B. J. 1963. Equipment and techniques for radio-tracking striped skunks. J. Wildlife Manag., 27:325-339.

ORIGIN OF COLOR OF AMERICAN INDIAN BLACK AND RED CERAMICS

M. Hyman and M. W. Rowe Department of Chemistry, Texas A&M University,

College Station, Texas 77843-3255

Abstract. — Results of studies on the chemical cause of the black coloration imparted to ferruginous clay ceramics that are fired under reducing conditions, and of the red color of these ceramics when fired under oxidizing conditions, are presented. Carbon and carbonaceous matter are ruled out as the principal blackening agent, as are iron and iron- oxide species of sizes greater than 300 nm, in the pottery studied here. The iron and iron- oxide system is likely responsible nonetheless, but if it is, the particle sizes of any such species must be less than 300 nm. In a red pottery sample studied, the principal coloring agent was hematite.

Early in this century, Hewett (1909a, 1909 b) excavated archaeological sites on the Pajarito Plateau in New Mexico (an area that later became widely known with the establishment of the Manhattan project and then the Los Alamos National Laboratory). During excavations, Dr. Hewett was intrigued by pottery found at the sites, which was matte black with shiny polished black designs. In order to more nearly establish the technical aspects of its production, Hewett requested the aid of Maria Martinez, a native American potter, to attempt to duplicate this pottery. Maria, along with her husband Julian, who had been a laborer on Hewett’s dig, were not successful, however, until shortly after the end of the First World War. After their rediscovery of the technique, this type of pottery became, and has remained, a popular style of “traditional” pottery in the San Ildefonso and Santa Clara Pueblo Indian settlements.

Although the necessity for reducing conditions is well known, the specific chemical cause of the black coloration of ancient ceramics has only recently began to be well understood (Hofmann, 1962, 1966; Hegde, 1966, 1975; Shepard, 1971; Hedges, 1975; Longworth and Warren, 1975; Noll et al., 1975; Longworth and Tite, 1979; Rogers, 1979; Maggetti et al., 1981; Maggetti and Schwab, 1982; Gilles and Urch, 1983; Chazan and McGovern, 1984; Makundi et al., 1989). Gilles and Urch (1983) summarized and expanded on the usual attributions in potteries from other countries:

“Iron oxide and hydroxides present in ferruginous clays, or added in the form of ocherous earths, for example will, on firing under reducing conditions, form spinel phases which are predominantly black in colour. The most common of these are hercynite (FeOALCF) and its solid solution series, and magnetite (Fe304) (Hofmann 1962, 1966, Longworth and Tite 1979, Longworth and Warren 1975, Maggetti et al., 1981). Iron- rich silicates containing a high percentage of ferrous ions occur naturally

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THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. I, 1990

in some clays and are another possible cause of black colouration provided that they are not oxidised or destroyed during firing. In ceramics in which vitrification has occurred, a black colour could be produced by ferrous ions in a glassy matrix (Hegde 1975).

“Clays rich in organic material, or to which carbonaceous material has been added, will fire black in colour under reducing conditions, owing to the formation of carbon. Alternatively, carbon can be deposited in the surface of wares by firing in a smoky atmosphere (Noll 1977) or by application of a ‘carbon’ paint (Shepard 1971) or possibly bitumin (Rahtz and Greenfield 1977).”

Almost all of the above references dealt with the black paints used on ancient ceramics from various regions. Most of these styles differ considerably in detail from the American Indian potteries studied in this work; we restricted our study to pottery that is black (or red) throughout. Black coloration of ancient and modern American Indian ceramics is generally attributed to the presence of carbon (or carbonaceous matter), or to manganese oxide, or to one of the several iron oxide compounds. A critical examination of reported studies reveals little unambiguous or even strong evidence supporting these conclusions when the discussion is limited to ceramics with both black surface and fabric.

We will discuss these possibilities below as we describe our own studies attempting to understand the coloration in modern, “black on black” (B/ B) pottery of the San Ildefonso and Santa Clara Pueblos in northern New Mexico, and by implication, ancient American Indian ceramics, and other archaeological ceramics throughout the world with black color occurring throughout the pottery. We are not considering the surface blackening of ceramics with nonblack fabric, which occurs during use over fires, or by painting with the various agents mentioned above, although the cause of the black coloration may be closely related in some cases; studies referenced above are almost all of this situation. We took broken, modern American Indian pottery, a red on red pot and a black on black pot from the Santa Clara, New Mexico, Pueblo as our objects of study and supplemented those with pottery samples prepared from local clay and fired to produce both red and black colored ceramic samples.

Experimental Methods

Two broken modern American Indian ceramic pots were purchased for study. One was a carved, polished black on matte black Santa Clara Indian pot. Upon breaking the pot into smaller samples for study, the black coloration was seen to extend throughout the pot. It was uniformly black with no red or paler coloration being observed. For comparison, a polished red on red Santa Clara pot also was purchased for study.

We also prepared a series of both red and black ceramic samples from a local clay for comparative studies. For these, a sample of dry clay was sifted through an Allen-Bradley

COLOR OF AMERICAN INDIAN CERAMICS

57

no. 35, 500-micron screen and mixed well to ensure homogeneity. The sifted dry clay then was mixed with water to create a wet, moldable clay that was rolled out into a rope of approximately one centimeter in diameter, cut into buttons two to three mm thick, and allowed to dry in a desiccator for two weeks. Twenty-eight such buttons were made and adjacent buttons were selected to be fired alternately black and red, that is, under reducing and oxidizing conditions, respectively. A few interspersed buttons were saved unfired.

For firing the pottery, a homemade kiln fueled with propane was used. The samples to be fired under oxidizing conditions were simply laid on the kiln floor and heated. The samples to be fired under reducing conditions were placed in layers of sawdust and charcoal in a two-inch diameter steel pipe. These remained in the pipe throughout the firing and subsequent cooling processes. A charcoal-sawdust mixture was added to the pipe as it burned away during firing. The firing temperature of 840° C was reached approximately 30 minutes after the kiln was lit and was held for one hour to ensure equilibration of the samples within the pipe. After cooling, we removed the red pieces, still too hot to touch, which were at that time black, but which turned red as they cooled and oxidized in air. Most of these red buttons had broken during firing and cooling as we had not added temper. We allowed the black pieces to cool overnight inside the pipe. Removal of the black buttons the next day confirmed that they were indeed black and none of these had broken during either the firing or the cooling process. Subsequently, we broke open some red buttons and some black buttons to confirm that the surface coloration occurred throughout the samples.

Results and Discussion

We utilized a number of analytical methods in an attempt to elucidate the chemical origin for production of the black color arising from a red clay, which is fired under reducing conditions, including chemical analysis of carbon and manganese, thermomagnetic analysis, x-ray diffraction, and low-temperature/ low-pressure oxygen and hydrogen plasma react¬ ions. Each of these techniques was chosen because of its utility in examining various possibilities. Nonetheless we cannot yet say unequivocally in a detailed way what precise chemical reaction(s) are responsible for the color transformation. We will discuss below each of the techniques we used and the information indicated by each. In large measure, we were only able to eliminate some of the possibilities often ascribed as the reason for the production of the black coloration.

Carbon and Manganese Analyses

Our first experiment was designed to test the suggestion that firing clays in environments rich in organic material, as in the American Indian technique of smothering a fire with dried horse manure, will cause the incorporation of carbon into the ceramic matrix and, therefore, lead to black coloration. This was the explanation given to us by several archaeologists in the Santa Fe, New Mexico, area when we initially enquired whether the cause for this transformation in color was known. However, the elemental carbon analyses we conducted indicated that the carbon content of both the red and the black potteries were too low to cause significant blackening. Values of less than 0.4 percent carbon were

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THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

found in all samples with no significant difference found between red and the black ceramics. In some analyses, the red had slightly higher carbon content and in some carbon was higher in the black samples. To establish the degree of blackening that might be expected by the presence of elemental carbon, we added 1 percent carbon black to a white (kaolinite) clay; only slight darkening compared to the color of the black pottery samples was observed. The white clay became off-white, not even darkening to a notable gray color. We conclude that formation or implantation of elemental carbon is not a factor in determining black coloration in potteries of the sort studied here. This conclusion confirms that of Makundi et al. (1989). We will include more about this in our discussion of the plasma reactions, where this conclusion is strengthened and extended to include carbonaceous matter in general.

Although we have measured the manganese contents of several varieties of Etruscan ceramics from Murlo, Italy, in our laboratory (Tobey et al., 1986), it is not clear that elemental analyses of manganese will shed much light on the suggestion that reduction of manganese oxides causes black coloration in some ceramics. It is likely that both the oxides and metallic manganese will be black in the finely divided state encountered in the pottery samples. We see no reason to suspect that metallic manganese will color ceramic fabric significantly darker than will oxides. Our analyses showed no significant enrichment of manganese of Etruscan ceramics. Manganese and its oxides do not play a significant role in the coloration of the ceramics of the type considered here.

Thermomagnetic Analysis

We previously had utilized thermomagnetic analysis of carbonaceous chondrites to establish the presence of magnetite (Fe304) in those black colored meteorites similar, at least superficially, to the color of the black pottery and used saturation magnetization to measure percent of magnetite in those samples (Larson et al., 1972; Watson et al., 1973; Herndon et al., 1974; Hyman and Rowe, 1983, 1986). The presence of metallic iron also can be established by the same technique. For instance, a clear indication of the presence of both metallic iron and magnetite in the carbonaceous meteorite, Novo-Urei, is demonstrated in Figure 1. Magnetite is indicated in Figure 1 with a Curie temperature of approximately, 590° C; that is, the magnetization curve drops rapidly near 590° C as the magnetic magnetite is rendered nonmagnetic by the thermal agitation. Metallic iron also is illustrated in Figure 1 by the presence of a second Curie temperature near 790° C, that is, the point at which the magnetization shown by the second plateau disappears. Hematite (Fe2C>3), which is less magnetic than either the magnetite or metallic iron, is characterized by a Curie temperature at 675° C; as

COLOR OF AMERICAN INDIAN CERAMICS

59

1.2

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Curie Point of Magnetite

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3 h a

□ □

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° □

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NOVO-UREI

200

400

600

800

TEMPERATURE, °C

Figure 1. Saturation magnetisation versus temperature curve for the Novo-Urei carbonaceous meteorite. Thermal analysis clearly shows both the presence of magnetite (Curie temperature causing an inflection at approximately 590° C) and metallilc iron (Curie temperature at 790° C) in this sample.

indicated by the thermomagnetic curve for Novo-Urei pictured in Figure 1, no hematite is present.

Because the partial reduction of the red iron oxide, hematite, to the black oxide, magnetite, has been suggested as producing the black color¬ ation in ceramics fired under reducing conditions, we decided to subject various red and black pottery samples to thermomagnetic analysis. Budd- ington and Lindsley (1964) have shown that hematite will not reduce to magnetite at 500° C unless the oxygen fugacity is less than 1(T18 atmospheres. However, Kobayashi and Schwartz (1966) found hematite converting to magnetite in a red sandstone in a thermomagnetic run in air. It thus is probable that, under conditions used by native American Indian potters, magnetite (and even metallic iron in some cases) will form. In the simplest possible case, that is, if pure hematite were the cause of the red coloration on red pottery, the samples should lose their magnetization near 675° C. The black pottery, on the other hand, if blackened by the conversion of hematite to magnetite, should lose its magnetization near 590° C. A typical sample of the red Indian pottery showed a Curie temperature of approximately 675° C, as seen in Figure 2B, indicating the probable presence of hematite. Results on black Indian pottery did not show the presence of magnetite (or metallic iron), however. For example, a black Indian pottery sample studied showed a Curie temperature of 525° C, as set forth in Figure 2A. In none of the

60

THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Z 1.5

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LU

Z 1.0

o

<

z

o

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0 c

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TEMPERATURE, °C

Figure 2. Saturation magnetisation versus magnetisation curves for a sample of black Indian ceramic (A), a sample of red Indian ceramic (B), and a homemade black pottery button, (C) (see text). Whatever causes the curve for the black sample of Indian pottery, it is not predominantly magnetite nor metallic iron. The curve C for the homemade black button was suggestive of a small component of magnetite (minor inflection at 590° C) and metallic iron (Curie temperature at 790° C). See Figure 1 for comparison, where both magnetite and metallic iron are clearly indicated.

■ _ C

a n ♦ ♦

Curie Point of Magnetite

♦ ■

□ Santa Clara B/B a

♦ Santa Clara B/B . TAMU Black

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samples of black Indian pottery selected for thermomagnetic analysis did we see a Curie temperature that would suggest that either magnetite or metallic iron was responsible for the thermomagnetic curve observed. A thermomagnetic curve (2C) of a homemade black pottery button (TAMU Black) did show a hint of an inflection at 590° C, which most probably was caused by the presence of a slight amount of magnetite, as well as a pronounced Curie point at approximately 780° C, which was almost certainly due to metallilc iron. It seems that the reductions we carried out using the charcoal-sawdust mixture were at a lower oxygen fugacity, that is, a more highly reducing environment, than the Indian pottery samples we investigated. Oddly, after sitting in the open, these black buttons slowly reoxidized to red. Apparently, the iron metal we produced was more easily oxidized in the atmosphere back to hematite than is the blackening agent in the Indian ceramics, which do not redden under the same conditions even after 70 years or more.

We conclude that reduction of hematite, which is present in the red pottery, to relatively pure magnetite does not play a dominant, if significant at all, role in producing the black coloration, in ferruginous clays fired under the conditions typically used by the modern American Pueblo Indians. Our thermomagnetic evidence does not preclude

COLOR OF AMERICAN INDIAN CERAMICS

61

formation of another, nonmagnetic, iron oxide, for instance wiistite (FeO), which may have a strong effect on the black color. Wiistite was ruled out by x-ray diffraction, however (see next section). There is a critically important caveat that we must place on the conclusion that magnetite (or other iron oxides and other spinels) plays only a minor role in causing black color. Thermomagnetic analysis will not detect magnetite particles that are superparamagnetic, that is, particles below approximately 300 nm in diameter. It is conceivable, even likely, that reduction of the hematite in ceramics results in the formation of magnetite particles less than 300 nm in size, which may nonetheless contribute to the black color. A scanning electron microscope survey of the black American Indian pottery indicated that fine magnetite was feasible because of the fine-grained nature of the pottery fabric. Magnetite grains that were greater than one micrometer would have been visible with the scanning electron microscope.

The possibility of superparamagnetic magnetite in the black ceramics could be investigated through use of low-temperature (liquid nitrogen) Mossbauer spectroscopy, a technique not available to us at this time. Other workers (Hedges, 1975; Longworth and Tite, 1979; Longworth and Warren, 1979), using Mossbauer spectroscopy on black potteries of a different sort, found that magnetite played a significant role in determining the color of black glazes in Chinese, Greek, and East Indian, and finally in Etruscan ceramics. A recent report by Makundi et al. (1989) using Mossbauer spectroscopy concluded that black coloration in early Cypriote and Nubian C-group black-topped potteries, coloration that seemed to have been achieved by the firing process as is the case for the Indian potteries studied in our work, “is due to Fe2+ ions in non¬ magnetic compounds. . . . The refiring experiments confirmed that the presence of carbon is not necessary for the black color.”

X-Ray Diffraction

X-ray diffraction is another technique that is informative concerning the presence of crystalline materials, and that exhibits characteristic peaks for magnetite and other iron oxides. We began with x-ray diffraction spectrometry of a sample of red Santa Clara pottery. The major peaks were, as expected, due to quartz and hematite as is shown in Figure 3. This was followed by an x-ray diffractometry spectrum taken on a sample of black Santa Clara pottery that gave quite unexpected results. Only one small unidentified peak, except for those of quartz, was seen above background. In a further attempt to investigate this puzzling situation, we ground a piece of the black pottery, subjected it to a magnetic separation, and then ran an x-ray diffraction spectrum on the magnetic fraction. The results are shown in Figure 4. Even after the magnetic separation, there is only weak indication for the presence of

RELATIVE INTENSITY

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THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Figure 3. X-ray diffractometry spectrum of a sample of red Santa Clara Indian pottery. Major peaks are seen due to quartz and hematite, as expected.

magnetite, Fe304, and also for maghemite, a different form of Fe304. The large peak at 2.88 A is unidentified and did not appear in the run before the magnetic separation. No FeO peaks were observed in the x-ray diffraction spectra. As in thermomagnetic analysis, the presence of extremely minute particles of magnetite or maghemite, or both, would not be detected by x-ray diffraction. Again with the critical caveat, that is, that magnetite or maghemite, or both, may be present as fine particles of less than 300 nm, we conclude that neither magnetite nor maghemite are responsible for the black coloration in the Indian ceramics studied here. The lack of distinctive x-ray peaks in the black pottery remains a mystery, unless the compounds responsible for the color are present as extremely small particles, a situation that seems likely when all the evidence presented in this paper is considered. An additional x-ray diffraction spectrum showed minor peaks due to hercynite, FeOAh03. Unless the iron oxides are present as particles less than 300 nm, we are left with a quandary. In what chemical form are the iron oxides after reduction in the black pottery that were demonstrated before reduction to be present in red pottery as hematite?

COLOR OF AMERICAN INDIAN CERAMICS

63

A

Quartz 2.46 A

MAGNETIC

SEPARATE

BLACK

POTTERY

2.88 A?

A

/

g gj

o m A ^ 2.81 A

f /

# /

2.97>|3-78J/

Quartz 3.34 A

Quartz 4.26 A

4? /

/ /

/ * „ c 3.76 A 3.55 A

k,

36 34 32

28

20

26 24 22 20 18

Figure 4. X-ray diffractometry spectrum of a magnetic separate from a sample of black Santa Clara Indian pottery. Even in the magnetic fraction, there is only a weak indication of magnetite and maghemite.

Low-Temperature I Low-Pressure Plasmas We have subjected pottery samples to repeated reductions and oxidations in low-temperature/ low-pressure oxygen and hydrogen plasmas in a reaction chamber similar to that shown, schematically in Figure 5. We produced an oxygen plasma for reaction of a black ceramic sample. In the conditions we used, the primary reactive species was atomic oxygen, present as approximately 10 to 20 percent of the particles in the reaction gas chamber. A majority of the particles remained as molecular oxygen, O2. Only a small percent of the oxygen species were ionized (for example, 02+,2_, 0+’~, and so forth), and the ceramic sample was far enough removed from the external plasma electrodes so that ionic recombination was expected to be virtually complete at the sample. The primary reaction of interest that we expected, therefore, was of atomic oxygen with iron oxides. After reaction with an oxygen plasma, the initially black pottery was rendered reddish-brown in color. A similar situation existed for the hydrogen plasma; there, of course, the expected reaction is the reduction of the hematite to the black oxides of magnetite/ maghemite or wiistite, or both. The initially red-brown pottery

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THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Figure 5. Schematic diagram of the low-temperature/ low-pressure oxygen plasma chamber. The gas could easily be switched to hydrogen and back to oxygen to yield reactive alternating oxidizing and reducing conditions within the chamber.

turned black in hydrogen plasma. Such samples could be repeatedly recycled from black to red and back.

One of the properties of an oxygen plasma, and one of interest here, is that it is quite reactive with virtually all varieties of carbonaceous materials. If the black coloration of these ceramics were due to carbon or even to the carbonaceous furfurans, as suggested by Rodgers (1979) for black smudges on pottery used over open fires, then it would be expected that the oxygen plasma would react with the carbonaceous matter to form CO2, CO and H20. However, the samples were recycled from black to red to black to red, and so on, by subjecting the sample first to an oxidizing oxygen plasma, then to a reducing hydrogen plasma, back to an oxygen plasma, and so on. Inasmuch as the black color reappeared each time upon reduction, it was not due to carbon or other carbonaceous matter that would have been converted to gaseous CO2, CO, and H20 during the course of this oxidation-reduction cycling. Rather it was more likely due to a metal oxide system as that of iron, possibly including other spinels such as hercynite, in which the changes between the various coloring agents (for example, red hematite contrasted with black magnetite, or maghemite, or wiistite, or some combination thereof) are reversible. We reaffirm, however, that if magnetite, or maghemite, or wiistite are really involved, they are present as particles smaller than the sizes that would respond to thermomagnetic analysis and x-ray diffraction.

COLOR OF AMERICAN INDIAN CERAMICS

65

Conclusions

Our work, like that before us, has not been totally successful in pointing to the precise chemical reactions most responsible for producing the black coloration in ferruginous clay ceramics, fired under reducing conditions, such as those used, for example, by modern American Indian potters. It is probable that the various reactions are complicated enough that, in fact, the precise reactions will depend critically on the exact extent of the reducing conditions to which the particular ceramic was exposed. We have, however, been able to rule out carbon and other carbonaceous material as the principal blackening agent in cases where the black coloration extends throughout the fabric of the ceramic material. Our evidence does not preclude that the iron and iron-oxide system may be responsible; if it is, however, the particle size of the black- reduced metallic iron or iron oxides must be less than 300 nm in diameter. Considering the present evidence, we think that this is likely the case. It appears to us that the definitive experiments are the low (liquid nitrogen) temperature Mossbauer spectroscopy experiments, which can detect and differentiate the iron-iron oxide species of interest even at extremely small particle sizes. Both of the Mossbauer studies on black and red surface paints on Chinese, Greek, and East Indian ceramics have in fact concluded that magnetite was important in causing black coloration and that the magnetite particles were quite small (Hedges, 1975; Longworth and Tite, 1979). In a third study, one involving Etruscan ceramics, the same conclusion was reached (Longworth and Warren, 1979). More germane to our studies here, however, was the Mossbauer study by Makundi et al. (1989), who found that the black coloration in the Cypriote and Nubian C-group ceramics, a product of the firing conditions, not paints, was due to the formation of ferrous ions in nonmagnetic compounds. Our thermomagnetic analyses are consistent with their Mossbauer results.

Note Added in Press

In a report of a Mossbauer study on red-black pottery from Early Bronze Age Cyprus just received from Waern-Sperber (1988), the same conclusions were reached as stated in our summary. She concluded: “To sum up, the results of the analyses indicate that the black colour of the EC II Red Polished black-topped bowl is not entirely due to carbon as suggested. Neither is it mainly due to manganese. Furthermore, positive evidence of magnetite in the black slip could not be obtained. The investigation indicates that the red part contains haematite and has been subjected to strongly oxidizing firing. The black part, though resembling haematite [in the Mossbauer spectrum] is different in that it also contains Fe2+ and has, it seems, been affected by more reducing conditions.”

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THE TEXAS JOURNAL OF SCIENCE— VOL. 42, NO. 1, 1990

Obviously, further investigations are necessary for a complete under¬ standing of the chemical cause for the black coloration in American Indian ceramics and in others worldwide.

Acknowledgments

This work was supported in part by the National Science Foundation Grant no. EAR- 8720837, the Robert A. Welch Foundation Grant A- 1062, and the American Chemical Society Grant ACS-PRF no. 20252-AC8. The support of the staff of the Texas A&M University Electron Microscopy Center is appreciated. We gratefully acknowledge partial support of this research by the Regents of Texas A&M University through the AUF- sponsored Materials Science and Engineering Program.

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Gillies, K. J. S., and D. S. Urch. 1983. Spectroscopic studies of iron and carbon in black surfaced wares. Archaeometry, 25:29-44.

Hedges, R. E. M. 1975. Mossbauer spectroscopy of Chinese glazed ceramics. Nature, 254:501-503.

Hegde, K. T. M. 1966. Electron microscopic study of the northern black polished (N.B.P.) ware of India. Curr. Sci., 24:623.

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Hewett, E. L. 1909a. The Excavations at Tyuonyi, New Mexico, in 1908. Amer. Anthropol., 11:434-455.

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Hofmann, U. 1962. The chemical basis of ancient Greek vase painting. Angew. Chem. Internat. Edit., 1:341-350.

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Hyman, M., and M. W. Rowe. 1983. Magnetite in Cl chondrites. J Geophys. Res., 88 (Suppl. Proc. 13th Lunar Planet. Sci. Conf., part 2):A736-A740.

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Kobayashi, K., and E. J. Schwartz. 1966. Magnetic properties of the contact zone between upper Triassic red beds and basalt in Connecticut. J. Geophys. Res., 71:5357-5364. Longworth, G., and M. S. Tite. 1979. Mossbauer studies on the nature of the red or black glazes on Greek and Indian painted ware. J. de Phys., 40(C2):460-461.

Longworth, G., and S. E. Warren. 1975. Mossbauer spectroscopy of Greek “Etruscan” pottery. Nature, 255:625-627.

Maggetti, M., and H. Schwab. 1982. Iron age fine pottery from Chatillion-S-Glane and the Heuneburg. Archaeometry, 24:21-36.

Makundi, I. N., A. Waern-Sperber, and T. Ericsson. 1989. A Mossbauer study of the 61 black colour in early Cypriote and Nubian C-group black-topped pottery. Archaeometry, 31:54-65.

Noll, W., R. Holm, and L. Born. 1975. Painting of ancient ceramics. Angew. Chem. Internat. Edit., 14:602-613.

Rahtz, P. A., and E. Greenfield. 1977. Excavations at Chew Valley Lake, Somerset. Dept. Envir. (Great Britain) Archaeol. Rept., 8: 1-392 (see p. 348).

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Rodgers, R. N. 1979. The chemistry of pottery smudging. Pottery Southwest, 6:2-4.

Shepard, A. O. 1976. Ceramics for the archaeologist. Publ. Carnegie Inst. Washington, 609:1-414.

Tobey, M. H., E. O. Nielson, and M. W. Rowe. 1986. Elemental analysis of Etruscan ceramics from Murlo, Italy. Pp. 115-127, in Proc. 24th Internat. Archaeom. Symp. (J. S. Olin and M. J. Blackman, eds.), Smithsonian Inst. Press, Washington, D.C., 517 pp. Waern-Sperber, A. 1988. Mossbauer spectroscopy and quantitative chemical analyses of early Cypriote blacked-topped pottery. Opuscula Atheniensia, 27:191-197.

OBSERVATIONS ON OBTAINING WHITE-TAILED DEER FAWNS FOR EXPERIMENTAL PURPOSES

Isaac M. Ortega, Lance D. Perry, D. Lynn Drawe, and Fred C. Bryant

Department of Range and Wildlife Management, Texas Tech University, Lubbock, Texas 79409 (IMO, FCB), Department of Zoology and Wildlife, Auburn University, Alabama, 36849 (LDP), and Welder Wildlife Foundation, P.O. Drawer 1400, Sinton, Texas 78387

(DLD)

Abstract. — In an attempt to obtain fawns for experimental purposes, 26 adult female white-tailed deer were captured using a drive net at the Welder Wildlife Refuge, San Patricio Co., Texas, on 12 March 1986. Deer were held in a 1.6-hectare enclosure until the end of the fawning season. Twenty-two does survived and at least 15 produced fawns. Thirty fawns (1 1 females, 19 males) were born alive between the dates 24 May and 23 June 1986. These results suggest that obtaining fawns from wild-trapped does held temporarily is feasible. Further, the stress of trapping, handling, close confinement, and daily disturbance apparently did not adversely affect birth rates. Key words: white-tailed deer; Odocoileus virginianus; Texas; captives; does; fawns.

Tame white-tailed deer ( Odocoileus virginianus) are used to study physiology, nutrition, growth rates, and behavior or food habits (Robbins and Moen, 1975; Bryant et al., 1981). Fawns generally are removed from their does shortly after birth to facilitate the taming process. Techniques used include capturing fawns in large enclosures where does are maintained throughout their lifetimes (Downing and McGinnes, 1969) or catching them in the wild by observing the behavior of wild does (White et al., 1972).

This paper describes the results of obtaining fawns from wild-trapped females held temporarily in an enclosure and then released back into the wild soon after birth of their young. The purpose of the project was to obtain fawns that were sufficiently tamed at one year of age to obtain data on food habits through direct observation.

Methods

Twenty-six adult female white-tailed deer were captured using a drive net (Beasom et al., 1980) at the Welder Wildlife Refuge, Sinton, Texas, on 12 March 1986. No attempt was made to determine if does were pregnant. Each animal was injected with one milliliter of acepromazine maleate (10 milligrams per milliliter) at the time of capture. Deer were transported two to nine kilometers by trailer to a 1.6-hectare enclosure within one to five hrs after capture. When the effect of the tranquilizer wore off, the deer often became excited and ran into enclosure fences, resulting in some minor injuries (lacerations and bruises). In mid-April, two semi-tame does were baited-in, captured and placed with the rest of the does.

The enclosure consisted of four interconnected pens (64 by 32 meters each, Fig. 1 A). By keeping the internal gates open, deer were allowed access to the entire 1.6-hectare enclosure. Outside fences were V-mesh net wire, 2.5 meters high. Inside fences were common net wire

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A

B

X

Brush

Brush

piles

Water

holes

Buffer

area

Figure 1. Pen used this study (A) and that suggested for similar studies (B).

(15 by 15 centimeter mesh). To minimize human interference and disturbance by free- ranging cattle, portions of the exterior fence were covered with black plastic sheeting to create visual obstruction. We also covered some internal fences with black plastic to provide a visual obstruction between the deer and people setting out feed or working in the pens. The holding pens were cleared of understory brush and mowed one week prior to capture of does to facilitate searching for fawns. A 30-meter line of brush was piled two-meters high in the fourth pen to provide screening cover.

Because deer are not fed supplements on the refuge, wild-trapped does had no experience eating pelleted feeds. Whole corn was offered ad libitum for 14 days but deer refused this feed. For the next eight days, two-centimeter pellets of cottonseed cake (35 percent crude protein) and 0.6-centimeter pelleted deer food were offered ad libitum. Deer did not feed regularly until alfalfa ( Medicago sativa) hay was offered on 3 April. Once deer began eating alfalfa hay, they also began eating the pelleted deer food. The captured deer refused to drink from a trough that was 0.5 meters high as evidenced by the lack of deer tracks at the trough. When deer tracks were observed at a small puddle formed by a leaking water hose, we buried a pan at ground level and allowed a water hose to drip into it. The watering hole was used the same day it was installed and every day thereafter.

Five does died during the first three weeks of captivity. A sixth doe escaped by jumping the 2.5-meter-high fence. The remaining 22 does survived and produced fawns. Some does showed a loss of weight during their captivity but had regained normal body condition at the time of their release back into the wild.

Daily searches for fawns were begun in mid-May. Newborn fawns were tagged with numbered aluminum poultry tags. Males were tagged in both ears and females in one ear. Male fawns were left with does, whereas female fawns were removed 48 to 72 hours after

OBTAINING WHITE-TAILED DEER FAWNS

71

birth. All were bottle-raised in concrete-floor cages until weaning following the guidelines of Kirkpatrick and Scanlon (1984). Female fawns were moved to a 0.5-hectare pen at 14 weeks of age, and fed alfalfa and horse-mule feed or deer pellets (16 percent crude protein) until one year of age.

Results and Recommendations

Thirty fawns (11 females, 19 males) were born alive in captivity. One set of twins was aborted early in May. However, at least 15 of 22 (68 percent) does gave birth to normal fawns.

Two of the 1 1 female fawns were born with hind leg abnormalities (abnormal tarsus). This may have been caused by physical damage to the fetus when handling wild-trapped does at capture. The fate of one abnormal fawn released with its dam at approximately one month of age is unknown. The second abnormal fawn was bottle raised but died at six months of age. At death, this fawn weighed the same as a fawn raised under confinement that was two to three months old.

Apparently confinement of the dams for three to four months prior to parturition did not affect birth weights of normal fawns. Fawn birth weight averaged 2472 grams and is comparable to the birth weights mentioned by Haugen and Davenport (1950) and Rue (1978).

These results suggest that obtaining fawns from wild-trapped does held temporarily is feasible. They also suggest that trapping and transporting deer in late gestation did not markedly affect birth rates.

The following are suggestions for maintaining captive, pregnant does.

1. The pen where does are enclosed should be at least 1.5-hectare in size and should include brush or other visual obstructions (1.5 meters high by 15 to 20 meters long) placed in at least one location to provide screening cover for the deer (see Fig. 1 B).

2. The enclosure should be located away from areas of human disturbance, and at least 100 meters from the nearest roadway.

3. A buffer area at least 50 meters wide should separate the pen from grazing livestock. The buffer area should contain enough brush or tall vegetation to minimize outside interference.

4. The pen should be fenced with V-mesh net wire at least 2.5 meters high. This wire should be buried 30 centimeters to protect the does and fawns against predators. The pen should be divided for separation of the deer.

5. The pen should be cleared of brush and tall grass before capture of does to facilitate location of fawns.

6. A water source should be installed at ground level.

7. The pen should contain tall trees to provide shelter against weather. The does in this study avoided man-made structures such as a three- meter high roof over feeding areas.

8. Although native foods are best, obtaining them may be labor intensive. If native forage is not provided, alfalfa hay should be provided

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as soon as the does are placed in the pens. Pelleted feed should be provided to insure a balanced diet.

Acknowledgments

We thank the many volunteer students from the wildlife classes at Texas A&I University, Kingsville, for helping capture does. We also thank J. Cox and B. Martinez from Welder Wildlife Refuge and B. Rust for their assistance. Funding was provided by the Rob and Bessie Welder Wildlife Foundation, P. O. Drawer 1400, Sinton, Texas 78387. This is a Welder Wildlife Foundation contribution no. 351 and Technical Article T-9-497 of the College of Agricultural Sciences, Texas Tech University.

Literature Cited

Beasom, S .L., W. Evans, L. Temple. 1980. The drive net for capturing western big game. J. Wildlife Manag., 44:478-480.

Bryant, F. C., C. A. Taylor, and L. B. Merrill. 1981. White-tailed deer diets from pastures in excellent and poor range condition. J. Range Manag., 34:193-200.

Downing, R. L., and B. S. McGinnes. 1969. Capturing and marking white-tailed deer fawns. J. Wildlife. Manag., 33:71 1-714.

Haugen, A. O., and L. S. Davenport. 1950. Breeding records of whitetail deer in the Upper Peninsula of Michigan. J. Wildlife Manag., 14:290-295.

Kirkpatrick, R. L., and P. F. Scanlon. 1984. Care of captive whitetails. Pp. 687-696, in White-tailed deer: ecology and management. (L. K. Halls, ed.), Stackpole Books, Harrisburg, Pennsylvania, xxiii+870 pp.

Robbins, C. T., and A. N. Moen. 1975. Milk consumption and weight gain of white-tailed deer. J. Wildlife Manag., 39:355-360.

Rue, L. L. 1978. The deer of North America. Crown Publishers, New York, xiii+463 pp. White, M. F., F. Knowlton, and W. C. Glazener. 1972. Dam-newborn fawn behavior on capture and mortality. J. Wildlife Manag., 36:897-906.

A NOTE ON SOME ASPECTS OF PITMAN NEARNESS

John W. Seaman, Jr, and Dean M. Young Department of Information Systems, Baylor University, Waco, Texas 76798-8005

Abstract. — Peddada (1985, 1986) and Berry (1986) have given sufficient conditions for an estimator that has smaller mean squared error or smaller mean absolute error than a competing estimator to be Pitman nearer. This note corrects a technical error and improves Peddada’s and Berry’s results through the use of the Cantelli-Frechet-Uspensky inequality. Finally, we note some variations on the definition of Pitman nearer that have appeared in the literature and the consequences of the differences in these definitions. Key words: mean square error; mean absolute error; Cantelli-Frechet-Uspensky inequality; Gauss inequalities; unimodality.

Peddada (1985) discussed the relationship among minimum mean square error, minimum absolute error, and Pitman nearness. For a given loss function L(T,0), he defined an estimator Ti to be closer to 0 than an estimator T2 in the Pitman nearer (PN) sense if P[L(Ti,0) < L(T2,0)] > 1/2. In his Theorem 2.2, Peddada provided sufficient conditions on the difference U = L(Ti,0) - L(T2,0) to imply that Ti is closer to 0 than T2 in the PN sense. These included the following moment conditions: E(U) = uQ < —2.67 and E(U - uG)j < j! for j = 1,2,. . . .

Berry (1986) used the Bienayme-Chebyshev inequality to prove that weaker conditions on the moments suffice in Peddada’s Theorem 2.2. Specifically, Berry’s conditions were that uG < -2.67 and a2 = Var(U) < 14(2. 67)2.

We point out a slight technical flaw in the arguments of both Berry and Peddada and provide a new, less restrictive, sufficient condition for the PN criterion to hold based on the Cantelli-Frechet-Uspensky inequality. We also discuss a potential further improvement via the assumption of unimodality and a Gauss-type inequality. Finally, we consider variations on the definition of PN that have appeared in the literature and some consequences of these variations.

A New Sufficient Condition for Pitman Nearness

Berry’s proof relied on the following implication: P( | U - uo | < 2 1/2 a) 22 1/2 = > P(U < uo + 2l//? a) > 1/2. This implication need not hold. For example, suppose U - uo has an absolutely continuous distribution function with support [— 2^2 a, + 00) and that P( | U - u0 | < 2^2 o) - 1/2. Then we must have P(U - Uo < 2'2 a) - 1/2 as well and the implication above does not hold. Of course, this problem can be alleviated by placing conditions on the distribution of U or by modifying the PN criterion by requiring only P(U < 0) > 1/2. This version of the PN criterion may be found, for example, in Mood et al. (1974), where L is taken to be

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absolute value loss. Indeed, in Peddada and Khattree (1986), the criterion is taken to be P(U < 0) > 1/2. We discuss variations of the definition of PN in a following section.

Peddada (1986) established an even weaker condition on the moments of U as he showed that the moments need only satisfy (uo/a) < — 2^2. However, again, we must either assume that P(U < -2I/2 a) > 0 or suitably modify the PN criterion.

We now establish a sufficient condition on the moments in Peddada’s Theorem 2.2, which are weaker than those presented by Berry and Peddada. We shall use the PN criterion in the sense of Mood et al. (1974) mentioned above, but under general loss. The condition follows from the Cantelli-Frechet-Uspensky (CFU) inequality (Frechet, 1950:137; Uspensky, 1937:198). For a random variable X with mean and variance /jl and t2, respectively, and a constant k, the CFU inequality is P(X - n > kr)< 1/ (1 + k2).

Theorem. — Let Ti and T2 be estimators of 6. Let L be a loss function, U = L(Ti,0) - L(T2,0), E(U) = uo, and Var(U) = a2. If (u0/a) < -1, then Ti is closer to 6 than T2 in the PN sense.

Proof: By the CFU inequality, P(U < Uo + o) > 1/2. If (uo/a) < — T, then P(U < 0) > 1/2 and the theorem is proved.

Note that the condition (uo/a) < — 1 is a considerable relaxation of the condition (u 0/0) < — 2'2, which was the best previous sufficient condition. Lee (1990) obtained a similar result.

A Sufficient Condition for Pitman Nearness Based on Unimodality

Berry’s improvement on Peddada’s sufficient condition was derived using Chebyshev’s inequality. Our improvement on Berry’s result has been obtained via a tighter probability inequality. We now consider what improvement may be had by assuming that U is unimodal, and employing a version of Gauss’s inequality.

Under the assumption of unimodality, Chebyshev’s inequality may be sharpened (though not uniformly). Probability inequalities that incorporate the assumption of unimodality are known as Gauss inequalities. We shall use the following Gauss inequality, which has been formulated by Vysochanskii and Petunin (1979). Let X be a unimodal random variable with mean n and variance r2. Then for all k > 0,

14 - k2 4

P( | X - n | > kr) < max < , —

1 3k2 9k

Now, let U be defined as before and suppose that it is unimodal. Set k

ASPECTS OF PITMAN NEARNESS

75

= da, for d > 0. In order that P(U < u0 + d0a) > 1/2, we must have do such that

Unfortunately, this inequality implies do > (8/5)^2 > 1. Thus, we obtain no improvement over the result obtained using the CFU inequality.

Variations on the Definition of Pitman Nearness

Let Ti and T2 be estimators of 6. Pitman’s original criterion, established in his 1937 paper, is that Ti is closer to 6 than T2 if P( | Ti - 0 | < | T2-0 |)> 1/2 for all 6.

Peddada (1985) and Rao et al. (1986), among others, used a generalized version of the PN criterion in which a general loss function is allowed. For a loss function L, the generalized PN criterion is that, for all 6, P[L(Tj,0) < L(T2,0)] > 1/2. This generalization affords the treatment of multiparameter estimation problems.

The generalized criterion may be further modified by changing one or both of the inequalities to allow equality. The four possible variations are listed below:

PN1: P[L(Ti,0)<L(T2,0)]> 1/2 PN2: P[L(Ti,0) < L(T2,0)] > 1/2 PN3: P[L(T,,0) < L(T2,0)] > 1/2 PN4: P[L(Ti,0)<L(T2,0)]> 1/2.

A cursory search of the literature reveals that PN1, PN3, and PN4 have been employed. In some cases, the same author used different versions in different papers. For example, Peddada and Khattree (1986) used PN3, whereas Khattree (1987) used PN4. Mood et al. (1974) used PN4 in their widely read text on mathematical statistics.

Our purpose in this section is to note that different results may be obtained using different versions of the PN criterion. We illustrate this fact with an example.

Let X be a unimodal random variable with support [a, b] and variance a2. Upper bounds on the variance of such random variables have been established by several authors; see Seaman and Odell (1988) for an overview. Seaman et al. (1987) considered the use of such bounds in small sample variance estimation. For example, when sampling from a distribution that is known to be symmetric unimodal, it can be shown that a2 < (b - a)2/ 12. The U-statistic for estimating variance in this case is the usual unbiased sample variance, S2. By truncating S2 at the upper bound, the mean square error (MSE) may be reduced for small (n < 10)

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sample sizes. If the criterion is minimum MSE, then the truncated estimator is superior. One may ask whether the truncated estimator, call it St, is Pitman nearer to o 2 than S2. The answer depends on which version of the PN criterion is employed.

Consider the following illustration. We have simulated 1000 samples of size 4 from a beta distribution with shape parameters three and three. This distribution is symmetric unimodal and therefore has variance not exceeding 1/12. We find that the ratio of the MSE for St to the MSE of S2 is approximately .910. However, using PN1 or PN4, we have P( | St - a2 1 < | S2 - a2|) « .047, so that S2 is PN to o 2 than St. By contrast, using PN2 or PN3, we have P( | St - cr2| < | S2 - a2|) = 1, so that one concludes that St is PN to o2 than S2. The two versions of the PN criterion, therefore, lead to opposite conclusions.

Literature Cited Berry, J. C. 1986. On Pitman nearness. Amer. Stat., 40:257.

Frechet, M. 1950. Generalites sur les probabilites. Variables aleatoires. Gauthier-Villars, Paris.

Khattree, R. 1987. On comparison of estimates of dispersion using generalized Pitman nearness criterion. Commun. Statist. -Theor. Meth., 16:263-274.

Lee, C. 1990. On the characterization of Pitman nearness. Stat. Prob. Letters, in press.

Mood, A. M., F. A. Graybill, and D. C. Boes. 1974. Introduction to the theory of statistics.

McGraw-Hill, New York (3rd ed.), xvi + 564 pp.

Peddada, S. D. 1985. A short note on Pitman’s measure of nearness. Amer. Stat., 39:298- 299.

— — . 1986. Reply to Berry’s letter to the editor. Amer. Stat., 40:257.

Peddada, S. D., and R. Khattree. 1986. On Pitman nearness and variance of estimators. Commun. Statist. -Theor. Meth., 15:3005-3017.

Pitman, E. J. C. 1937. The closest estimates of statistical parameters. Proc. Cambridge Phil. Soc., 33:212-222.

Rao, C. R., J. P. Keating, and R. L. Mason. 1986. The Pitman nearness criterion and its determination. Commun. Statist. -Theor. Meth., 15:3173-3191.

Seaman, J. W., and P. L. Odell. 1988. Variance upper bounds. Pp. 480-484, in Encyclopedia of statistical sciences (S. Kotz and N. L. Johnson, eds.), John Wiley and Sons, New York, 9:xxi + 1-762.

Seaman, J. W., P. L. Odell, and D. M. Young. 1987. Improving small sample variance estimators for bounded random variables. Industrial Math., 37:65-75.

Uspensky, J. V. 1937. Introduction to mathematical probability. McGraw-Hill, New York, ix + 41 1 pp.

Vysochanskii, D. F., and Yu. I. Petunin. 1979. On a Gauss inequality for unimodal distributions. Theor. Probab. Appl., 27:359-361.

PHYLLODONT (PARALBULINAE) FISH TOOTHPLATES FROM THE LOWER CRETACEOUS GLEN ROSE FORMATION OF CENTRAL TEXAS

Henry H. Huggins

Department of Physical Sciences, Tar let on State University, Stephenville, Texas 76402

Abstract. — Seven partial phyllodont toothplates from the Lower Cretaceous (Comanchean) Glen Rose Formation of Erath County, Texas, represent the largest collection of phyllodont toothplates known from a single locality in the state. Both parasphenoid and basibranchial toothplates, referrable to the Paralbulinae, are known from the site. The specimens compare favorably with Casierius heckelii. Key words: phyllodont; Casierius ; fish; Lower Cretaceous.

Collections of vertebrate fossils made by the author and by field crews from Southern Methodist University and Tarleton State University have been reported from fossil localities on the Huggins Ranch, 5 mi. (8.6 km.) NW Hico, in Erath Co., Texas. These localities have been referred to as the Huggins Localities or SMU nos. 108-1 and 108-2 (Winkler et al., 1990). Vertebrate fossils reported from the fossil localities include sharks, Hybodus sp., Pseudohyp olophus mcnultyi, Rhinobatos sp., and Leptostyrax bicuspidatus, and an amphibian, Prosiren elinorae, as well as other Caudata and Anura (Winkler et al., 1990). Most of these collections are from the locality designated as SMU no. 108-2. Seven toothplates (described here) were collected earlier from the site by the author and Chester Huggins.

Toothplates were found at the contact between the Glen Rose and the overlying Paluxy Formation or within a three-meter interval immediately above the contact. As the collection comes primarily from sediments turned up during the excavation of a stock pond, the precise stratigraphic horizon could not be determined. The Glen Rose Formation here is a dense, fossiliferous limestone that represents the Upper Member (Boone 1968; Perkins 1987). The Paluxy Formation immediately above the contact consists of poorly consolidated quartz sandstone.

Casierius has been reported from brackish-water, freshwater, and marine environments. The locality reported herein was an estuarine or transitional marine-terrestrial environment (Winkler et al., 1990), thus supporting the supposition that Casierius was diadromous.

General Description

Seven fossil toothplates were discovered at the site that show a phyllodont type of tooth replacement, with superposed sets of replacement teeth (Estes 1969a). All seven toothplates are believed to belong to the phyllodont subfamily Paralbulinae, although there are

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minor differences in the morphologies of the plates. Three of the toothplates represent major portions of toothplates, the other four are fragmentary.

The teeth are irregularly stacked and are subcircular to oblong in occlusal view. In lateral view, they appear flattened or shallowly cupped, with the concave surface toward the attachment surface. There is no evidence of sculpturing on the surfaces of the teeth, although some show wear due to trituration. Wear on these teeth probably is due to predation on molluscs and crustaceans by these fishes (Estes 1 969a). The teeth vary in size from 0.7 to 2.1 mm. The larger teeth are typically in the center of the plate and the smaller teeth toward the edges. In lateral view, the toothplates taper anteriorly. No sigmoid curvature of the occlusal surface of the toothplates is present, as seen from both occlusal and lateral views, thus differentiating them from members of the Phyllodontinae.

Of the seven toothplates, six have a convex occlusal surface whereas the seventh is concave. The concave plate (SMU 72340; Figs. 1A, IB, and 1C) differs from the other toothplates in many respects. In particular, it has six layers of irregularly stacked teeth, whereas the other specimens have only three or four. Occlusal view of the specimen reveals moderate tooth wear. The lateral edges of the plate are raised, with parallel grooves perpendicular to the occlusal surface, which may represent the symphyseal attachment surface, as seen in Figure IB. On the dorsal portion of the plate, a pair of raised facets may be for articulation with other bones of the skull. The bulbous dorsal attachment surfaces, raised lateral edges, and concave occlusal surface indicate that this is a parasphenoid toothplate (Estes 1 969a). The concave occlusal surface would have fit around a smaller basibranchial plate in the frontal mandibular area. Thurmond (1974) referred to a fragmentary concave phyllodont plate from the Garvin Church local fauna in the lower Paluxy Formation of Wise County, Texas, as a pterygoid plate from Casierius heckelii. Although one of the characters used to define the Phyllodontidae is the presumed lack of pterygoids (Estes 1969 b), this is only a superficial definition and cannot be supported due to a lack of other cranial and skeletal bones.

Convex basibranchial tooth patches, which lack sigmoid curvature of- the occlusal surface, are also characteristic of the more primitive genera of the subfamily Paralbulinae (Estes 1969a). The other six specimens (SMU 772341, 72342, 72343, 72344, 72345, 72346) are consistent with this morphology and are, therefore, also believed to be referrable to the Paralbulinae. These plates are similar in regard to tooth size, shape, the irregular stacking of the replacement teeth, and the mode and extent of tooth wear. The specimens have convex occlusal surfaces and, where present, flattened ventral attachment areas.

FISH TOOTHPLATES FROM CRETACEOUS OF TEXAS

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Figure 1. A. Casierius heckelii (anterior to left), occlusal view of fragmentary parasphenoid, showing trituration of tooth surfaces (SMU 72340).

B. C. heckelii, (anterior to left), right lateral view of SMU 72340 showing parallel grooves of lateral edges, presumed symphyseal attachment surface.

C. C. heckelii, medial view of broken anterior end, showing multiple sets of replacement teeth and concave structure (SMU 72340).

D. SMU 72341, medial view of broken anterior end of basibranchial toothplate showing high-domed occlusal surface, C. heckelii.

E. SMU 72342 (anterior to left), lateral view of basibranchial toothplate showing flattened occlusal surface and smaller teeth toward edge, C. heckelii.

F. SMU 72342, medial view of broken anterior end.

Specimen SMU 72341 represents the posterior portion of a toothplate. It has a high-domed dorsal surface (see Fig. ID). The posterior edge of the specimen is three millimeters thick, and the central portion of the toothplate thickens to seven millimeters. The larger teeth are found in the middle two-thirds of the plate.

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Specimen SMU 72342 has a much flatter occlusal surface than in specimen SMU 72341 (see Figs. IE, IF). Also, teeth are smaller in comparison (1.0 to 2.1 mm in 72341 as opposed to 0.7 to 1.8 mm in 72342). Both SMU 72341 and 72342 have approximately four layers of teeth. Inasmuch as the toothplates are approximately the same size, it cannot be determined whether these represent different taxa, different toothplate series, or differences in preservation. However, the ventral surface of SMU 72342 is flattened, whereas the ventral surface of SMU 72341 is slightly concave, also revealing significant differences between these plates of similar size.

Specimens SMU 72343, 72344, 72345, and 72346 consist of poorly preserved fragments with convex occlusal surfaces. However, tooth morphology and replacement indicates placement within the paralbulines.

Discussion

The extinct phyllodont fishes may be divided into the subfamilies Phyllodontinae and Paralbulinae. There are clear distinctions between the two subfamilies. The Phyllodontinae are characterized by toothplates with sigmoid curvature and in having multiple sets of teeth that are directly successional in stacks on the main portion of the plate. Conversely, the Paralbulinae are phyllodont fishes with basibranchial and parasphenoid plates that lack a sigmoid curvature. Other characteristics of the subfamily include the presence of abundant interstitial matrix between the teeth, which is composed of a calci-phosphatic replacement material. The assumed presence of interstitial matrix is due to the

irregular stacking of the toothplates in the Paralbulinae (Estes 1969a).

All specimens described herein possess characteristics of the Paralbulinae. Of the genera described from the subfamily, the SMU specimens most closely resemble those described as Casierius sp. A single basibranchial plate of C. heckelii has been described from the Glen Rose Formation of northern Bosque County, Texas (Johnson, 1972). The curvature of the toothplate and the irregularity of tooth stacking is

identical between Johnson’s plate and those reported here. In addition,

tooth shape, size, and basic morphology are comparable. The Huggins toothplates differ in size from the specimen described by Johnson, although this is not necessarily a constraining factor. SMU 72340

represents the first parasphenoid plate described and figured from the Lower Cretaceous of Texas.

Acknowledgments

I would like to thank Chester Huggins for assistance in collection of the fish toothplates, and Michael Cope for drawing the illustrations. I am also grateful to Dr. Phillip A. Murry (Tarleton State University) for his comments on the manuscript and Dr. Dale Winkler of Southern Methodist University for supplying comparative material. Funding was provided

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by National Geographic Society Grant 3403-86 and National Science Foundation Grant

BSR-8700539 to Drs. Dale Winkler and Phillip Murry, and Tarleton State University

Organized Research Grant 15-639 to Dr. Murry.

Literature Cited

Boone, P. A. 1968. Stratigraphy of the basal Trinity (Lower Cretaceous) sands of central Texas. Bull. Baylor Geol. Stud., 15:1-62.

Estes, R. 1969a. Studies on fossil phyllodont fishes: interrelationships and evolution in the Phyllodontidae (Albuloidei). Copeia, 1969:317-331.

- . 19696. Studies on fossil phyllodont fishes: Casierius, a new genus of albuilid from

the Cretaceous of Europe and North America. Eclogae geol. Helvetiae, 62:751-755.

Johnson, G. D. 1972. Phyllodont toothplate from the Lower Cretaceous of Texas. Texas J. Sci„ 24:67-74.

Perkins, B. F. 1987. Alternating marine and terrestrial environments in the Comanche Cretaceous lower Glenrose Formation, Somervell and Hood counties, Texas. Field Trip Guide, 1987 Southwest Sect. Amer. Assoc. Petrol. Geol. Convention, Field Trip 1, pp. 9-37.

Thurmond, J. T. 1974. Lower vertebrate faunas of the Trinity Division in north-central Texas. Geoscience and Man, 8:1 19-120.

Winkler, D. A., P. A. Murry, and L. L. Jacobs. 1990. Lower Cretaceous (Comanchean) fossil vertebrates of central Texas. J. Vert. Paleo., in press.

IMPORTANCE OF CANOPY POSITION FOR GROWTH OF CELTIS LAEVIGATA SEEDFINGS

O. W. Van Auken and R. J. Fohstroh Division of Life Sciences, The University of Texas at San Antonio, San Antonio, Texas 78285

Abstract. — Seedlings of Celtis laevigata (Willd.), Texas sugarberry, were planted in the field to determine affects of canopy position, root competition, nutrient addition and herbivory on their growth. Growth was greatest under a mature Acacia smallii (Isley), huisache, canopy and least in open grassland between the mature Acacia trees. Mortality was highest in the open grassland compared to below the Acacia canopy. Other factors examined had no significant effect on C. laevigata seedling growth. Celtis laevigata is a mature community species that is shade tolerant and requires high levels of soil nitrogen. As such, growth should be limited in areas typical of early successional communities and promoted by conditions typical of mature communities. Key words: competition; herbivory; light levels; nutrients; shade; Texas sugarberry; trenching.

Secondary succssion in parts of southern Texas may begin with abandonment of farmland. If farming is stopped, colonization by various annual species occurs quickly, followed by establishment of woody plants like Acacia smallii (Isely), huisache, within five years (Van Auken and Bush, 1985). Acacia smallii dominates the savanna-woodland for the next 25 to 35 years, after which it declines. It is shade intolerant (Bush and Van Auken, 1 986a) and grows poorly below its own canopy (Fohstroh and Van Auken, 1987). During this time period, Celtis laevigata (Willd.), Texas sugarberry, becomes established and then dominates the mature community (Bush and Van Auken, 1984). Celtis laevigata is shade tolerant as might be expected for mature community species (Bush and Van Auken, 1986c).

Although C. laevigata is tolerant of low light levels in the greenhouse, we have not identified any reports of its ability to grow in the field or its site requirements for establishment. Fate successional species are usually tolerant of low levels of light (Grime, 1965; Loach, 1967, 1970), but may require higher levels of nutrients (Bormann, 1953; Tilman, 1982; Van Auken et al., 1985). Additional factors that could be important in determining plant establishment and dominance are herbivory and ability to compete for resources below ground (Weaver and Clements, 1966; Harper, 1977; Smith, 1980).

This study was designed to determine the importance of canopy position, nutrient addition, herbivory, and root competition on the growth of seedlings of C. laevigata in the field.

Methods

Fruits of Celtis laevigata were collected in February 1985 from trees located along the San Antonio River, Bexar County, Texas. Seeds were soaked in giberellic acid for 3.5 hours

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to break dormancy (Kahn, 1968) and then rinsed thoroughly with distilled water. Three seeds were sown in each 15-centimeter diameter by 15-centimeter deep plastic pot containing 1400-gram sieved, Frio clay-loam soil (Taylor et al., 1966) from an early successional site. The soil was a Mollisol, classified as a fine, mixed, thermic, Cumulic Haplustoll, low in carbon (1.29 + 0.45 percent, by weight), nitrogen (0.14 + 0.03 percent), and phosphorous (8 ± 2 milligrams per kilogram). Calcium was high (2400 + 800 milligrams per kilogram) and magnesium and potassium were at intermediate levels (30 + 1 1 milligrams per kilogram, 1 1 + 5 milligrams per kilogram, respectively) (Bush and Van Auken, 19866). Rainfall data were taken from the NOAA local climatological data summary (NOAA, 1985).

Seven weeks after sowing, plants were sized according to height and vigor and thinned to one per pot. One-hundred-nineteen plants were used in the experiment. Seven randomly selected plants were kept in the greenhouse and the remaining plants were transplanted to a field site along the San Antonio River. Seven randomly selected plants were placed into each of the 16 treatments. All plants were given 300 milliliters of distilled water every third day for 10 days (equilibration) before initial measurements. Greenhouse plants were harvested for initial measurements by cutting at the cotyledon scars and drying (70° C) to a constant weight.

The experimental design was 24 factorial with canopy position (under an Acacia smallii canopy or in the open), nutrients (added or not added), root competition (roots present or trenching, no woody plant roots), and herbivory (herbivores present or insecticide treatment, no herbivores) as factors. With all combinations, there were 16 treatments. The canopy position was beneath five mature A. smallii trees and the open area was adjacent. Mean light levels were 515 ± 153 /xM-rrf^sec1 under the canopy and 2173 ± 174 juM-m”2- sec 1 in the open. Light levels were measured with a Li-cor® LI- 188 integrating quantum sensor. Nutrient treatment consisted of 250 milliliters of a complete nutrient solution (Van Auken and Kapley, 1979) once per week starting at zero time for 12 weeks. No nutrient- treatment plants were given 250 milliliters of distilled water at the same time intervals. Root competition was reduced (no competition treatment) by trenching a one square meter perimeter around the plants, to a depth of 20 centimeters. Root competition treatments were untrenched. No herbivory plants were sprayed with malathion (2.95 milliliters per liter) once each week for the duration of the experiment. A four-sided cardboard box prevented the insecticide from being blown onto other test plants. Herbivory treatment plants were sprayed with an equal amount of distilled water at the same time intervals.

The experiment was terminated 12 weeks after initial measurements were taken (1 November 1985). Plants were cut at the cotyledon scars and dry weights were measured. Data were statistically analyzed by an analysis of variance (ANOVA) procedure and means were separated using the Least Significant Difference test. Mortality data was analyzed with a X2 test (Steel and Torrie, 1980; SAS Institute, 1982).

Results

Position was the only main factor tested that was significant (ANOVA, F - 64.03, P < 0.0001, Fig. 1). Herbivory, competition, or nutrient additions were not significant (ANOVA, F - 0.00-3.59, P > 0.05). Of the 10 two-way and three-way interactions, only the herbivory-nutrient interaction was significant (ANOVA, F - 4.37, P - 0.0397). Mean aboveground dry weight of Celtis laevigata seedlings grown in the open (grassland, full sunlight) was 0.25 ± 0.21 grams, whereas mean aboveground dry weight of plants grown below the Acacia canopy (shade) was 0.96 ± 0.45 grams. Mean aboveground dry weight of the plants harvested at the start of the experiment was 0.14 ± 0.08 grams.

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Figure I. Bar graph presenting mean aboveground dry weight (+ SD) of Celtis laevigata seedlings grown for 12 weeks under various field conditions. Treatments included the following: 1, under canopy; 2, open; 3, herbivory; 4, no herbivory; 5, competition; 6, no competition; 7, nutrient supplemented; 8, no nutrients added. Mean Dry weight of plants in treatment 1 was significantly different from those in treatment 2 (ANOVA, LSD, P < 0.05). There were no significant differences between the other treatments.

Thus, plants in the open (grassland) increased in dry weight 1.79 times, while those in the shade below the Acacia canopy increased 6.86 times.

Means of all treatments were separated using the least significant difference test (Table 1). The main differences were in canopy position. Mean values for all plants in treatments in the open were lower than means for plants in treatments below the Acacia canopy. Other differences are slight and should not be considered because other main effects were not significant when tested with ANOVA.

Overall seedling mortality in the field was 36 percent, and 98 percent of these mortalities occured in the open (grassland, full sunlight), which was highly significant (X2 - 36.1, P < 0.005). Rainfall during August 1985 was only 0.45", which was 83 percent below normal, and this was the time when almost all of the Celtis mortalities occured (Fig. 2). Rainfall during September and October was 28 percent and 36 percent above normal, and only one mortality occurred during this time.

Discussion

Many abiotic and biotic factors affect the establishment and growth of woody plants (Harper, 1977). However, the present field study identified

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Table 1. Mean aboveground dry weight (± SD) for Celtis laevigata grown in 16 treatments and initial measurements. Mean values followed by the same letter are not significantly different (ANOVA, LSD P> 0.05).

Treatments	Dry-weight (grams)
Zero time	0.14	±	0.08A
Open, herbivory, competition, nutrients	0.18	+	0.31 A
Open, herbivory, competition, no nutrients	0.08	±	0.03A
Open, herbivory, no competition, nutrients	0.23	+	0.28AB
Open, herbivory, no competition, no nutrients	0.35	+	0.36ABC
Open, no herbivory, competition, nutrients	0.23	+	0.42AB
Open, no herbivory, competition, no nutrients	0.17	±	0.19A
Open, no herbivory, no competition, nutrients	0.40	+	0.35ABC
Open, no herbivory, no competition, no nutrients	0.28	±	0.45AB
Under canopy, herbivory, competition, nutrients	1.00	±	0.55DE
Under canopy, herbivory, competition, no nutrients	0.91	+	0.29CDE
Under canopy, herbivory, no competition, nutrients	0.85	+	0.36CDE
Under canopy, herbivory, no competition, no nutrients	0.93	+	0.19CDE
Under canopy, no herbivory, competition, nutrients	0.88	±	0.49CDE
Under canopy, no herbivory, competition, no nutrients	0.79	+	0.41BCD
Under canopy, no herbivory, no competition, nutrients	1.41	+	0.93E
Under canopy, no herbivory, no competition, no nutrients	0.90	±	0.42CDE

C. laevigata seedling position relative to an Acacia smallii canopy as the major factor. Growth of C. laevigata seedlings below the Acacia canopy was 3.84 times higher than that of seedlings in open grassland. Reciproically, mortality of C. laevigata seedlings in the open grassland was 39 times higher compared to those below the Acacia canopy.

Light intensity, an obvious variable influenced by the presence of a woody plant canopy, may be reduced by more than 90 percent by the A. smallii overstory (Bush and Van Auken, 1986a). But, low light levels should reduce, not stimulate, C. laevigata growth. Apparently, the stimulation of C. laevigata seedling growth is mediated by another factor associated with the A. smallii canopy. Soil temperature, soil nutrients, and soil water content all have been shown to change below the canopies of certain woody species (Tiedemann and Klemmedson, 1973a, 19736, 1986; Bush and Van Auken, 19866). Higher levels of soil nutrients may be a factor involved in stimulating the growth of C. laevigata. However, we did not find a nutrient stimulation in the present experiment. Soil nutrients, especially nitrogen, have been shown to increase during successional events (Gorham et al., 1979), and total soil nitrogen has been demonstrated to increase during secondary succession in this area (to 2.5 + 0.7 g*kg_1, Bush and Van Auken, 1986b). In addition, total soil nitrogen is almost twice as high under a 15-year-old Acacia smallii canopy compared to open grassland. Low light levels occur below the canopy, but higher soil nitrogen levels also are found. Added nitrogen,

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Figure 2. Bar graph presenting weekly rainfall and weekly mortality of Celt is laevigata seedlings during the experiment.

rather than other nutrients, has been shown to stimulate C. laevigata growth in the same Frio soil in greenhouse experiments (Van Auken et ah, 1985). Thus, soil nitrogen seems to be a major candidate for causing the increased growth observed.

Other factors examined, including competition, nutrient supplements, and herbivory, did not cause significant changes in C. laevigata growth. Competition for soil-borne resources, mainly water and nutrients, can play a significant role in establishment and subsequent growth of woody species (Harper, 1977). Trenching effectively eliminates root competition (Ehrenfeld, 1980; Horn, 1985). But, we did not see a difference in C. laevigata growth between trenched and nontrenched plots and concluded that competition between adult and seedling plants was unimportant. Competition with woody plant roots may have been masked by competition with herbaceous plants that were not removed from the study plots (both canopy and open plots). Competition between woody plant seedlings and herbaceous plants can cause significant growth

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reduction in woody plants (Glendening and Paulsen, 1955; Van Auken and Bush, 1989).

Nutrient additions were expected to have stimulatory effects on the growth of C. laevigata as seen in previous studies (Van Auken et al., 1985). However, additional nitrogen in the canopy soil could have masked that effect. The lack of stimulation of growth of C. laevigata in the open when nutrients were added was probably the result of the early drought and later stimulation of associated herbaceous species by the removal of a nutrient limitation. With the growth limitation removed, the herbaceous species may have used up a nutrient required for C. laevigata growth.

Herbivores can have drastic effects on plant populations and change successional sequences (Harper, 1977; Gilbert, 1985), although not shown in the present study. This study was completed in the autumn and was of limited duration. A study completed over a longer time and including both the spring and autumn growing seasons might show significant growth reduction due to herbivory, if herbivores are present.

Celtis laevigata appears to be a late successional species, requiring high soil nitrogen and capable of growth in canopy shade. Although it can grow in disturbed areas or grasslands, it would do so at reduced rates, depending on the presence of competition and soil nitrogen levels. A more favored site would be in the reduced light and higher soil nitrogen environment below the canopy of A. smallii.

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Bormann, F. H. 1953. Factors determining the role of loblolly pine and sweetgum in early oldfield succession in the Piedmont of North Carolina. Ecol. Monogr., 23:339-358.

Bush, J. K., and O. W. Van Auken. 1984. Woody species composition of the upper San Antonio River gallery forest. Texas J. Sci., 36:139-145.

- . 1986a. Light requirements of Acacia smallii and Celtis laevigata in relation to

secondary succession on floodplains of South Texas. Amer. Midland Nat., 1 15:1 18-122.

- . 19866. Changes in nitrogen, carbon, and other surface soil properties during

secondary succession. Soil Sci. Soc. Amer. J., 50:1597-1601.

Ehrenfeld, J. G. 1980. Understory response to canopy gaps of varying size in a mature oak forest. Bull. Torrey Bot. Club, 107:29-41.

Gilbert, L. E. 1985. Ecological factors which influence migratory behavior in two butterflies of the semi-arid shrublands of south Texas. Pp. 724-747, in Migration: mechanisms and adaptive significance (M. A. Rankin, ed.), Contrib. Marine Sci., 27 (suppl.): 1-868. Glendening, G. E., and H. A. Paulsen, Jr. 1955. Reproduction and establishment of velvet mesquite as related to invasion of semidesert grasslands. USDA Tech. Bull., 1 127:1-50. Gorham, E., P. M. Vitousek, and W. A. Reiners. 1979. The regulation of chemical budgets over the course of terrestrial ecosystem succession. Ann. Rev. Ecol. Syst., 10:53-84.

Grime, J. P. 1965. Shade tolerance in flowering plants. Nature, 208:161-163.

Harper, J. L. 1977. The population biology of plants. Academic Press, London, 892 pp.

Horn, J. C. 1985. Responses of understory tree seedlings to trenching. Amer. Midland Nat., 114:252-258.

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Kahn, A. A. 1968. Inhibition of gibberellic acid-induced germination by abscisic acid and reversed by cytokinins. Plant Physiol., 43:1463-1465.

Loach, K. 1967. Shade tolerance in tree seedlings. I. Leaf photosynthesis and respiration in plants raised under artificial shade. New Phytoh, 66:607-621.

- . 1970. Shade tolerance in tree seedlings. II. Growth analysis of plants raised under

artifical shade. New Phytol., 69:273-286.

Lohstroh, R. J., and O. W. Van Auken. 1987. Comparison of canopy position and other factors on seedling growth in Acacia smallii. Texas J. Sci., 39:233-239.

NOAA. 1985. Local climatological data, annual summary with comparative data, San Antonio, Texas. National Climatic Data Center, Asheville, North Carolina, 8 pp.

SAS Institute. 1982. SAS user’s guide. SAS Institute Inc., Cary, North Carolina, 584 pp.

Smith, R. L. 1980. Ecology and field biology. Harper and Row, New York, 835 pp.

Steel, R. G. D., and J. H. Torrie. 1980. Principles and procedures of statistics: a biometric approach. McGraw-Hill, New York, 633 pp.

Taylor, F. B., R. B. Hailey, and D. L. Richmond. 1966. Soil Survey of Bexar County, Texas. USDA Soil Conserv. Serv., Washington, D.C., 126 pp.

Tiedemann, A. R., and J. O. Klemmedson. 1973a. Nutrient availability in direct grassland soil under mesquite ( Prosopis glandulosa ) trees and adjacent open areas. Soil Sci. Soc. Amer. Proc., 37: 107-1 1 1 .

— — . 19736. Effects of mesquite on physical and chemical properties of the soil. J. Range Manag., 26:27-29.

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bulk density. Soil Sci. Soc. Amer. J., 50:472-475.

Tilman, D. 1982. Resources, competition and community structure. Princeton Univ. Press, Princeton, New Jersey, 296 pp.

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Van Auken, O. W., E. M. Gese, and K. Connors. 1985. Fertilization response of early and late successional species: Acacia smallii and Celtis laevigata. Bot. Gaz., 146:564-569.

Van Auken, O. W., and J. K. Bush. 1985. Secondary succession on terraces of the San Antonio River in south Texas. Bull. Torrey Bot. Club., 1 12:158-166.

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SOLAR POND FEASIBILITY FOR LOW COST ENERGY AND WATER PRODUCTION

M. A. K. Lodhi

Department of Physics, Texas Tech Univesity, Lubbock, Texas 79409

Abstract. — The High Plains region in Texas and New Mexico seems to be a favorite site for salt-gradient solar ponds due to its potential for overcoming the major drawbacks in cost effectiveness of these ponds. Strong points for this region are the availability of a 1) naturally level ground with pre-existing large alkaline lake basins, 2) high insolation almost year round at a height of one kilometer above sea level, 3) salt from alkaline lake basins and oil-brine emulsion from oil fields, 4) soils with good impervious and insulating properties, and 5) demand of low to moderate temperature heat and fresh water in the area. The feasibility of production of electricity from low-temperature heat also is discussed. Key words: solar pond; energy cost; High Plains; energy sources.

In a recent paper published in a previous issue of Texas Journal of Science, the principle and potential of salt-gradient solar-pond (SGSP) were discussed with reference to the High Plains region of western <