Appendix 2 ‘A DESCRIPTION AND COMPARISON OF FIELD METHODS USED IN STUDYING AND CENSUSING DESERT TORTOISES Kristin H. Berry Because field methods affect results, an understanding of procedures influences both analysis and interpretation of data. This appendix presents a detailed description of the methods used in collection of data from all existing tortoise study plots for which ve have inforzation, Seven important factors to consider in examining the field methods of a tortoise study are: (1) time of year or season the study was conducted, (2) the season(s) tortoises were active (3) number of days spent by each investigator in the field, (4) the time of day searches vere mide, (5) the qualifications of the investigator(s), (6) the number of investigators, and (7) the study plot size. Field survey methods used in studies by the Bureau of Land Management (BLN and other investigators at the forty-seven plots can be grouped into six principal categories: (1) the 30-day spring census, (2) the 60-day spring census, (3) the 20-day fall census, (4) the three- or four- census or year-long survey, (5) winter den surveys, and (6) special surveys. The first three categories were the most common survey methods and were developed by the author for BIM. Some study plots received two or three types of surveys, others only one type. A summary of surveys undertaken at each plot is presented in Table 1 with notes on any modifications in the basic census or study techniques. 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‘eats 201d res pue Buyzds Apna Teyoeds Apnas SuoT-209k Apnas Suot-1e9k kpnae Buot-a99k 108-6L6T 208-L6T 208-LU6T 208-LL6T dha pur 19k ensue (1961 yaesng) “3TTe *a03930g (aorrespunmeos Teuosred ‘pavddous tog6t axemyo pur upmyor) Ksoydyaed - 11 pur 1 ‘zyay ‘edots meq 22080g (woyaeoyunmmo> yeuosaed ‘pieddays fog6T axemo Pur teRyor) 11 *2¥2y ‘dots weq 19A82q (vor eoyunso> Teuosxad preddaus 0961 23emo pur uemioH) 1 *Ay3y ‘edoTs weg 10K0og ‘souexayea pus omeu aa7s, panuyauog “Tt eTqey ‘ eats 42-7is discussed. THE 30- AND 60-DAY SPRING CENSUS TECHNIQUES Factors Considered in Developing Census Techniques Descriptions of the 30- and 60-day spring census techniques have not been published elsevhere and are worthy of attention here, Sone background on the developaent of the techniques is appropriate also, From 1973 co 1979, the BLM, California Desert Program, conducted a massive inventory of resources vithin the California deserts. The purpose of the inventory vas to gather data on all resources for a major land-use plan for public lands. Collection of data on the distribution and status of selected animal species such as the desert tortoise was part of the program. The author considered five factors in developing the 30-day and 60-day spring census techniques: funding, permanence of sites, minim size of sites, site location, and duration and timing of study. Each factor is discussed briefly belov. Funding. A series of short-tern, low-cost studies were a prerequisite because of Limited funding and its incremental nature. The BLM received an annual budget for the California Desert Program inventories, However, the snount of funding was uncertain until approximately January of each year in which the studies were to be conducted, Field personnel had to be hired and the field work initiated with a fev-weeks notice. Permanence of Sit The sites were selected by BLM personnel and were chosen where permanence of the habitat seemed assured. The sites were on lands scheduled for retention as public land and where no major 2-8surface-disturbing activities were anticipated in the foreseeable future. The single exception (site 9) on private property was an error in selection. Minimum Size of Plots. We selected the one sq.-mile as the minimum size racher than the smaller 62- to 247-acre (25- to 100-ha) quadrats described by Bury and Luckenbach (1977) for several reasons. First, the plots were to be used for determining not only demographic features but population condition and trend over long time periods. The sample size of tortoises had to be adequate for such determinations. The considerable differences in population densities in the geographic range vere already a matter of record (Berry 1974, 1975; Burge 1977; Ron Marlow, personal communication; Woodbury and Hardy 1948), Population densities ranged from approximately 10 to over 200 tortoises per sq. mile. Most sites vere chosen in areas expected to have more than 50 tortoises per sq. mile. When densities were suspected of being lower, the sq. mile study area could be enlarged. Second, we wanted to determine the relationships of tortoise distribution and abundance to various types of soil, vegetation, and topography. We thought that such information could be obtained more easily through large study plots, which were likely to contain subtle differences in topography, soils, and vegetation. Location of Plots. Four factors were considered in selecting locations: (1) adequate tortoise densities for sampling, (2) habitat typically occupied by tortoises, (3) lack of human disturbance, and (4) data from strip-transect studies on relative densities and distribution of tortois First, in order to save time and funds, sites vere selected where A2-9viable tortoise populations were known to be or where soil, vegetation, and elevation were expected to support viable populations, No plots were placed at sites where tortofses were rarely observed, such as at elevations above 3,500 ft or in such vegetation types as pinyon-juniper woodland. Sone plots were chosen on the basis of coments from BLM staff or the public, such as, “there are lots of tortoises near Chemehuevi Wash" or, "we used to see tortoises piled up against the railroad tracks near Coffs." cond, the plots had to have vegetation and soil types typical of Large expanses of lowland desert and repr satative of a major ecosystem (western Mojave Desert, eastern Mojave Desert, Colorado Desert, etc.). Between 1976 and 1981, an almost equal number of plots were chosen in the Mojave and Sonoran (Colorado) deserts of California. We hoped to extrapolate the findings from a single site to much larger, surrounding areas. ‘Third, the plots chosen were in areas with a minimum of recent or current human disturbance. The selected sites were considered the least disturbed examples of the representative habitat type of ecosystem. With two exceptions, all plots were located at least 0.5 mile from paved roads. It is important to note that the entire lowland de rt where tortot: cccur has had some human impacts, hovever. Fourth, some plots vere selected on the basis of strip-transect data acquired from the California deserts between 1976 and 1978 (see Chapter 2). Sone sites vere chosen in areas with little or no tortoise sign, while others were picked here sign levels were moderate to high. The purpose in such selection was to relate the strip-transect data to actual tortoise densities and distribution at specific sites. 2-10Duration and Timing of the Study. The 30- and 60-day spring censuses were centered around the peak, above-ground activity period for tortoises. Census periods generally involved 30 to 60 calendar days to allow for variations in tortoise activicy, The length of the 30-day census permitted capture of over 120 tortoises, a number initially considered adequate for a population sample. The 60-day census pernitted capture of even more individuals. The intent of the field work was to find as many live tortoises and carcasses as possible on each plot. Each fieldworker walked the terrain for about eight hours per day during key tortoise activity periods, recording tortoise observations. The fieldvorkers vere encouraged to avoid "tortoise watching" and to resume search immediately after recording data pertinent to captured individuals, We hoped such an effort would provide population samples where problems associated with immigration and emigration, hatchlings, and mortalities would be minimized. ‘The 30-day Spring Census The 30-day census technique was developed gradually over a three-year period, and refined sonevhat over the years. Thirty-day censuses vere undertaken on 19 plots between 1977 and 1980 (Table 1). Variations in field methods are discussed below by year. In 1976, Berry began experizenting with an accurate and inexpensive method for gathering quantitative data on tortoise density, population structure, sex ratios, mortality, and tortoise distribution within various elected sites in California. A on habitat types at qe-mile plot was established in Fremont Valley (Site 2), and a single investigator spent 18 A211days in May attempting to capture and mark every live tortoise on the plot and collect all carcasses. In that time period, 118 individuals vere marked and 43 carcasses collected, Because this technique appeared successful in obtaining a large sample of tortoises, we adopted the 30-day spring census in 1977 for BLM-sponsored tortoise studies in California. ‘The 1977 Spring Census Season, Six plots were established in California: Fremont Peak (2 plots), Ivanpah Valley, Goffs, Chemehuevi Wash, and Chuckwalla Bench (Sites 5, 6, 14, 15, 20, and 23, respectively). With the exception of Site 20, Chemehuevi Wash, all sites covered one aq. mile. Plot boundaries usually followed the U.S. rectangular survey system (township, range, and sections). All tortoises were assigned a number and marked following a notching system used by Berry (1974, 1975) and Burge (1977), and later recognized by the Desert Tortoise Council (Fig. 1). Notches vere made on the marginal scutes with a file, or, in the case of juveniles, with fingernail or toenail clippers. The notches vere deep within the scute or close to the underlying bone, The following measurements were taken: maximum carapace Jength at the mid-line (MCL) plastron length from the tips of the gulars to the tips of the anals (PIT); plastron length from the gular notch to the anal notch (PIN), width at the flare of the third marginal (43), width midway along the edge of the sixth marginal (M6), height at the third central, and weight. The measurements were mostly made with Browne and Sharpe calipers, Weights vere taken with Chatillon spring scales. Each tortoise over 180-mm MCL was assigned 4 sex, using combination of characteristics: tail length, degree of development of the chin glands (Rose, Drotman, and Weaver 1969), and shapes of the gulars and plastron. 2-12ume: ysten for 13 on perm ay in California and torto! sber 34.Investigators located tortoises by walking the plot, Most fieldworkers walked "transects," frequently covering a quarter section a day. Systematic searches were conducted to cover the plot evenly over the 30-day census period. Capture date and time, location on the plot, and behavior were recorded for each tortoise observed. Locations were plotted on maps made either from aerial photographs or by the field investigator. All carcasses were collected and their locations plotted. At a later time, data for each carcass were recorded on a standardized Desert Tortoise Shell Data Card (Fig. 2) developed by Lori Nicholson and Berry. Tortoises on three of the six plots were photographed with 35-am color slide fil: Refinements of these initial procedures are discussed below in connection with the individual years or census types. ‘The 1978 Spring Census Season, In the spring of 1978, twelve more sites were established in California: Calico (Site 8), Stoddard Valley (Site 9), Kingston Wash (Site 12), Shadow Valley (Site 13), Amboy (Site 17), Cadiz Valley (Site 18), Lower Ward Valley (Site 19), Pinto Basin (Site 21), Cottonwood Springs (Site 22), Chuckwalla Valley I (Site 24), Ford Dry Lake (Site 25), and Cargo Michacho (Site 27). Field procedures were further refined. Again, each investigator was assigned a plot with a minimum size of one sq. mile. If an investigator was unable to find or mark 30 tortoises within that area, the plot was enlarged to three of more sq. miles. Investigators used standard forms (Fig. 3) developed by Burge and Berry to record data for each live tortoise encountered. All field investigators filled out Desert Tortoise Shell Data Cards (Fig. 2) for each carcass encountered, and collected the remains. Ad-14‘DESERT TORTOISE SHELL DATA caro [> a Froroararme auao [ po pores yee amar ‘CONDITION OF SKELETON evrae aor eve su co ree Eee conoimion or scures owe rossaue wontauny racrons | ee eee Gir a ese prt meta ae Bes ie Fig. 2. The Desert Tortoise Shell Data Card. The top card is side 1, the bottom card is side 2. Developed by Lori Nicholson and Kristin #. Berry, A215More body measurements vere taken in 1978 than in 1977, Additional measurements were wade of carapace width at the midline of the fourth marginal (M4), at the yam between the seventh and eighth marginals (7-8), and at greatest carapace width (Fig. 3). Fieldvorkers also drew abnormalities and/or damaged areas of the carapace, plastron, and limbs onto figure in the data form (Fig. 3). All fieldworkers photographed the carapace and plastron of each tortoise with 35-mm color slide file. These body measurements/and procedures becane the standard BIM techniques for all surveys on permanent plots. Additional notes were taken for each tortoise o temperatures of the air at about 1 @ above ground surface, at 1 cm above the soil, and on the ground, Temperatures vere taken with a quick-recording Schultheis thermometer, with the bulb shaded. General weather conditions at each site were recorded also. In addition, fieldvorkers were asked to record information on all burrows associated with a marked tortoise and observations on feeding. The 1979 and 1980 Spring Census Seasons. vo 30-day study plots, Sheep Mountain and Piute Valley (Sites 29 and 31), vere established in Nevada in 1979 and a third, Last Chance (Site 28), in 1980, Site 5, Frenont Peak, in California, also received a 30-day census. The sane recording techniques were used as in 1978. Expertise of Field Investigators. During the course of the 1977-1979 studies, a number of fleldworkers were employed to undertake the censuses. These fieldworkers had a wide range of expertise-some had five or more years of field experience with collecting similar data on wild desert tortoises, whereas others were familiar only with tortoise captives and the 2-16fea) ieee Re(aus eae re tea site of epoxted Fig. 3. The standardized data sheet for live tortoises used at most California, Nevada, Arizona and Utah study sites since 1978. Developed by Kristin H. Berry and Betty L. Burge. A217Literature. Some were recognized herpetologists with field experience in the California desert, vhile a few were recent college graduates who were familiar with desert reptiles and amphibians and who had degrees in biology. We soon realized that sone individuals could find tortoises and perform a census satisfactorily, vhereas others could not. This ability was important in choosing fieldvorkers for the 60-day census type and in selecting data for analysis (see Appendix 3). The 60-day Spring Census after a preliminary evaluation of population data from 17 sites with 30-day spring censuses (Berry and Nicholson 1979), we judged that 60 days of effort would provide larger and more reliable measures of most population attributes being evaluated. A 60-day census resurvey was conducted on six existing study sites in California in 1979: Fremont Valley (Site 2); Desert Tortoise Natural Area Section 11 (Site 3); Stoddard Valley (Site 9); Ivanpah Valley (Site 14); Chemehuevi Wash (Site 20); and Chuckwalla Bench (Site 23). Also, a new, three sq.-mile plot was established at the Desert Tortoise Natural Area Interpretive Center. In 1980 and 1981, there were eight additional 60-day censuses, five vere at new sites-—Kramer (Site 8), Lucerne Valley (Site 9), Johnson Valley (Site 10), Upper Ward Valley (Site 16), and Chuckwalla Valley II (Site 26)--and three at previously established plots at Fremont Valley (Site 2), Stoddard Valley (Site 9), and Goffs (Site 15). Fremont Valley and Stoddard Valley were resurveyed again in 1981 because the field work was unsuitable. Fremont Valley appeared to have many tortoises with double numbers, mistakes in sex identity, and inaccurate measurements. The 42-18investigator at Stoddard Valley in 1979 used nonrandom search methods, and population density could not be calculated. ‘The field methods were the same as those described for the 30-day spring censuses in 1978, Additional information was gathered on shell-skeletal remains (photographs in situ prior to carcass collection) and on shell wear (see Appendix 4). All investigators were required to collect more data on tortoise burrows and on annual and perennial vegetation. In 1979 and 1980, field investigators were asked to spend a few days in careful searches for hatchling and small juvenile tortoises, These searches were unsuccessful and were not conducted in the 1981 censuses. With one exception, all field investigators selected for the 60-day studies had prior field experience with the 30-day censuses in‘California. The 20-day Fall Census Berry (1974, 1975) and others reported that desert tortoises were generally much less active in sumer and fall than in spring. Not only were fever individuals found per unit of search effort, but there appeared to be a bias in favor of adults. By mid-1979, we had become increasingly concerned about the low numbers of sightings of hatchling and juvenile tortoises and our apparent inability to find more small animals during the 30-day and 60-day spring censuses. We speculated that hatchlings might be nore active and readily observable soon after emerging from nests in fall than in spring. The 20-day fall censuses in October were designed to determine if hatchlings and snall tortoises vere more observable in the fall and to quantify observations on the size classes of tortoises that A2-19were active. Seven study sites were surveyed in California: Fremont Valley (Site 2); Desert Tortoise Natural Area, Section 11 (Site 3); Fremont Peak (Site 5); Stoddard Valley (Site 9); Ivanpah Valley (Site 14); Chemehuevi Valley (Site 20); and Chuckwalla Bench (Site 23). Procedures for collection of field data vere as described earlier. OTHER TYPES OF SURVEYS Year-long or Multi~season Studies Seven year-long or multi-season studies have been conducted in the Southwest at Argus, California (Site 1); Arden, Nevada (Site 30); the Beaver Dam "Slope" (Sites 44-46) and the Picacho Mountains in Arizona " Utah (Site 33). Each study is (Site 42); and the Beaver Dam "Slope, discussed separately below. Site 1, Argus, California, The site was used in a study of wild tortoises relocated from a highway construction project (Berry 1974). The site received intensive surveys during 1971 and 1972 to locate che transplanted tortoises, Data also were collected on resident tortoises. Sixty-one tripe were made between July and Novesber 1971. One to four people worked in the field on each occasion; work hours per trip ranged from two to 20, In 1972, 46 trips were taken between February and June 30, and 33 tripe were taken between July 1 and December 30. One to five individuals worked in the field on each occasion; work-hours per trip ranged from two to 30. The initial survey technique involved location of tortoi: » capture, marking (by notching), and release. Sizes of tortoises (seven parameters), weight, and sex were recorded. Time of day and period of observations, 42-20behavioral activities, location and orientation, daily moveneats, construction and use of burrows, tines of emergence and retreat, associations with other tortoises and other species, and weather conditions were recorded. Site 1 s unlike most of the other 46 sites, ‘The plot has soft windblown sands in many areas. The survey method involved tracking tortoises. We located tortoise footprints and followed tracks to the animals. Fieldworkers recorded locations and behaviors of tortoises using the tracks. Wind gusts over 15 to 20 niles per hour and rain occasionally obliterated all traces of tortoise activities. Such circumstances force Eieldworkers to seek tortoises by systematically surveying the study plot ‘The extent of the study area was gradually enlarged from fall of 1971 through 1972, reaching a final size of 5,3 sq. miles. It is believed that most, if not all, resident tortoises within the study area were marked by the end of 1972, The roster of tortoises used in the data analysis represents the individuals occupying the plot from fall 1971 through fall 1972. Site 30, Arden, Nevada, This study site was established by Burge as part of a thesis project (Burge and Bradley 1976; Burge 1977). Data were collected between May of 1974 and October of 1975. Radio transmitters vere placed on some tortoises. Burge (1977) reported spending 188 calendar day: (936 hours) in the field during that time, Visits generally were made at one- to three-day intervals for five to seven hours each day, Fever visits occurred between November and February. Although the emphasis in the field was on locating tortoises fitted with radio transmitters, every effort was made to locate and to recapture other resident tortoises (Burge and Bradley A2-211976). Burge (personal communication) prepared a roster of individuals to represent the population on the study plot in spring, summer, and fall of 1975 for this analysis. Sites 44-46, Beaver Dam Slope, Arizona I, II, and IIT, This group of study sites had tvo principal field investigators, Hohman (Hohman and Ohmart 1980) and Sheppard (1980, 1981). fchman and Ohmart (1980) were funded by the BLM in 1977 and 1978 to determine population density and structure on the Beaver Dam Slope in Arizona and study dietary overlap between desert tortoises and cattle. Sheppard continued the work from 1979 ro 1981. Neither Hohman nor Sheppard had any field experience with desert tortoises prior to the study, The numbers of days spent per month surveying for tortoises between 1977 and 1980 varied from 0 to 20. The number of days spent in the field by Hohman in the spring of 1977 and 1978 exceeded those for a 30-day spring census but fell short of the 60-day spring census. The numbers of field days for Sheppard in 1979 and 1980 fell short of a 30-day spring census. Field time was divided equally between Sites 44 and 45, the control and exclosure, respectively. Both Hohman and Sheppard provided rosters of the first encounters of individuals for each year for this study. We have pooled the data for the sites for most analyses. ‘The census of tortoises on these sites was not conducted with systematic search patterns. Sheppard (personal communication) reported that tortoises with radio transmitters were sought first, then searches were made for other resident tortoises, with emphasis on places of probable 2-22Site 42, Picacho Mountains, Arizona, Tortoises at this site were studied from 1975 to 1979 by Schwartznann for a thesis project (Schwartznaan, a.d.), Over the four-year period, Schwartzman spent 130 calendar days making observations: nine days in January, four in February, six in March, 13 each in April and May, nine in June, seven in July, 15 in ‘August, 17 4a September, 29 in October, five in November, and three in December. During this same tine period, 53 individual tortoises were captured and marked. Eleven were fitted with radio transmitters. Our roster of tortoises was provided by Schwartzmann (personal communication) and includes information on first encounters of the 53 individuals for the period from 1975 to 1978. Site 33, Beaver Dam Slope, Utah II, This site provided one of the more difficult data sets (Coombs 1974, 1977a, 1977b), Coombs studied tortoise populations on the Beaver Dam Slopes from 1973 through 1976 (time period inferred from Coombs 1977b, Table V-13). There is no complete roster of tortoises for the study period in any of his reports or papers. A partial list of individuals can be obtained from the data sheets in Coombs (1974); some have more specific localities noted on the data sheet: It is not clear from the study site descriptions or discussion of methods (Coombs 1977a, 1977b) how auch area was surveyed on the Beaver Dam Slope how many days were spent in the field, and when, which, and how many tortoises were captured in the “intensive study areas" and "high density areas" shown in Coombs (1977a, Fig. 2), or in Coombs (197b, Fig. V-5). Our data base for this study area is incomplete. 2-23Winter Den Surveys Winter den surveys have been conducted at two study sites, one on the Beaver Dam Slope, Utah (Site 32) and the other on Tiburon Island, Sonora, Mexico (Site 43). Site 32, Beaver Dam Slope, Utah I. This is the classic study of Woodbury and Hardy (1948) on an approximately two sq. ile area in southwestern Utah, It is Saportant to recognize at the outset that this study vas done very differently from other desert tortoise studies in the United States and for different reasons. In their methods section, Woodbury and Hardy (1948) stated: "This study 1s an outgrowth of Anvestigations made on snake dens and was undertaken to provide comparative material for interpretation of snake behavior in concentrating in winter dens." Ninety trips of one to five days duration were made to the site between November of 1936 and April of 1946 (Woodbury and Hardy 194 Hardy, personal communication). Hardy (personal communication) provided a list of trip dates. Fifty-three trips were made between November and the end of March and 47 between April and October. More tortois were found per trip during the November to March period than in the April to October period. Most tortoises were captured by pulling them from dens. Hardy (1976) returned to the site on several occasions after the initial study period to recapture and mark additional tortoises, but at no time repeated surveys similar in extent to the 1936-1946 period. Woodbury and Hardy (1948) collected data on sizes and sexes for sone of the 281 tortoises marked during the 1936 to 1946 period. Data for 126 tortol, 1 and 2 of (both marked and unmarked) were presented in Table 82-24their pay Our roster of tortoises is taken from those tables. Most of the over 200 observations listed in their tables vere nade between Novenber and March. Site 43, Tiburon Island, Sonora, Mexico. We have placed this site in the winter-den survey category, although Reyes Osorio and Bury (1982), sho surveyed the area in 1978 and 1979, might disagree. They surveyed 35 transects, each 10 a wide and 3 to 7 ka in length, between Septenber 1 and Novenber of 1978 at five sites on the southeastern end of Tiburon Island: Valle del Chalate, Calle del Cerro Prieto, Ensenada de Don Punta Ballena, and Valle de los Espanoles. Our roster of 120 tortoises comes from these five sites (Reyes Osorio, unpublished manuscript). There are neasurenent data for another 12 tortoises from other sites on Tiburon in Reyes Osorio and Bury (1982) from the 1979 surveys, but those are not included in our roster, since we lack this information. Reyes Osorio and Bury (1982) reported that tortoise activity appeared to be most concentrated in the cooler fall season: sssIn the present survey, 32% of all tortoises seen were out in September, 45% in October, and 23% in November...Few are seen at other times of year. Unfortunately, their report does not address the following questions: (1) what were the nunbera of search days and person-days of effort per month? (2) how many tines were the transects checked? (3) how many tortoises were found per day of search effort, per month? (4) how many tortoi: were recaptured? and (5) who conducted the surveys from December through August and for how long? (we are referring to the statement above chat “Few are seen at other times We are aware of only one report of other surveys on Tiburon Island. Bury, Luckenbach, and Munoz (1978) found 42-25a single tortoise above ground on Decenber 9, 1977, during two days of secting up study plots and walking transects during that month. They found Lieele other sign of tortoises during their survey. Reyes Osorio and Bury (1982) noted that few very young tortoises and juveniles were found (<180-mm MCL), and that the ratio of adults to nonadults was 9:1. They reported that the population profile resembled that from che Beaver Dam Slope in Utah during the Woodbury-Hardy study effort. We agree that the size class structure of tortoises encountered is similar, Reyes Osorio and Bury (1982) suggested that juveniles and immatures (<180-mn MCL) might have been difficult to find because of a “natural phenomena” or the dense vegetation. We disagree, particularly, with the latter explanation. One plot in California with dense vegetation typical of the Sonoran Desert (Site 23, Chuckwalla Bench) has a high tortoise density and a relatively high proportion of young individuals in the population. The comparatively dense vegetation in that area is similar in some respects to Tiburon Island and has not hindered encounters. While there are a number of possible explanations for the extremely low percentage of small individuals found, we favor the survey technique—the time of year the work was done--and the limited expertise of the investigators a! possible explanations. First, ve doubt that fall is the principal activity period for the reasons discussed earlier, Second, the importance of field expertise cannot be underestimated, especially in finding the smaller tortoises. Until we are confident that the population profile reported by Reyes Osorio and Bury (1982) is an accurate reflection of the tortoise populations on Tiburon (or at least similar to a 60-day spring census), we will call this a "winter den" survey, 2-26Special Surveys Nine study sites with their data sets vere placed in this category: two sites on the Beaver Dam Slope, Utah (Sites 34 and 35) (Minden 19803 Minden and Keller 1981); six sites in southern Arizona (Sites 36-61) studied by Schnedder (1981); and the Barstow study area, Site 47 (Fusari 1981). Site 34, Beaver Dam Slope, Utah III, Minden (1980) surveyed approximately 148 sq. miles between April 9 and June 26. Although the study effort covered 60 days and 500 work hours, it vas not the typical 60-day spring census described earlier, The study area vas extrenely large. Minden (1980) conducted quadrat surveys, tracked adult fenales with radio transmitters, surveyed dens, and made tortoise surveys over a very large area. He had not worked with desert tortotses in the field prior to this experience. Site 35, Beaver Dam Slope, Utah IV. Minden and Keller (1961) established a one-sq.-mile census plot in the spring of 1981 within Site 32, the two-sq.-mile, Woodbury-Hardy study area, Minden and three other investigators spent 26 days on site, and 532 hours (equivalent to about 66 eight-hour workdays). With the exception of Minden, the other fieldvorkers apparently had no experience with tortoise censuses. About 185 hours were consumed in subdividing the plot into 100 quadrats and another 77 hours in sampling vegetation. Although tortoises were found during the-vegetation studies and setting up of the grid, only 270 hours (equivalent to 35 eight-hour days) vere spent in "systematically censusing the plot" for tortoises. (Ia the typical SLM 60-day spring census, no more than two days each were consumed in making vegetation transects and setting 42-27up a grid). While this plot certainly had survey time in excess of the typical 30-day spring census, it did not receive the minimum nuuber of calendar days (about 40) for a 60-day spring census and had only one experienced field surveyor. Sites 36-41, Arizona: Alamo Hill, Stout Mountain, Little Shipp Wash Harcuvar Mountain, Black Butte, and Ragged Top. These sites vere all surveyed in the spring of 1980 between March and mid-June by Schneider (1981) and four fielévorkers for "124 field days." The four other fieldvorkers apparently had no desert tortoise experience. We are uncertain how many of the 124 days were spent searching for tortoi on the plots themselves, because some time was spent fitting tortoises with radio transmitters and tracking them, making plant transects, and searching areas adjacent to the plots for tortoises. ‘There is no information on the number of calendar days of searching effort for each site. The number of "field days," which may be equivalent to person-days or workdays is shown in the text. The number of field days and timated number of calendar days are shown in Table 2. ‘Two sites, Alamo Hill and Little Shipp Wash, had the most search effort. The data from Little Shipp Wash may be comparable to a 30-day spring census, but that from Alamo Hill is questionable, with apparently 13 calendar days and 22 field days, Stout Mountain was abandoned for Alano Hill, which had higher tortoise densities Site 47, Barstow, Calif. Fusari (1981) undertook a special study for the California Department of Transportation on the efficacy of fences as barriers and culverts as "tortoise underpasses" for protecting tortoise populations adjacent to highvays. There is a cumulative list of 60 42-28Table 2. Numbers of field and calendar days spent at six desert tortoise study sites in southern Arizona, as calculated from Schneider (1981). Site name Field days Estimated . Live Dead (shown in text) calendar days* tortoises tortois Alano Hill 22 43 (2 in April, 29 8 10 in May, 1 in June) Stout Mountain 8 6, all in April 8 ° Little Shipp Wash 40 27 (24 in April, SL 8 ‘and May, 3 later) Harcuvar Mountain as 1 1 ° Black Butte 12 3 4 ° Ragged Top 27 7 7 o a, This number may be the minimum; it was determined by using the appendices in Schneider (1981). There may be days of survey not shown in this appendix because no tortoise captures were nade 42-29tortoises found at the site in March, May, September, and October of 1978, and March, April, and May of 1980 in Appendix II of Fusari's report. Another List of 25 tortoises found on April 24 on a “transect extending for 1/2 mile south of the Sidewinder Road exit on I-15 on the west side of the freeway fence” is presented in Appendix III of Fusari's report. No details are offered on how much time and how many people vere involved in each rch effort. Data from Fusari's Appendix II appear to have been acquired during behavioral studies of tortoises contacting an experimental fence and culvert system. We have decided not to use any data from Site 47 because of the limited amount of data gathered and the technique used. 2-30REFERENCES CITED Barrow, J. 1979, Aspects of the ecology of the desert tortoise, Gopherus ‘agassizii, in Joshua Tree National Monument, Pinto Basin, Riverside County, California. Pages 105-131 in E. St. Amant (ed.), Proc. 1979 Symposium of the Desert Tortoise Council, Long Beach, Calif. Berry, K. H. 1974, Desert Tortoise Relocation Project: Status Report for 1972." Division of Highways, State of California. Contr. No. F-9353, Section III, C. 3. Berry, K. H. 1975. Desert Tortoise Relocation Project: Status Report for 1973. Division of Highways, State of California. Contr. No. F-9353, Section III, C. 4. Berry, K. H. and L. Nicholgon. 1979. ‘The status of the desert tortoise in california. Draft report. U.S. Dept. of Interior, Bureau of Land Management, California Desert Plan Program, Riverside, Calif. Berry, K. H. 1980, State Reports--California. Pages 61-67 in K. A. Hashagen (ed.), Proc. 1980 Symposium of the Desert Tortoise Council, Long Beach, Calif. Burge, B. L. 1977. Movements and behavior of the desert tortoise, Gopherus agassizi. M.S. Thesis. University of Nevada, Las Vegas. Burge, B. L., and W. G, Bradley. 1976. Population density, structure, and feeding habits of the desert tortoise, Gopherus agassizi in a low desert study area in southern Nevada. Pages 57-74 in N. Engberg, S. Allen, and R, Young (eds.), Proc. 1976 Symposium of the Desert Tortoise Council, Long Beach, Calif, Bury, R. B., and R. L, Luckenbach. 1977. Censusing desert tortoise populations using a quadrat and grid system. Pages 169-178 in M, Trotter and C. G. Jackson, Jr. (eds.), Proc. 1977 Symposium of the Desert Tortoise Council, San Diego, Calif. Bury, R. Bs, Re As Luckenbach, and L. R, Munoz, 1978. Observations on Gopherus agassizii from Isla Tiburon, Sonora, Mexico. Pages 69-74 in WM. Trotter and C. G. Jackson, Jr. (eds.), Proc. 1978 Symposium of the Desert Tortoise Council, San Diego, Calif. Coombs, E. 1974, Utah cooperative desert tortoise study Gopherus agasei2ii. Unpublished M.S. on file at U.S. Dept. of Interior, Bureau of Land Management, Cedar City, Utah. Coombs, E. M. 1977a, Status of the desert tortoise, Gopherus agassizii, in the State of Utah, Pages 95-101 in M. Trotter and C. G. Jackson, Jr. (eds.), Proc. 1977 Symposium of the Desert Tortoise Council, San Diego, Calif. 2-31Coombs, E. M. 1977b, Wildlife observations of the hot desert region, Washington County, Utah, with emphasis on reptilian species and their habitat in relation to livestock grazing. Report to U.S. Dept. of Interior, Bureau of Land Management, Cedar City District, Utah from Utah Div. Wildlife Resources, Fusari, M, 1981. Feasibilicy of a highway crossing system for desert tortoises. Final Report to the California Department of Transportation, Sacramento, Calif. Hardy, B. 1976, The Utah population--a look in the 1970's. Pages 84-88 in'N. J, Engberg, S. Allan, and R. L. Young (eds.), Proc. 1976 ‘Symposium of the Desert Tortoise Council, Long Beach, Calif. Hohman, J, and R. D. Ohmart. 1980. Ecology of the desert tortoise Gopherus agassizii) on the Beaver Dam Slope, Arizona. Report to U.S. Dept. of Interior, Bureau of Land Management, Arizona. Contr. No. YA~S10-PH7-54, Karl, A. 1979a. An ecological study of a population of desert tortot: Gopherus agassizi, in Clark County, Nevada. Report to U.S. Dept. of Interior, Bureau of Land Management, Denver, Colorado. Contr. No. YA-512-CT9-90, Karl, A. 1979b. An ecological study of a population of desert tortoises, Gopherus agassizi, in southern Nevada. Report to U.S. Dept. of Interior, Bureau of Land Management, Denver, Colorado. Contr. No. YA-512-CT9-90, Karl, A. 1980, The ecology of a population of desert tortoises, Gopherus ‘agassizi, near Pahrump, Nevada, Report to U.S. Dept. of Interior, Bureau of Land Management, Denver, Colorado, Contr. No. YA-512-CT9-90. Minden, R. L. 1980, Investigations of the desert tortoise (Gopherus agassizii) on the Beaver Dam Slope, Washington County, Utah. Utah State Div. Wildlife Resources, Publ. No. 80-21. Minden, R. L. and $. M. Keller. 1961. Population analysis of the desert. tortoise (Gopherus agassizii) on the Beaver Dam Slope, Washington County, Utah, Utah State Dept. Natural Resources and Energy, Div. Wildlife Resourc Reyes Osorio, S. n.d. Aspectos biologicos de 1a tortuga del desierto (Gopherus’ agassizii) in Isla Tiburon, Sonora. M.S. on file at the Bureau of Land Management, Riverside, Calif. Reyes Osorio, S., and R. B. Bury, 1982. Ecology and status of the desert tortoise (Gopherus agassizii) on Tiburon Island, Sonora. Pages 39-49 in R. B. Bury (ed.), North American Tortoises: conservation and ecology. WS. Dept. of Interior, Fish and Wildlife Service, Wildlife Res. Rept. 12, Washington, D.C. 2-32Schneider, P. B. igert, a population analysis of the desert tortoise Gopherus agassizii in-Arizona. Report to U.S, Dept. of Interior, Bureau of Land Managenent, Phoenix District Office, Arizona. Contr. No. AZ~950-CT9-0014, Schwartzmann, J. L, n.d. Untitled draft for Masters Thesis, Arizona State University, Teape, Arizona. Sheppard, G. P. 1980. Status of the Arizona Beaver Dam Slope population of desert tortoise. Pages 23-32 in K. A. Hashagen (ed.), Proc. 1980 Symposiua of the Desert Tortoise Council, Long Beach, Calif. Sheppard, G. P. 1981, Desert tortoise population of the Beaver Dan Slope in northwestern Arizona, Pages 25-47 in K. A. Hashagen and E. St. Anant (eds.), Proc. 1981 Symposium of the Desert Tortoise Council, Long Beach, calif. Woodbury, A. M. and R. Hardy. 1948, Studies of the desert tortoise, Gopherus agassizii, Ecol. Monogr. 18:156-200. 1, This report has been distributed in several drafts, each with the same title and date. We refer to the draft distributed in February, 1982. 42-33Appendix 3 METHODS USED IN ANALYZING DESERT TORTOISE POPULATIONS Frederick B. Turner and Kristin H. Berry This appendix covers a number of topics: selection of data sets for use in the principal analys. of populations; corrections of these data sets; arraying population data for live tortoises by size and sex; analyzing densities in selected populations; and clustering population data by regions within the geographic range. Each topic is discussed separately below. SELECTION OF DATA SETS FOR USE IN PRINCIPAL ANALYSES OF DESERT TORTOISE POPULATLONS Data sets for 46 study sites were compiled. For some sites only one season of information was available, whereas for others several years of data existed. A summary of the dates and types of censuses and surveys for each of the 46 sites was presented in Table 1, Appendix 2. We selected only 27 data sets for the principal analy! (Table 1). Four criteria vere used in determining which data sets should be included in the analysis: (1) the abundance of tortoises on the study plot, (2) experience of the fieldworker or workers, (3) the methods used by the workers, and (4) duration of the study. Abundance of Tortoises Data from thirteen study plots were not used because of low tortoiseTable 1. Study sites, census type, and data analysis on the desert tortoise, selected for principal data Site no. Site name Census year and type 1 Southern Argus Range, Calif. ‘1971-1972: year-long 2 Fremont Valley, Calif. 1981: 60-day spring census 3 Desert Tortoise Natural 1979: 60-day spring census ‘Area (Sec. 11), Calif. 4 Desert Tortoise Natural + 60-day spring census ‘Area Inter. Center., Calif. 5 Fremont Pesk, Calif, 30-day spring census 7 Kraner, Calif, 1980: 60-day spring census 8 Calico, calif. 30-day spring census 9 Stoddard, Calif, 1981: 60-day spring census a0 lucerne Valley, Calif. 1980: 60-day spring census u Johnson Valley, Calif. 1980: 60-day spring census a3 Shadow Valley, Calif. 1978: 30-day spring census uu Ivanpah Valley, Calif. 1979: 60-day spring census 1s Gofts, calif. 1980: 60-day spring census 16 Upper Ward Valley, Calif. 1980: 60-day spring census 20 Chemehuevi Wash and 1979: 60-day spring census Valley, Calif. a Pinto Basia, Calif. 1978: modified 30-day census 23 Chuckwalla Bench, Calif. 60-day spring census 26 Chuckwalla Valley II, Calif. 1980: 60-day spring census 29 Sheep Mountain, Nev. 1979: 30-day spring census 30 Arden, Nev. 1974-1975: year-long 31 Piute Valley, Nev. 1979: 30-day spring census Aa2Table 1. Study sites, census type, and data selected for principal da: analysis on the desert tortoise. (Continued) Site no. Site name fensus year and type 32 Beaver Dam Slope, Utah T 1936-1946: winter den study 35 Beaver Dam Slope, Utah IV 1981: spectal eurvey (spring) 36 Alamo Hill, Ariz. 1980: spectal survey (spring) 38 Little Shipp Wash, Ariz. 1980: special survey (spring) 43 Tiburon Island, Mexico 1978: fall and winter surface and den study 44-46 Beaver Dan Slope, Ariz. 1977: year-long 1978: year-long 1979-1980: year-longpopulations or no tortoises (Table 2). There were eight such sites in California, one in Nevada, and four in Arizona. The site with the smallest population sample used in the analysis was Alamo Hill, Arizona (Site 36), with 29 individuals. Experience of Fieldworkers General experience and/or the ability to find tortoises is a critical element in a census. Two data sets, Stoddard Valley and Chenehuevi Wash and Valley (Sites 9 and 20), were not used because of the denonstrated inability of surveyors to find tortoises; a third set (Site 34) was discarded for this and other reasons (Table 2). Some herpetologists and field biologists readily find tortoises, while others cannot. Inexperienced field personnel with unusual and outstanding abilities also exist. One problem may be the number of search dmages che investigator aust keep in mind: numerous sizes of tortoises, ranging from approximately 4S-mm MCL for a hatchling to 300-mm MCL for a Large adult; and the different kinds of tortoise sign which might direct the observer to a nearby tortoise such as burrows, scats, tracks, courtship rings, eggshells, and drinking sites. Variation in visibility of tortoises in different habitats within the geographic range and variation in the kinds of sign in different habitats compound the problem. Expertise also appears to be a strong factor in finding small tortoises <100-am MCL. Our best locators of small tortoises have spent several years in the field surveying permanent study plots and their abilities to find tortoises may be improving with experience. : 1. We plan to check this statistically in the fall of 1983. AB-4Methods Used by the Fieldworkers and Duration of Studies Where several data sets were available for a single site, we chose the set most likely to provide the best information on apparent population structure. The order of preference was: (1) a 60-day spring census and/ or selected year-long studies, (2) 30-day spring census, (3) special studies in the spring, and (4) winter den studies, Some reasons for our preferences are presented below. The 60-day vs. 30-day Spring Census. We speculated that 60-day spring censuses might favor capture of small tortoises and provide more reliable samples of the actual tortoise populations than the 30-day census. Three areas with both 30-day and 60-day spring censuses were compared using x teste: Ivanpah Valley, Goffs, and Chuckwalla Bench (Tables 3 and 4). The findings vere equivocal. More small tortoises were found at Goffs and the Chuckwalla Bench during the 60-day censuses. The two samples from Ivanpah Valley were almost identical and did not differ significantly. ‘At this point we cannot state that relatively more snall tortoises will always be found during a 60-day spring census than during the 30-day census. Aside from the actual relative abundance of small tortoises, at least two other factors might affect the size composition of the samples: (1) the abilities of fieldworkers, and (2) weather conditions during the time of the census, Although the tests reported in Table 4 are not wholly conclusive, we believe that longer censuses are more likely to reflect true size distributions of populations than shorter ones. We did not include Reasons for Excluding the 20-day Fall Census any data from 20-day fall censuses because fall samples vere significantly different from those data acquired during the 60-day spring censuses. In Aa-7uasuay sedexe9 96T Zot 6tz Stt ert SL ont ast ont < yaBuet sordeses 69 n re) a 9 st 9 ” oot > 6L6t——_LL6T oget Let sist Let Test _6L6T (w=) fep-o9 _&ep-0g K-09 KeP-oc © KeP=09 Kep-or = ep-09 KP ~09 ozya youeg PTTeRyaRYD 83309 KotteA yeduent AorTea qwowsaa aeforz01, snsua> jo 3vak pue ‘ad42 snsuss ‘es 208-mm MCL) tortoises observed at 18 sites in California, three in Nevada, three in Arizona, and one in Utah. To compare observed ratios with an sumed sex ratio of 1:1 we computed ~statistic for each site: Z = (p~ 0.5)/(0.25/n)!/2 @ 3-17with p the proportion of males in the sample and a the total sample size. Z 4s equivalent to the square root of x”, but is simpler to understand because Z is distributed normally with unit variance. Hence, 95 and 99 per cent confidence intervals for Z are 41.96 and 42.57, respectively. Data from four sites in California indicated statistically significant departures from a sex ratio of 1:1. The probability of observing statistically significant differences is increased as uore data are available. Hence, it is worth looking at the distribution of Z-values from all 25 sites. Does this distribution have a mean of zero and unit variance? We also considered observed sex ratios of subadult tortoi: 5 and adult tortoises separately. Because male tortoises attain larger sizes than females, it is obvious that the apparent sex ratio can be influenced by how size categories are established. Does the available evidence suggest that setting @ lover size Limtt of 208-mn MCL for “edulte" has any eftect on apparent sex ration? To teot the mean of a distribution of Z-values GZ), one computes Ze) '/2, wnich itself has an expected mean of zero and uit variance. To test the variance of a distribution of Z-values, Var(Z)(n-1) is compared with x? (g-l degrees of freedom). Table 7 gives data for such comparisons, based on observed numbers of (1) all ‘les and females >180-nm MCL in 25 populations, (2) adult males and females >208-mm MCL in 25 populations, and (3) subadult male and females 180- to 207-am MCL in ‘15 populations. Subadult tortoises from ten populations were not included because of their small numbers (Argus, Calico, Johnson Valley, Shadow Valley, and Pinto in in California; Sheep Mountain in Nevada; Beaver Dam Slope, Alano Ag-18oz"0~ 06°8t 919°0- ser 6sT“0- st 19m ™~207 0 -08t) seTemay pus ‘sotem atnpeqns 100"0 81S fe "T ore 99z"0 sz (Tow wa-goz J) soTemoy pur SeTem 3TNPy 20°0~ 9e"6e s68"0 2" eit"o sz (row moet Z) seteRs ‘pue soTem TTY : pezktour _ 7 oo sontea-Z ——suoyaetndod a DOr — y7DZ Daeg yo wea Jo z9quny, sénox8 2275 ssooueyaea 3Fun 30 3603 303 uaayS axe Z 30 senteA *sasyor103 q198ap jo sdnox8 277s 9e3y2 03 BuyUTera—d SanTeA-Z Jo SuOFINgTAISTP Jo SUOFIENTEAT */ PTAPL A319M111, and Little Shipp Wash in Arizona, and Beaver Dan Slope, Utah). The teats of Z-values in Table 7 are interesting ta thet none of the values of Z(a)l/2 is statistically significant. Hence, the neaas of the various groups anélyzed do not differ from zero, On the other hand, tests of the dispersions of Z-values give contrasting results. The variance of the distribution of 15 values pertaining to eubadult tortoises does not differ signigicantly trom one, Thus, there is no evidence that the observed numbers of subadult male and fenale tortoises differ from an assuned ratio of 1:1. The distributions of larger tortoises both exhibited variances significantly greater than one, The distributions are normal but too long in the tails, Hence, there are statistically significant differences between populations and departures fron balanced sex ratios. The effect is principally expressed anong tortoises >208-an MCL. ore specifically, the differences Lie in four samples from California: Argus and a portion of the Desert Tortoise Natural Area (with a preponderance of females), and Goffs and vard Valley (with a preponderance of males). The foregoing analyses have assuned that the expected sex ratio among desert tortoise populations is 1:1, and that significant departures from this expectation may be indicative of suboptinal conditions, If ales and females impose identical costs to their parents, natural selection acts to produce sex ratios (Fisher 1958, Planka 1974). Most vertebrates exhibit such proportions, and sex ratios differing significantly from 1:1 are nore prevalent among invertebrates (Nichols and Chabreck 1980). We now know, however, that temperature may affect the sex of developing turtle embryos, leading to sex ratios differing conspicuously from (ull 1980). 43-20Present understanding of sex determination in reptiles has been reviewed by Bull (1980), and the following quotations relating to turtles are drawn from his paper. Two factors in reptile sex determination have been studied: sssthe presence or absence of heteromorphic sex chromosomes, and is.the influence of temperature. Recognizable sex chromosomes are ‘sapparencly rare in turtles..,Temperature-dependent sex determin— ation...is common in turtles.,,.Present findings...suggest that... temperature actually determines sex rather than simply causing differential mortality, and...temperature controls sex determina- tion in nature as well as in the laboratory, Only one study...has convincingly demonstrated the latter. Sex ratios in turtles were discussed by Gibbons (1970), who pointed out that because a number of papers indicated an apparent predominance of females in turtle populations Carr (1952) had jerted that "...in many apecies...females outnumber males." Gibbons analyzed sone of these data, focusing particular attention on problems in sampling bias and difficulties in determining the sexes of turtles. Gibbons concluded that "...there are insufficient data to support the notion that turtles often have unequal sex ratios. Improper methodology and selective sampling...have led to this conception, Unequal sex ratios may indeed occur in natural popula~ tions of turtles, but the data presently available...do not substantiate rhe assumption. Gibbons (1970) commented on two studies in which investigators selected an inappropriate size limit in attempting to determine relative numbers of males and fenales. Both studies involved species in which males and females matured at different body sizes. In the firet ca Sexton (1959) described a population of Chrysemys picta composed of about 57 per cent males. Gibbons argued that had Sexton used a lesser plastron length as a criterion of female maturity the sex ratio would have been A321close to 1:1 because previously excluded mature females would have then been included in the analysis. Cagle (1942) apparently erred in the opposite direction in his analysis of a sample of Pseudemys scripta reporting an excess of females when the sex ratio (based on a larger size of females at maturity) was really 1:1. A ore common difficulty is that males and females may differ in the ease or frequency with which they are observed. Finally, it may be difficult to discriminate between the sexes on the basis of secondary sex characteristics, Gibbons (1970) gave examples of both kinds of problems. More recent discussions of sex ratios in turtles are included in Auffenberg and Iverson (1979) and Bury (1979). Comparing 30-day and 60-day Spring Censuses of Torto: In 1977, five areas were inspected for 30-day periods during the spring. Two to three years later, three of these areas were examined again for 60 days in spring, Table 8 gives numbers of males and females >180-am MCL registered in these areas in different years. Tests of the 30-day and 60-day census data from these sites shoved that the population structure of larger tortoises inferred from 30-day censuses was no different from that revealed by 60-day censuses. chi-squared values (3 4.f, in all cases) were 2.2 for Ivanpah Valley, 4.1 for Goffs and 4.0 for Chuckwalla Bench. Regional Comparisons of Observed Sex Ratios of Tortoises. Table 9 gives data pertaining to 985 male and 968 female tortoises >180-mm MCL grouped according to geographic locale. sits represented in the western Mojave Desert are Argus, Fremont Valley, both Desert Tortoise Natural Are sites, Fremont Peak, Kramer Hills, Calico, Stoddard Valley, Lucerne Valley, and Johnson Valley, Sites from the eastern Mojave Desert are ShadowTable 8. Comparisons of samples of desert tortoises 2 180-mm MCL from three areas taken during 30- and 60-day spring censuses. Site site Census Subadults ‘Adulte no. name Males Fenales Males Females Totals 16 Ivanpah Valley, 30 days, 2 6 a1 26 35 calif, 1977 60 days, 7 8 39 3 ar 1979 15 Goffs, calif. 30 days, 2 u a2 28 99 1977 60 days, uw cry 95 35 186 1980 23° Chuckwalla 30 days, a 9 33 35 88 Bench, Calif. 1977 60 days, 9 ub 70 60 153 1979 3-23Table 9. Numbers of male and female tortoises 2180-mm MCL recorded in six parts of thé geographic range of the desert tortoise. Region Number of Males Females z sites Western Mojave 10 456 517 desert, Calif. Eastern Mojave 5 256 201 2.37 desert, Calif. Southern, 3 11g 113 0.40 desert, Calif. Nevada 4 1 78 0.99 Central and 6 31 39 0.97 western, Arizona Beaver Dam slopes, 2 32 20 1.66 Arizona and Utah Ag-24Valley, Ivanpah Valley, Coffs, Upper Ward Valley, and Chemehuevi Wash. Southern Desert sites are the Pinto Basin, Chuckvalla Bench, and Chuckwalla Valley. Nevada is represented by three sites (see Chapter 8), as vell as data pertaining to nine tortoises from Last Chance. The Arizona samples represent principally Alamo Hill and Little Shipp Wash (see Chapter 10), but also include 15 tortoises from four other areas examined in 1980 (Schneider 1981). Samples from the Utah and Arizona portions of the Beaver Dam slopes are considered jointly. When considered over the entire range of these particular sampl total ounbers of males and fenales recorded were essentially identical. However, a test of the relative numbers of males and fensles in samples from six regions shows the distributions to differ significantly (x° = 15.1, X2y,01 * 15-1). About 70 per cent of the total x derives from the western and eastern Mojave samples, exhibiting more or less complenentary patterns of relative abundance. A test of these two areas alone gives a X° of about 10.1, and an associated probability of around 0.001 that the collective samples could have been dravn from the same populations. Any geographic differences existing in the data analyzed are again the result almost entirely of tvo highly unusual samples: the De: rt Tortoise Natural are in 1979 (with an enormous preponderance of females), and Goffs in 1980 (with many more males). ‘The Goffs' 1980 sample is perplexing because none of the work carried out in previous years is consistent with the outcome. For example, in 1977, a 30-day census revealed 54 male and 45 female tortoises >180-mm MCL. A year later, a total of 22 male and 13 female subadult and adult tortoises were observed during a brief inspection. 43-25,Site 4 at the Desert Tortoise Natural Area is near Site 3, which is also in the Natural Area, As shown in Chapter 5, the 1979 census of Site 3 indicated equal numbers of tortoises >180-um MCL. It is puzzling, chen, that in the same year so many more females vere registered at Site 4. The work at Site 4 in 1979 was carried out over an area of three sq. miles by three different people working for 60 days. This differs from a one-person examination of one sq. mile for 60 days. However, it is not clear how this difference could affect the sample composition. Site 4 differs from Site 3 in the levels of human disturbance, which may account for some differences in sex ratios. Site 4 has numerous human uses: (1) 2 major dirt road with heavy vehicle traffic and several minor dirt roads; (2) one of the higher levels of off-road vehicle use in the California deserts; (3) documented illegal collecting and vandalism of tortoises; and (4) sheep grazing. Part of Site 4 is fenced for protection from vehicle use and sheep grazing; the other part receives heavy impacts (see Appendix 1). ESTIMATING DENSITIES OF DESERT TORTOISE POPULATIONS The present abundance of the desert tortoise in various parts of its geographic range is a clearly important element in our understanding of the biology of this species. Future land management decisions will require reliable information as to the density of tortoises, particularly in California. For example, Southern California Edison Company is pré ntly considering a number of areas in southeastern California as locales for energy facilities. Ivanpah Valley has been considered a possible site for a large coal-burning power plant. The construction of a 100-tW, solar 83-26‘thermal power plant in Johnson Valley is also being considered. In both of these instances, the California Energy Commission has identified the desert tortoise as an important biological issue. The Commission has requested estimates of the absolute abundance of tortois at both sites, and some vork along this line has already been done in Ivanpah Valley (Nicholson, 1980; Turner et al. 1982). Information as to the abundance of tortoises in areas to be sacrificed for energy facilities has important bearing on mitigating actions required by the Comission. From a broader point of view, public concern exists for the tortoise in all of the western stat evidenced by the Federal listing of che tortoise in southwestern Utah in 1980, Questions have been raised as to the status of the tortoise in other areas, and these questions have been the principal motivation for this docunent. The question of present status unavoidably raises questions as to past status, Can we compare current situations with those existing in the past? It is a matter of record that populations of turtles may experience long term changes in abundance--resulting either from natural changes in habitats or owing to direct or indirect effects of human actions. Auffenberg and Iverson (1979) described a population of gopher tortoises (Gopherus polyphenus) in Lake County, Florida, whose density declined from about 1,900/sq. miles in 1960 to only 100/sq. miles in 1971. During this time, the vegetation of the area changed from an old field community to mature sand pine scrub, Stickel (1975) described changes in a Maryland population of box turtles (Terrapene carolina) between 1945 and 1955. Turtle numbers were apparently stable between 1945 and 1955, but the density of the population was reduced by half between 1965 and 1975. No 83-27causes for this change could be clearly identified. A Michigan population of painted turtles (Chrysemys picta) was reduced from around 981 to 186 individuals between 1954 and 1972, possibly because of increased predation resulting from a reduction in the number of safe basking sites (Wilbur, 1975). Unfortunately, few comparable data are available for the desert tortoise. Data from the pioneering work of Woodbury and Hardy (1948) on the Beaver Dam Slope in Utah between 1936 and 1945 can be compared with data collected between 1973 and 1981 by Coombs (1977a, 1977b), Minden (1980), and Minden and Keller (1981). However, there were substantial differences in the size and location of study areas and in the sampling methods (Appendix 2). Release of captive tortoises in and near the study sites in the last ten years has contributed to further confusion in interpretation and comparison of data from the samples. We have divided the data by site and investigator (Sites 32-35) and have discussed the differences in some detail in Chapter 12, One-time density estimates are available in the literature and unpublished reports for an additional 18 sites: Site 1, Argus (Berry 1974); Site 20, Chemehuevi Wash (Schneider 1980); Site 21, Pinto Basin (Barrow 1979); Site 28, Last Chance (Karl 1980); Site 29, Sheep Mountain (Karl 1979a); Site 30, Arden (Burge 1977: Site 31, Piute Valley (Karl 1979b); some southern Arizona sites (Schneider 1981), Site 42, Picacho Mountains (Schwartzman, n.d.); Site 43, Tiburon Island (Reyes Osorio and Bury 1982); and Sites 44 and 45 (Hohman and Ohnart 1980; Sheppard 1980, 1981). The estimates were nade using best professional judgnent, actual counts, and estimators such as the Lincoln Index, Stratified Lincoln Index (Overton 1971), Peterson 43-28method (Peterson 1896), Schnabel method (Davis and Winstead 1981), Schumacher~Eschmeyer method (Schumacher-Eschmeyer 1948, Davis and Winstead 1980), and Linear regressions (Hayne 1949, Marten 1970). The estimates from these sites are presented in the chapters for each state (Chapters 5, 8, 10, and 12. Densities of Desert Tortoises at Thirteen Sites in California We estimated densities of tortoises at 13 sit im California using data taken during spring censuses. These censuses were ordinarily based on about 60 person-days of effort (i.e., the work was carried out by one person over a period of about two months, or two or three people worked for shorter periods of time). For 12 sites, we used census data taken during a single year--either 1979, 1980, or 1981. For Fremont Valley (Site 2), data were available for both 1979 and 1981. Two census periods were established for each set of data, dividing the overall effort into two roughly equal periods of time. Tortoises observed and marked during the first of these time periods made up the first sample of the population (@- The second sample, composed of n tortoises, included some marked during the first census period (x) and others which were not marked during the first effort, With no further elaboration these data could be used to estimate spring density as recommended by Bailey (1952): ae at Diet) a Table 10 gives basic observations for the 14 sets of data used, together with population estimates computed using Equation (1). When recapture data (1.e., the second samples from each site) were examined in detail, we found that larger tortoises (those > 208-mm MCL) 43-29suopazaaz0> 8eyq NITA XOPUT TeuoTIz0do24 aya BuySN pentz9p 9rBN saIeMTISy “e oy SoT sot aca ws 09 ve AoW ~ 21 2H 086T oT 9% wt lz 06E 69 Let SET 9t Ae - 2 28H 6L6T wT ez se att o1z oO” zOT 96 oz Ae - 6 20H 66T eT oz ® ow oz «9 | 6S ST me ~ Oe aK OBST ot ot cot ove om OFT EZ WOT TT UE - Gz AH ORT ot st 2 ez 61 SOFT 26 me OT aR BET ot n 1 eet at Oke ST ARH OBST ot tu e ost nt m6 ST ae zt TH oneT ot ot OL ezt 8zt Te 9S cae at one - 17 adv TS6T oT 6 *9 vet wet eT ETE MOH 6 HORE ot “ eT ost ose 1ST ely 887 Te AW - € 28H 6L6T oe ” ort we soz 88) HT GOT. st Mr BH GLET rt € tot 92 a i a ee z 80T 667 667 es 69T s6 Te Aen - T 2H 6L6T oT Zz (errs “8/0 samme — (etya *bs/u) yo Aqtsuap ‘AQysuap a ae Be TeareIUT (@TTm *bs) paqeurisy peremtied = 7 u t sneua> TeqoL 2e2K eazy arts fe TR6T PUP 6z6T woonaaq eyurop Teg UE s9aye CT 3e Seszo2I03 aze89p 50 AITSUEP jo seawMTase exMadeDsx-emnadey “OT *TaPL 43-30were recaptured more readily than tortoises less than 180-mm MCL. If tortoises of different sizes are sampled with equal efficiency, one would expect-—on average--the size distribution of marked tortoises in second samples to match the distribution of marked tortoises at risk in the population. A hypothetical example, illustrating perfect correspondence, is given in Table 11. We can set up simple contingency tables to examine this relationship. Table 12 gives actual sampling data from Fremont Valley in 1961. Expected values are in parentheses. Total x", with 3 d.f., 48 about 11.8, with an associated p value of ~0.01. In this particular sample, it is clear that relatively more marked tortoises > 208-nm MCL were taken during the second census. Smaller tortoises were underrepresented. This pattern was characteristic of all 14 sets of sampling data. The situation reflected in Table 12 could arise in either of two ways. First, there could be a higher probability of recapturing a large tortoise once it had been marked. This would occur if field workers deliberately searched in areas where they had marked tortoises, expecting to find the same individuals again. This possible source of bias has never been carefully explored, However, we judge it more likely that sampling data of the sort pr ated in Table 12 indicate that a higher fraction of the subpopulation of tortoi: > 208-mm MCL was marked during the first census, while a smaller proportion of the subpopulation of small tortoises was marked. Data acquired at the Desert Tortoise Natural Area (Site 4) in 1979 and discussed by Shields (1980) are suggestive of this difficulty. One approach to this problem is to stratify the sampling data (t.e., group captures and recaptures of individuals in different size groups). A331Table 11. Hypothetical recaptures of tortoises of different sizes in the absence of a size effect on collecting efficiency. Tortoise Number of marked Marked tortoises size ranges tortoises at risk in second sample (am NCL) (warked in first sample) <100 a2 4 100-179 18 6 180-207 24 8 >208 36 2 Totals 90 30 83-32Table 12. Representation of marked tortoises of different sizes in the second sample of the population of Fremont Valley in 1981. Size Tortoise Marked tortoises Marked tortoises Total nunber of group size range in second sample not represented _— tortoises marked (am HCL) in second sample in first sample <1 100 4 (7.0) 8 (5.0) 2 2 100-179, 22 (28.2) 26 (19.8) 48 3 180-207 6 (6.5) 5 (4.5) n >4 208 52 (42.3) 20 (29.7) n Totals 86 59 143 43-33Schneider (1960) analyzed data from Chenehuevi Valley in several ways, including by means of the Stratified Lincoln Index (Overton, 1971). We applied this technique to the sampling data set forth in Table 10. Data pertaining to different size components of the population were segregated and used to estimate an overall population density. Consider, for example, sampling data as given in Table 13. The first step is to arrange the recapture data so as to allow for tortoise grouth from one size group to another between censuses (Table 14). The colums of Table 14 show recaptures in terms of original size groups. The rows allow for growth. Note that by the time the second census was taken, one of the two tortoises originally in size group 1 had grown into size group one tortoise in each of size groups 2 and 3 also grew into larger size groups, The remainder of the analysis involves the creation of a succession of matrices based on the numbers given in Table 14, One eventually ends up with a set of adjusted values of a, (see Table 13), which are used in connection with original values of n, and x, to estimate the abundance of ay each size group in the population. In this particular example, the adjusted values of a, are 6,39, 32.94, 16.39, and 41.28. The total population is then estimated as: fe Fany/ry @ or 295. Had ve estimated this population simply as G+ faen, + v/@r, +d @ we would have obtained 208. The 95 per cent confidence range endpoints for population estimates based on stratified samples are computed as: N (@r, +2) ¢2 (er, + 11/2) « Se A334Table 13. Hypothetical sampling data pertaining to four size groups of desert tortoises. Size group® Number marked Total taken in Number of marked in first census second census individuals in second census wo @ @ 1 a u 1 2 30 2 8 3 4 23 10 4 40 42 28 95. 104 ] a, Size groups are as in Table 43-35Table 14, Rearrangement of data in Table 13 to allow for growth during the time betweeh censuses. Size groups at time of Recaptures based on original size groups recapture” 1 2 3 4 1 1 ° ° ° 2 1 7 ° ° 3 ° L 9 0 4 ° ° L 27 a. Size groups are as in Table 43-36Population estimates (adjusted to a sq. mile basis) and confidence ranges based on stratified samples are given in Table 15. Discussion Most of the density estimates given in Table 15 have fairly broad confidence intervals. The lowest observed density (Stoddard Valley) differs significantly from nine of the other 12 sites, Densities in Ivanpah Valley and Chemehuevi Wash are gnificantly less than those reported for five of the other 11 sites, The three highest densities (che Desert Tortoise Natural Area, Goffs and Ward Valley) differ from five to six of the other 10 sites. Three sites (Frenont Valley, Kramer, and Chuckwalla Valley) with interaediate densities differ significantly only from Stoddard Valley. Luckenbach (1982) commented on the abundance of tortoises at two places in the western Mojave: the Desert Tortoise Natural Area (OTWA) and Minkley (near Barstow). Data from the DIVA were based on work by Marlow. These estimates were described as "density" in the text but as "relative density" in a table. The figur given are 927/sq. mile for the DINA and 562 /sq. mile for Hinkley. The estimates were derived from numbers of live tortoises observed in transects coubined with counts of active burrows. Without judging the reliability of these figures, it is certainly true that they far exceed other observations in the western Mojave and, in particular, at Sites 3 and 4 at the DINA. Barrow (1979) estimated the presence of around 75-80 tortoises/sq. mile in the Pinto Basin (Joshua Tree National Monument), but pointed out that not all small tortoises were captured. Future estimates of densities of desert tortoise populations will 43-37Table 15. Estimated densities of desert tortoises at 13 permanent study sites in California.* site Year Estimated density 952 confidence (a/sq. atle) snterval 2 1379 286 216 ~ 37% 1981 292 235 ~ 363 3 1979 353 272 ~ 460 4 179 22 252 - 339 7 1980 248 at ~ 362 9 1961 no 17 = 157 10 1980 21 162 - 328 u 1980 1st 95 - 274 1 1979 187 143 = 245 3 1980 BU 353 - 501 16 1980 a6 34a - 665 20 1979 187 aah ~ 226 23 1379 339 267 = 431 26 1980 249 161 - 383 a. Densities were computed using the Stratified Lincoln Index. A338require improvenents and or modifications of past procedures. The use of tortoise signs counted along transects probably can never be refined to the point vhere it can provide the sort of density data presently sought. More complex transect techniques have been described by Burnham, Anderson, and Laake (1980), which depend on counts of animals and the determination of a “detection function," expressing the probability of an animal being observed as a function of its distance from the line, The problem with this approach is that one rarely observes many live tortoises along transects, and the power of such a procedure is severely weakened when few animals are counted. Capture-recapture analysis is an attractive alternative because it requires neither measurenents of distances, nor derivations of detection functions. The problems lie in departures from standard ssumpt ions underlying capture-recapture models (e.g., Shields 198 Schneider 1980; Turner et al, 1962). The ost serious source of error in application of capture-recapture models is almost certainly unequal capture probabilities among various size classes of tortoises. Saal tortoises are simply not sampled with the sane efficiency as larger ones. The Stratified Lincoln Index {s designed to deal with this problea, tut ay create problems vith very low recapture frequencies. When no marked individuals in a stratum are recaptured, the estimated abundance of this stratum is taken as zero. On the other hand, if a very small number of marked individuals (e.g, 1) in a stratum is recaptured, the estimated abundance of this group will be large. This explains why the population estimates for Fremont Valley (Site 2), Stoddard Valley (Site 9) and Ivanpah Valley (Site 14) in Table 10 are larger than the population estimates given for these sites in Table 15. 3-39Wo juvenile tortoises were recaptured at these thr sites, the juvenile strata vere set equal to zero, and the total population estimates were reduced, Conversely, at Site 16 (Ward Valley) one juventle tortoise vas recaptured and the stratified index estimate (476/sq. mile) vas more than twice the proportional index estimate (216/sq. mile). These examples show how potently the capture-recapture experience of one size component of a population can affect the total estimate. In considering this problem, Schneider (1980) recomended estimating the density of only adult and subadult tortoises (>180-am MOL). He argued that "The snaller size classes are harder to census and inclusion of data [pertaining to] these tortoises compromises the accuracy [of the estimate]." We have presented density estimates based on all four strata (Table 15), but these figures must be carefully gauged in light of the foregoing problea. What do ve know of densities of other testudinids occupying arid environsents? Probably the closest analogs providing comparative data are other species of Gopherus in the southwest (G. ber: \dieri in Texas and G, Elavomarginatus in northern Mexico). Auffenberg and Weaver (1969) reported densities of G. berlandieri ranging from around 6,000 to over 30,000/sq. mile, but these figures apply to small areas. Preliminary surveys of G. flavomarginatus suggest densities of around 250/ sq. mile-- comparable to those of G. agassizi in some parts of California (Gary Adest, personal comunication). In more mesic environments in the southeast, G. polyphemus may sustain densities of 1,000 to 5,000/sq. mile in favorable habitats (Auffenberg and Iverson 1979). As gr cover diminishes, densities of thie species fall to some hundreds/sq. mile. As with G, berlandieri, che high abundances are observed only in the best 43-40areas and extrapolation to sq. mile extents may be misleading. CLUSTERING OF TORTOISE POPULATION DATA BY REGIONS ‘The tortoise population data were clustered by region within the geographic range of the species for analytical purposes. A description of the regions is presented, followed by a discussion of the assignment of study sites to specific regions. A Description of Regions within the Geographic Range of the Desert Tortoise Desert tortoises are found in two major deserts of the southwestern United States and Mexico: the Mojave and the Sonoran. These two deserts include several ecosystems, which can be defined on the basis of vegetation, topography, elevation, and faunal composition. The regions are described by state for convenience. Cali fornis Berry prepared a map and a brief discussion of ecosystems for desert areas in California for the BLM (U.S. Dept. of Interior 1980, 1981). The map delineations are used here. The term “ecosyste describes geographic regions of similar climate, elevation, topographic features, and plant and animal communities. The source materials for defining the boundaries of ecosystems were primarily unpublished data gathered by the wildlife, botanical, and soil staffs of the California Desert Program, Bureau of Land Management, between 1974 and 1980. They included: (1) maps of distributions of vertebrate species based on thousands of new distributional records gathered between 1973 to 1980; (2) Landsat images with 28 color-interpreted special classes at a scale ASMLof 1:250,000; (3) a California desert Landsat color composite mosaic at a scale of 1:500,000; (4) interpretations of generalized vegetation and floristic regions in the California deserts; (5) soil and landform map (6) wildlife habitat maps for Saline Valley, El Paso, Red Mountain, East Mojave, Yuha, and Santa Rosa Mountain planning areas; (7) vegetation maps of Death Valley and Joshua Tree National Monuments; (8) range maps of various plant species shown in Benson and Darrow (1954); and (9) information based on the personal experience of the wildlife staff at the California Desert Program office. ‘The Mojave Desert in California was subdivided into four ecosystems: (1) northern, (2) western, (3) central, and (4) eastern (U.S. Dept. of Interior 1981). The Sonoran Desert in California (or the Colorado Desert) was further split into six subdivisions: che Coachella Valley, north- central Colorado Desert, northeastern Colorado Desert, southern Colorado Desert, West Mesa, and East Mesa. We identified seven of these "ecosystems" as relevant to tortoises in California: northern Mojave, western Mojave, central Mojave, eastern Mojave, north-central Colorado, northeastern Colorado, and southern Colorado deserts. Each ecosystem is described briefly below (Pig. 1). Northern Mojave Desert—This region is part of the Basin and Range province and has a number of enclosed valleys, some of which are near or below si level (U.S. Dept. of Interior 1981), The valleys are surrounded by mountains rising to 7,000 to 11,000 ft. Rainfall occurs primarily in Late fall and winter. Vegetation ranges from creosote bush (Larr tridentata) at the lover elevations to sagebrush (Artemisia sp.) and bristlecone pine (Pinus longaeva) forests in the mountain ranges. A342Tortoises reach thetr northernmost distribution in California and parts of Nevada in this ecosystem. Western Mojave Desert: elevations of 2,000 to 3,500 ft (U.S. Dept. of Interior 1981), There is This is a region of broad valleys and fans at Little topographic relief, with only occasional low hills and mountains which do not exceed 5,000 ft. Rainfall occurs primarily in winter. Vegetation is typically creosote bush scrub, Joshua tree (Yucca brevifolia) woodland, and alkali sink (Atriplex spp.) types. The desert tortoise is a common and dominant species (dominant in terms of reptilian biomass). Ceatral Moj. Desert--This region extends from the southern boundaries of the Naval Weapons Center's Randsburg Wash and Mojave B test ranges and the ‘Amargosa Gorge south to the Barstow area (U.S. Dept. of Interior 1981). Elevations range from 2,000 to 5,000 ft. ‘The desert here is generally drier and of lover productivity than areas to the north, west, and east. Rainfall comes during winter. Vegetation is typically creosote bush scrub and alkali sink, often with windblown sand. Tortoise populations are much lower in density and more scattered than in the western Mojave Desert (Chapter 2). Eastern Mojave in the north to the Fenner Valley and Goffs areas in the south (U.S. Dept. -~This ecosystem extends from the Nopah Mountains of Interior 1961). It has several major valley and mountain systems, with elevations ranging from 3,000 to 7,000 ft. Over 200 water sources are present, most of which are in mountainous terrain. Rains occur in winter and summer. Vegetation at lower elevations is the typical creosote bush scrub and Joshua tree woodland communiti » but with more perennial grasses, cacti, and shrub-filled washes than in the western and central AaaMojave De: rt. Tortoise populations reach the northern limits of their geographic range in the northern part of this region; three moderate to high-density populations occur in Ivanpah, Piute, and Fenner Valleys further to the south (Chapter 2). jentral Colorado Desert-~This system extends from the south- eastern edge of the Newberry Mountains and the vicinity of Baker south and east through Joshua Tree National Monument to the Chuckwalla Valley and Colorado River (U.S. Dept. of Interior 1981). The region is one of the lower, drier, hotter, and les productive desert areas in California, Elevations range from 1,000 to 4,000 ft. The topography is characterized by broad desert basins and valleys with numerous playas edged by windblown sands, Tortoise populations are low in density and sparsely distributed (Chapter 2). Northe: rn Colorado Desert--This region contains broad valleys and stretches from Fenner Valley in the north through Ward and Chenehuevi Valleys to Vidal Valley in the south (U.S. Dept. of Interior 1981). The creosote bush scrub vegetation contains typical elements of the Sonoran or Colorado Deser: ocotillo (Fouguieria splendens) and teddy bear cholla (Opuntia bigelovii), with smoke tree (Dalea spinosa), palo verde (Cercidiun floridum), and catclaw (Acacia greggii) in the vashes, Elevations range from 1,000 to 1,500 ft along the Colorado River and adjoining valleys, and rise to 3,000 to 4,000 ft im the Turtle, Old Woman, and other mountain ranges. Rainfall occurs in both winter and sumer. Tortoti populations range from moderate to high densities in the valleys, becoming lower in density to the south. Southern Colorado Desert—This is a heterogeneous region with valleys, A345,‘expansive drainages, and ashes interspersed with rocky mountains and areas of desert pavement. Vegetation is creosote bush scrub, often with ocotillo and brittle bush (Encelia farinosa). Washes often contain dense microphyll woodlands with ironwood (Olneya tesot: palo verde, suoke tree, and mesquite (Prosopis sp.). Elevations range from 300 ft along the colorado River to 4,800 ft in the Little San Bernardino Mountains. Rainfall occure both in winter and summer. Tortoise populations are of low to moderate densities with a few small areas of high densities. Nevada. Most tortoise habitat in southern Nevada is in terrain and vegetation types similar to the eastern Mojave Desert of California. Some areas, particularly those north of Las Vegas, may be more typical of the northern Mojave or Great Basin deserts (U.S. Dept. of Interior 1981). Arizona. Tortotse populations occur both in the Mojave and Sonoran deserts. Populations in the Mojave Desert are limited to areas north of the Grand Canyon and occur in habitats similar to the eastern Mojave Desert of California. Mojave Desert habitats also occur near Kingman. Elsewhere in Arizona, populations of tortoises are found primarily in Arizona Upland communities of the Colorado Desert (Chapter 10). vee Tortoise habitat is confined to the extreme northeastern tip of the Mojave Desert (Woodbury and Hardy 1948). This area is similar to the eastern Mojave Desert described for California. ASSIGNMENT OF STUDY SITES TO REGIONS ‘The study sites occur in nine ecosystems, of which four are in the Mojave Desert and five in the Colorado Desert (Table 16). Three of these ‘ecosystems had few or no tortoises on the study plots. The northern Mojave 3-469 or oo TxeH ‘339890 uexouos ooyKan ‘a398eq uez0U0s ae ‘oe ty-96 euozy3y ‘az980q uesoN0S euoztay ‘33088 uezou0s yo puozyay ‘adots meq seanag uoztay ‘dots meq 19A09q AT pue se TT wean ‘111 ‘11 wean ‘edoTs weg soneog ze 1 wean 1 wean ‘edots meq z9avoq re-ez penen epenay ‘220800 2AefoH we-te uz *90 *€e-12 ax98aq Wanos 4@89q opPz0ToD u394I0S oz ‘or 3398aq operoqop wi93Se9y3I0N Sz tye ‘6TH-ut qx989q opexoToD TeII99H330N oz-et Oyby ‘oE~6z *ST-ET eaefoa asa 33980q eAefoH us02505 ze ‘aa980q aavfow Te22U29 ez ‘Tt ax080q eACfoW Ur8yI20N Trt 1-6 tit aaefon 260K, azeReg aaefog uxersey 1 ‘83a uy wnoys: suoy8a1 03 sn9364609 UT serys jo quseusyssy Suyazans90 $9375 ‘ameu woyfoy ueu m926ks00g *euoySex pur sue3sisova 03 se2zs 2Wan-TTaUs UaAds TTP UT Sasyors02 aAey SIoTd TT 3ON “4 ‘sesyoai01 uye2399 307 SePITS Jo YET SF Sas9"T> swaK-TTeYS YaTA S9TOUedoI087p 0 voseDI TedyoUTId ‘UL “stoqunu [e103 uy saqauedas9sTp puTy Kem OTA aueE a2 GOI] “D1 SASEETD azTS sMOTIeA ‘ua uF 898702203 10; Saqenu YIqA $e98eTD LeaR-TTaYs UF sasyors02 30} szequnu SuyIedN09 s1epeay “e o's ty tet set zy ‘doy peBiey ty sty over zy teaang 32eTE OF > 0861 ay txeanozeH 6 Lists ty tet tt set av ‘yen drys 913377 e ots ty tet 861 2v ‘upeaunoy 3098 LE Lig ts ty ett gst ZW ‘TUTH oety of ots ty fete tT 6tst aw ‘AerTeA 8amFa TE Lists ty tet tr auyseym 2808 SL6T-7L6T aN ‘upy of L198 's ty te te 6u6t Am “peaunoy dasys 62 oisty tt gst AX ‘e0ugo 2881 oz ois ty te tet o86t Yo ‘11 Kortea erremeny 92 sty tetz acer vo 'r Aertea erremyonn 92 Listy tesa tT 616 *BL6T yo ‘woueg eTTemxoMy) — £2 ost tee Let Vo ‘#Buyads poonuoaz0y zz ots teeta tt sist Vo ‘oyerg oma 1 (620118 30 s9qunu) quatox8 vo etqetrene oze Fey Suysspa arqettene. sou WoT 103 sosseTD avon-TTaYs 30 38F7 ase saprs s3eox ze sopyTs savan eave 2255 oats spamuzauos ‘sxo1d Apnae Tf ao3y OTgeTTVAP PaeP reAR-TTaYe aeTo3z0R, “1 oraer rryear was used. here gore than one year of data existed for an individual, the information for each year was treated separately. Changes in Shell-vear Classes. Tortoises (n = 508) on nine study plots were first photographed in 1977, 1978, or 1979 and then photographed again one to three years later. These tortoises were assigned shell-vear classes using the 35am slides. The following data were recorded for each tortoise: identification number, any change in carapace leagth and shell-wear class, and the time period over which the change occurred. If a tortoise showed a change in shell-vear class, the slides for each year vere compared to determine the nature of changes. Relationship of Bone and Scute Thickness and Configuration and Shell-wear Class. Carcasses of adult tortoises were examined (1) to determine the thickness and configuration of bones and scutes of the carapace, and (2) to compare these features in tortoises at various stages of shell wear, Carcasses were selected from the collection of over 1,500 shell-skeletal renains for study plots in Arizona, California, and Nevada (Table 1). Shells of several adult males and fenales representing four of the shell-vear classes were sawed in half with a six-inch, circular hard-board saw blade vith fine ceeth near the junction of the fifth and sections of the carapa sixth marginals to prepare cro RESULTS Definitions of Shell-vear Cli Distinct changes occur in tortoise shells sociated with age and abrasion, Changes are most readily apparent in growth rings, hatchling plates, and in the relief of scutes on the carapace. The changes are often AGBsubele and gradual. We divided shell vear into seven classes outlined in the key (page Ad-23) and further described below. We emphasize that these classes are artificial. Shell wear is continual, and distinctions between advanced stages of one class and early stages of an older class may not be clear-cut. In the ensuing definitions and figures, descriptions of shell wear are confined to changes in costal and vertebral scutes. Marginals were not included in the evaluation because predator damage is comon to these scutes, The parts of the cortoise used in sheli-vear class descriptions are shown in Fig. 1. Terms for the scutes are taken from Woodbury and Hardy (1948) and Loveridge and Williams (1957) as described in Zangerl (1969). Glass 1. Ar least 95 per cent of each scute 1s composed of the areolae, Granulations are distinct on areolae (Fig. 2a). This is the only class with a size limitation and includes tortoises up to 60-mn MCL. Glas Areolae begin to wear smooth. In general, the first scutes to show such wear are the first vertebral and first costals (Fig, 2b). The total amount worn is equivalent or less than the area of 1.5 areolae. For example, in advanced stages of Cl 2, either up to 75 per cent of the areola of any one scute (usually the first vertebral) and 25 per cent of the areola of one to three additional scutes (usually the first costals) may be worn smooth or up to 50 per cent of the areolae of three scutes may be worn smooth (Fig. 2b). Class 3. Even more hatchling plates show wear, generally from anterior to posterior. The middle stage of Cla; 3. is show in Fig. 3 and the advanced stage in Fig. 4. Areolae of all scuté may be worn smooth except for an area equal to 1.5 areolae, e.g., (1) 100 per cent of the Aun9or areola second right costal scute growth rings. third right costal scute fifth vertebral scute Fig. 1. A drawing of the desert tortoise showing (a) the areola of hatchling plate; (b) growth rings or annuli; (c) first, second, third, and fourth costal scutes; and (@) first, second, thrid, fourth, and fifth vertebral scutes. Ab-10desert tortoise.Dp (ie = Ne Ab-13,areola of at least one scute and 25 per cent of the areolae of at least two other scutes, or (2) all but 50 per cent of the areolae of three scutes Typically, the last to wear are the fourth costals and fifth vertebral. Growth rings may be unvorn or are just beginning to wear, usually on the centers of the first costals and vertebrals. Class 4, All areolae are worn snooth, except for an area equivalent to less than 1.5 areolae, If areolae on the fourth costale are depressed and generally unvorn, the equivalent area remaining rises to a maximum of 2.5 areolae. In most tortoises, chipping and flaking of growth rings is Limited and occurs primarily on the older, inner rings of each scute. Sone tortoises have chipping and flaking in some outer rings, but still have growth rings typical of a Cl 24, Less than 50 per cent of @ line across the width or length of a scute overlies sncothly worn areas which no longer show evidence of growth rings, However, if one scute 1s highly domed, then the saooth area may underlie 70 per cent of the line, The scutes most Likely to show wear are the second and third costals and vertebrals. Figs. 5 and 6 show middle and late stages of this class. 5, Flaking and chipping extend to outer growth rings; worn clas areas are expanding on scutes and are partially, if not entirely, the result of flaking and chipping of thin layers of laminae (Fig. 7). On two or more scutes, a smooth area underlies more than 50 per cent of a line across the width or length of the scutes, Wear on growth rings follows a pattern similar to that of the areclae: from anterior to posterior. The first, second, and third costals and vertebrals show the most wear, whereas the fourth costals and fifth vertebral show the least. Class 5 tortoiss still have slightly convex scutes and underlying Abasbone so that each scute is slightly arched or almost so (Fig. 8a). No "sinking" or depressions are evident. With "sinking," both acutes and underlying bone become depressed (Figs. 8b, 8c). Sinking appears to Javolve physiological changes in both bone and scutes. The underlying bone not only becomes depressed, but thins, often in patches, and is more porous, Where scutes join (the seams), the underlying bone is thicker, often more so than in earlier shell-wear classi Carapacial bones of adult females appear thinner and more porous than in male Glass 6. Chipping and flaking of growth rings often has expanded to peeling, which results in loss of thick layers of laminae over areas of 1.0 Growth rings may not be very worn (Fig. 9). Scutes are depressed to some extent. Depressions are present on eight or fewer scutes, usually appearing first on the lateral edges of the second and third vertebrals and on the upper portions of the costals where they meet the vertebrals, Spaces above the seams may appear to be widening and scute edges thickening (Fig. 8b). Both bone and scutes are no longer slightly convex or rounded in cross-section. Instead, the vertebral and costal scutes and underlying bone appear slightly flattened, particularly at scute centers. Scute material in the depressed areas is often thinner than in adjacent are: Bone may exhibit a slight thinning under scute centers as well as a slight thickening under scute seams. Glass 7. Scutes and bone are quite sunken and the space above the seams generally appear wide with scute edges thickened (Figs. &c and 10). Sinking is present on nine or more scutes. Most growth rings are worn smooth; those remaining are usually on posterior scutes and in depressions on the anterior and posterior portions of the second and third vertebral Ab-18a yscute bone / scute Fig. 8. Cross-sections of the carapace of the desert tortoise with (a) shell-wear Classes 4 or 5; (b) shell-wear Class 6; and (c) shell-vear Class 7. A419depressed area Fig. 9. A typical shell-vear Class 6 for the desert tortoise. The shaded areas denote depressed areas. of scute and bone, AG-20depressed area Fig. 10. A typical shell-wear Class 7 for the desert tortoise. The shaded areas denote depressed areas of scute and bone. 4-21scutes. In early stages of Class 7, sinking is apparent on the first, second, and third costals and second, third, and fourth vertebrals. In advanced stages, the first and last vertebrals become depressed and centrally the scute material very thin (Fig. 8c), Patches of bone may show through scutes. One possible explanation for the thinning of scute centers and the thickening of scute edges in Classes 6 and 7 is stress on the bonds between laninal layers caused by gradual sinking of underlying bone. in Classes 1 through 5, scutes and underlying carapacial bones are generally convex. With the gradual flattening and development of concave areas in the underlying bone, the inflexible laminae are pulled dowavard with the bone, weakening laminal bonds. Macroscopic air spaces or lightly-colored “pubbles" sometimes appear between laminal layers. The bubbles break open, then chip and flake. This action may cause the thinning of scute centers. scute material may remain thick near the seams because the underlying bone is relatively static here, Further deposition of laninal material may continue to build these edges As bone underlying scute centers continues to sink, laminae in the center may be further weakened and peel in large pieces. The result is the paper-thin scutes of the advanced Class 7 tortoises. ‘The apparent slowness of wear on the fourth costals and fifth vertebral scut may be explained by changes in relief of scutes throughout the life of the tortoise, The fourth costals typically are the least convex scutes on a young tortoise; they even may be slightly concave. Their flatness may reduce the relative potential for abrasion, As the tortoise ages, the fourth costals and fifth vertebrals may become 4-22depressed, but the relative change to the underlying bone is not as great a5 the change experienced by other costal and vertebral scutes and underlying bone. The fifth vertebral scute usually remains the most convex scute on the carapace throughout the life of the tortoise, especially in males, and the Last to wear in sone tortoises. Changes in the marginals from Classes 1 through 7 are worthy of note also and are show in Figs. 2 to 7 and 9 to 10, The marginals gradually wear avay on the edges and scute material is lost. Changes in the nuchal are of 23 particular interest; the nuchal extends beyond the marginals in Clas: and 4, but gradually wears avay and becones even with the other anterior marginals, The wearing avay and reduction of the nuchal may affect carapace length measurenents, especially in Classes 6 and 7 individuals. This is discussed under "Relationship of Tortoise Size to Shell-wear Class" below, The following key was used in assigning shell-wear classes. Key to Shell-wear Classes for Desert Tortoises! A <60-mm MCL; 95 per cent of each nonmarginal scute composed of areola. Granulations on areola distinct. Class 1 AA Not as above; > 60-mm MCL. 3 B Total amount of areola worn smooth is less than or equivalent to 1.5 areolae. For example, up to 75 per cent of one areola and 25 per cent of three others may be worn smooth, or 50 per cent of three areolae is worn smooth. Class 2 BB Not as above, or, an area equivalent to more than 1.5 areolae is worn smooth. c 1, Scutes mentioned here are the nonmarginal scutes of the carapace costal and vertebral scutes. Growth annuli are on these same scutes. Ab-23© An area up to or equivalent to 11.5 areolae is worn away. An area equivalent to at least 1,5 areolae must remain unworn. Growth rings are generally unworn. Class 3 CC Granulations on areolae mostly, if not entirely, worn smooth. An area equivalent to 1.5 areolae may remain, However, if areolae on the fourth costals are depressed and generally unvorn, a maximum equivalent of 2.5 areolae may remain. > D Chipping and flaking of growth rings are Limited and occur primarily on inner rings adjacent to areolae. Growth rings are distinct and in good condition otherwise. Less than 50 per cent of a line across the width or length of scutes overlies smooth worn areas. A smooth area may underlie 70 per cent of the Line of one scute. Class 4 DD Flaking and chipping extend to outer growth rings. On two or more scutes, the smooth area underlies more than 50 per cent of a line across the width or length of the scutes. E E Depressions present on scute surfaces. F EE Depressions not present on scute surfaces. Class 5 F Depressions present on eight or fewer scutes, primarily on the second and third costals and second and third vertebrals; generally patchy in distribution. Class 6 FF Depressions present on nine or more scutes and generally much more uniform, often involving all but the edges of entire scutes. The Relationship of Shell-wear to Age, Size, and Growth Evaluation of Tortoises of Known Age and Shell-wear Class, Information on tortoises of known age and shell-vear class was divided into two group: () data for tortot: s under 35 years of age, and (2) data for those over 35 years. A246For the group under 35 years, definitive data are available on growth but not on shell-vear class for 21 wild tortoises living in fenced plots at Rock Valley, Nevada since 1963 (Medica, Bury, and Turner 1975). Eighteen of these tortoises were young and small (52~ to 110-am MCL) and probably in shell-wear Classes 1 and 2 when first encountered (Medica, personal communication), Nineteen years later in 1982, they ranged from 156- to >214-mm MCL; Medica (personal communication) suspects that their wear classes are 4 or less, When these tortoises are re-captured in 1983 and their shell-wear classes are recorded for the first time, a significant new body of data will be available for wild individuals. Numerous owners of captive tortoises have observed the sequential changes in shell-wear patterns described for Classes 1, 2, and 3 in individuals raised from eggs to adulthood and kept in captivity: from 10 to 30 years (personal communications from Bayard Brattstrom, Betty Burge, Robert Patterson, Glenn Stewart, and Martha Young). Ne refer here only to captives with "normal" carapaces, not those with the "knobby" appearance associated with rapid growth, early maturity, a "stimulating environment," and “an excellent dietary regimen" (Jackson et al., 1976, 1978). For tortoises 35 years and older, we have data for one tortoise that has remained in the wild for many years, four wild tortoises taken as captives and kept in captivity for decades, and one tortoise hatched from an egg in captivity. Our age estimates are based on data presented on size, age, and growth rates for captive and wild tortoises in Patterson (1972), Patterson and Brattstrom (1972), and Medica, Bury, and Turner (1975). Tortoise No, 1. Female Bj (Woodbury and Hardy 1948) was captured initially on January 22, 1944. She measured 215-mm MCL. When recaptured Ab-25in November, 1968--22 years later--she measured 218-mm MCL was a Class 7 (Hardy slide 7454; Hardy, personal communication), If she was 20 to 30 years old in 1944 (an estimate biased toward youth and based on size), she may have been 45 to 55 years old as a Class 7 in 1969. If, however, she was considerably older in 1944 and a Class 4 or 5, then she may have been well over 60 years old. After discussions with Dr. Hardy, we surmised that the group of tortoises studied between 1936 and 1946 included few Class 6 or 7 tortoises. Woodbury and Hardy (1948) described only one tortoise, a female (Ce) with "sunken areas" on the first, second, and third vertebral scutes. Female Bj was probably not a Class 6 or 7 in 1944 because she vas not mentioned. Tortoise No. 2, A freeze-dried female specimen (338-mm MCL) in the San Bernardino County Museum, California. Notes on the museum tag (No. 2649) indicate that the tortoise was about the same size in 1931 when acquired by Bering Monroe. This tortoise died on February 18, 1981 at an advanced Class 4. Judging from the very large size, it was probably a minimum of 30 to 40 years old in 1931, and could have been 80 to 90 years old or more when it died. Tortoise No. 3. A 230-mm MCL male collected near Victorville, California in 1928 (Jennings 1981). No clear photographs showing shell wear are available until 1948. From 1948 through the mid-1960's, the shell-wear vas Class 5 (Jennings, photographs, drawings, and personal comunication). The 1966 and 1967 photographs indicate smoothing of and loss of growth rings and an advanced wear Class 5. No photos are available from 1968 to 1982. In 1982, after 54 years in captivity, the tortoise Ah-26measured 294-mm NCL and was an early Class 7. We suggest it might have been at least 25 to 30 years old in 1928, based on the 230-mm NCL adult size, and 75 co 80 in 1962. Tortoise No. 4. A 298-nm MCL captive fenale, Roxie (Klein 1977; Mary Trotter, personal communication), Roxie has been kept in captivity since 1920; she became the pet of Klein (1977) in 1936. Klein (1977) reported that she was an adult in 1920 and approximately 298-am MCL in 1936. In 1982, she was an early Class 4 (Mary Trotter, personal communication) and has been adult for at least 62 years. Tortoise No. 5. A 254-mm MCL female kept in captivity since 1925 in Pasadena and Thousand Oaks, California (Rosemary Sage, personal communication), This animal was collected by Arthur Vierick from the vicinity of Kelso, San Bernardino County, in about 1920, Sage reports that the tortoise has had a smooth shell with no evidence of growth rings (at least a Cl 5) since 1925 and has not changed appreciably in Length over a 57-year period, Photographs taken in 1977 and 1982 indicate that the tortoise is in the early stages of Class 6. Tortoise No. 6, A 30l-mm NCL sale, born in captivity in 1904, This tortoise vas kept in a backyard with its om burrow in San Bernardino until about 1960, when its omers moved to Lancaster, California in the Mojave Desert. ‘The tortoise was again kept in a backyard until about five years ago, when the owners moved to a mobile home park. The tortoise then lived in a “little house" instead of a burrow, In summer 1981, ommership was transferred to Leo and Gloria Nowak of Inyokern, California. This male is an advanced Class 4 at 78 years of age. Relationship of Tortoise Size to Shell-vear Class. The size ranges of Ab-27tortoises at 31 permanent study plots in the seven shell-wear classes are shown in Table 2, Only the first record of size and shell-vear class was used for each tortoise. Only shell-wear Class 1 was defined by size (< 60-mm MCL). A wide range of sizes exist for tortoises in shell-vear Classes 2, 3, and 4, from immature through adult. Where sex was determined, mean sizes of males and females increase directly with increasing shell-vear class (Fig. 11). For the undetermined group (Classes 1 through 4), the pattern is the same. Note that the sample size is very small for Classes 4 and 5 in the undetermined category (Table 2). Most tortoises in shell-vear Class 4 and all in Classes 5, 6, and 7 are considered sexually mature. Minimum size at sexual maturity for females appears to be about 197-mm MCL, judging from the number of females at or near this length with old, worn carapaces almost devoid of growth rings. Using the same criteria, the minimm size at sexual maturity for males is also 197-am MCL. These are smaller sizes than reported by Woodbury and Hardy (1948), Patterson and Brattstrom (1972), and Auffenberg and Iverson (1979), and smaller than Berry and Turner use elsewhere in this volume for estimating adult sizes (Appendix 3). If shell wear on marginals (particularly the nuchal) is taken into account, the 197-um MCL tortoises in Classes 6 and 7 would be considered a few millimeters longer. ‘The wide range of sizes in shell-wear Classes 2, 3, and 4 appears to be related to variation in growth rates. Some individuals grow rapidly from juvenile and immature stag to adulthood, whereas others grow Tortoise Slovly. Examples of extremes in slow-growing animals art No. 178 (Class 2), which grew only 8 mm, from 60- to 68-mm MCL, in 33 months; and Tortoise No. 118 (Class 4), which grew 19 mm, from 132- to Ab-28*sesyorioa a398ap jo syaBuey aoedese uvaw pur sasseT> IWaM-TIONS “IT “Sta SSP|9 12@M-|]OUS L 9 s ’ e 1 +1 ——___1—__ Peulwieyepun OS — ae eyewe} eeu os ost o0z osz ooe (ww) yj6ue] soedeseo ue A293Table 2. Shell-vear cli es and carapace lengths of tortoises. Observed carapace lengths (am) She11-vear 2 standard errors clase No. Sex Mean Range of means 1 128 undet. 52.4 41-59 0.8 2 850 undet. «11.0 60-177 2a 168 179.0 117-246 3.3 moo 168.5 96-231 3.6 3 63 under. © 146.2 93-178 4a ae 201.9 133-268 2.8 7 ot 199.0 115-248 28 4 3 under. 160.0 133-175 27.0 298 236.0 148-307 3.0 m3 219.5 142-267 2.6 5 2 under. 151.5 133-170 37.0 252 258.8 155-310 29 2638 229.5 176-293 2a 6 19 270.8 3 m of 235.8 193366 2.8 7 a8 272.3 235-330 6.0 2 o£ 245.9 209-290 9.3 4, These low carapace lengths appear to be the result of wear on the nuchal and pygal scutes. The 193-mm MCL is from a single female with an abnormally small nuchal (Site 15, Tortoise No. 426, April, 1977). This same female measured 197-an MCL in 1978 and 1980, and would have an even longer carapace length if the abnormality were taken into account. Ab-3015i-mm MCL, in 36 months and then remained at 151-mm MCL for an additional 24 months. An example of a more rapidly growing animal is fenale Tortoise No. 13 (Class 2), which gret 82 am, from 165- to 247-mm MCL in 60 months. ‘These examples are taken from the same study area (Site 2 in Fremont Valley, california), over the same general time period. The average growth rate for young tortoises in southern Nevada was about 9 um per year (Medica, Bury, and Turner 1975). ‘Thus, some large tortoises may be young and have low shell-vear classes, and some small tortoises may be old and may have relatively high shell-vear classes for their sizes. A corollary is that body size is not necessarily an indicator of age, even at the immature sizes. Patterson (1972) also found considerable variability in size with age in a study of growth in six captive desert tortoises over a thirty-year period. Gibbons (1976) has reviewed the size and age issue for reptiles and reported similar findings. Relationship of Shell-vear Class to Growth. The frequency of tortoises showing evidence of growth in each shell-vear class is presented in Table 3. Wich the exception of shell-vear Class 1, the frequency of tortoises showing evidence of grovth decreased as shell-vear class increased (Fig. 12). No tortoises in Class 7 shoved signs of grovth. Frequencies vere similar for Classes 1, 2, and 3 (80 to 86 per cent). When frequencies of tortoises showing signs of growth were compared by shell-vear class for the four spring months, some interesting patterns energed (Fig. 13). Samples of tortoises from shell-vear Classes 1, 2, and 3 have almost identical patterns and have high frequencies of growing tortoises in April, May, and June (78 to 100 per cent), whereas Classes 4, 5, 6, and 7 are each distinct and exhibit lover frequencies in a sequential, step-wise manner. ‘Ab=31*eaep sseT> ran-TToys aTqeTTEAe aya pue srOTd APMIS Jo ISTT B 305 T PTARL 295 ‘saqdues [TEMS YIFA s20Td BBoy2 ATTwFoadse ‘ssvTD AvaR—TTOYS YDeE UT Saszo1z02 eAry saoTd Kpnas TTe ION *e 0-0 6 a t on wer 8 ese 29 6z s 99 509 oc , 7-08 sz uw € 6-98 sett 82 z 6-08 wt mw 1 Wanoa 30 suze Suynous eeaep wa ‘eset ‘ature 30 309 304 son soars Kpnas Jo 20qmN awan-Trous +sseT> 1PaN-TToYs Y>ee UT YanorB Jo aouEPTAS SupAoYS ysfo2203 yo Kouenbarg “¢ eTaeL Au-32ssassea 28aA-TTOys UOADE Jo Uae UF yaMOI# Jo sU;S BUTAOYE saszor202 jo Kousnboag SSB|D J2OM-|]aUS L 9 3 v & z can “ra on oz oe ‘OF 0s OL og 06 01 (%) YIMOJH Bulmoys sesiojio, yo Aouanbes4 A433Frequency of tortoises showing growth (%) 100 90 80: 70 50 40. 30: 20 10. Fig. 13. Shel-wear classes PoL2on4 March April May June Frequency of tortoises showing growth in shell-wear Classes 1, 2, 3, 4, 5, and 6 in spring, Ab=34A preliminary examination was made of the amount of growth manifested by tortoises in each shell-vear class. This examination was based solely on changes ia length in recaptured animals over one~ to five-year periods. The pattern appeared similar to that described above for frequencies of tortoises showing evidence of grovth. This analysis will be completed in 1983. Tortoises in wear Classes 6 and 7 shoved Little evidence of growth. ‘Those in Class 6 (n= 148) showed a mean incr se of 1.1 um (range: -5 to #11 ma), whereas those in Class 7 (a = 21) showed a 0.18 mm increase (range: -7 to +3). Forty-six per cent of those in Class 6 and 50 per cent in Class 7 showed no or negative growth, respectively. Tortoises in these two classes may be experiencing shrinkage due to changes in bone structure, as vell as thinning and wear on the nuchal and pygal scutes. Changes in Shell-vear Classes. Data on changes in shell-vear class from nine study plots are shown in Table 4, Tortoises exhibiting changes im shell-vear class are compared with individuals showing ao such change. ‘The sample size of Class 1 is very small (n = 1), because of the difficulty in recapturing these snall aninals. Summarized data for tortoises advancing from one shell-vear class to another are shown in Table 5. More tortoises (19.8 per cent) moved from Class 2 to 3 than for any other class. However, we expect Clase 1 to have an even higher rate—-S0 per cent or nore--vhen more data are assembled. Class 4, 5, and 6 each have a very low proportion of tortoises changing wear class—-approxinately 1 per cent. These three classes are probably of considerably longer duration than Classes 1, 2, and 3, as inferred from the lower turnover rates. Changes in Bone and the Relationship of Shell-vear Classes. A number AG=35oe ros : ey . of eters on tenet cone ote Ab-36vt T 6 Log 60 T 60t gors 60 7 ot sole er bee 2% yore set wv we eoz ‘eueys Suyouazredxa SETS IvOM-TTAYS TeUTsTIO sevT> zvan-TTays sets Up saeyoaz0] yo quao aeq ‘Sup8ueys saszor103 Jo “oN 278 oTdues aeen-TTays UT e8uEID *(y 9TaeE 988) wyUI0; FTE UF s207d Apnas ouzu 103 paztiemns se sastorica paindese1 203 S9ssUTD I¥OR-TTOYS UE seBuEYD “¢ OTAEL Ab-37of changes occur in bones of the carapace between hatchling and adulthood and appear to be related both to size and to shell-vear class. Tortoises in shell-vear Class 1 have incompletely ossified skeletons and large fontanelles. By the time tortoises have been in shell-vear Class 2 for three to seven years and have reached 100- to 110-am MCL, their skeletons have ossified alnost completely. Growth in length and thickening of the bony skeleton typically continue until and extend beyond the time the tortoise reaches xual maturity, Tortoises in Classes 2, 3, and 4 are in chis category regardless of size. When tortoises have reached shell-wear Class 5 and for the time they remain in this class, they probably are at the peak of bone development (Fig. 8a). Tortoises in shell-vear Classes 6 and 7 show characteristics typical of aged vertebrates: little or no growth, deterioration and thinning of bone, and increased porosity of bone. Bones of fenales appeared to be thinner and aore porous than those of ales. DIscusstON The changes in shell-vear class that occur over the Life of the tortoise are a real phenovena and are a part of the growing and aging processes. Wear classes, in tura, appear to be affected by diet and possibly sex, but probably not terrain and soil texture, These factors interact in complex ways that are not yet fully understood. Our discussion is divided into three parts: (1) the changes in shell-wear patterns and 1, The shells cut in cross-section for this analysis were stolen from the Bureau of Land Management office in Riverside before measurements and analysis were completed. Ab-38sequence of changes, (2) chronological age of tortoises and the relationship to shell-wear classes, and (3) possible factors affecting duration of shell-vear classes Changes and Sequence of Changes in Shell-vear Patterns ‘The changes in shell-wear patterns and the sequence of changes described for desert tortoises may be typical of many species of tortoises and terrestrial turtles. Legler (1960) had similar findings for the ornate box turtle (ferrapene ornata Presence or absence of areolae on laminae of the shell indicated degree and sequence of wear. The anterior edge of carapace and plastron, and the elevated mid-dorsal line...wear smooth in some individuals before the first hibernation. Subsequent wear on the carapace proceeds posteriorly...when only one central areola remained, it was the fifth. The areola of the fifth central laminae, because of its protected position, persists in adult turtles that are well past the age of regular growth. Physiological proce 8 associated with shell-vear changes include: Browth and the associated increase in size, and changes in bones and scutes. Early shell-vear classes (1, 2, and 3) are marked by youth: high frequency of growth and high growth rates, substantial increases in size and length, development and ossification of bones, and thickening of scutes. The “niddle" wear classes-~4 and 5--can be identified by a reduced frequency in growth and growth rates, attainment of maximum size, and maximum development of bones and scutes of the carapace. In contrast, wear Classes 6 and 7 are typical of old age. Tortoises in these classes exhibit low frequency of growth, no growth, or shrinkage; thinning of bones and scutes; and increased porosity of bones. The physiological processes associated with shell-vear in desert tortoises may be similar for other turtles and tortoises. For example, Ab-39Legler (1960, 1963), in describing wear patterns in che ornate box turtle and the redfoot tortoise (Gecchelone carbonaria), reported that smoothly worn individuals were usually the larger, older individuals, and were not growing, He also noted that wear on the shell reduced the thickness of scutes and that greatly worn shells had only a thin layer of epidermis visible (Legler 1960). Auffenberg and Iverson (1979) in a review of Literature on terrestrial turtles stated that, "...those past the onset of maturity tend to grow much more slowly and sometimes not at all.” Chronological Age of Tortot: and the Relationship to Shell-vear Classes ‘The data presented so far for both captive and wild tortoises indicate that shell-wear classes cannot be used to assign chronological age and that shell-vear classes will be useful only for assigning relative ages to adults (e.g., young, middl ged, and old). As of this date, we have no examples of captive tortoises raised from the Class 1 to Class 7, nor of a single marked, wild tortoise that has experienced the same sequence. We ‘now that two captives (4 and 6) have lived about 80 years and are still in Class 4 (Fig. 14). Furthermore, two wild tortoises (examples 3 and 5), were captured as adults and kept in captivity for several decades. Tortoise No, 3 has spent at least 34 years in Cl. es 5, 6, and 75 Tortoise No, 5 has lived 57 years in Class 5 and early Class 6. The total longevity of captives might be crudely estimated at 115 to 130 years by combining data from Tortoises No. 4 and No. 6 with data from Tortoises No, 3 and No. 5. For example, captive No. 6 required 82 years to reach and remain in Class 4, while wild/captive No, 4 spent 57 years in Classes 5 and ‘The total duration of life in Classes 1 through 6 is 130 years. By A440vor 8a *sayo3103 eayadvo uo waup ArzAasuer SSB] JOM |]OUS A 9 § v € 2 7 € “ON @sjo101 eANded/PIM = ot s188k ve $ “ON es}o10) oandeo/puM Meee siBek 1S 9 “ON esIol04 eAandeo ¥ “ON as}o}i01 eandeg seek 08 Abmb1combining data from Tortoise No. 6 with Tortoise No, 3, the longevity is of other 115 years, These figures for longevity are consistent with tho: captive terrestrial turtles and tortoises mentioned earlier. Estimates for wild animals can be made from more fragmented informatio (1) the Woodbury and Hardy (1948) and Hardy (1976) capture~ recapture data; (2) growth data from tortoises observed over a 19-year period (Medica, Bury, and Turner 1975; Medica, personal communication); (3) capture-recapture data on shell-vear classes from nine study sites in california; and (4) the data on mean sizes for tortoises in the seven wear classes in Table 2. For example, we know from field observations that tortoises in Class 1 range from one to three years old. Those in Class 2 have chronological ages from about 3 to 20 years, based on carapace length. Class 2 tortoises have a mean carapace length of 179 mm for males and 168.5 for females. If the average growth rate of 9 mm/year (Medica, Bury, and Turner 1975) is used, then the mean Class 2 tortoise might be 13 to 14 years old and range from about 3 to 20 years. Similarly, mean Class 3 tortoises of 199- to 202-am MCL might be about 19 years old with most individuals in chronological ages of 15 to 25 years. Estimation of age for wild tortoises in Classes 4 through 7 is sore difficult. We know that (1) most tortoises over 200-am MCL grow much more slowly than the smaller, younger aninals (Medica, Bury, and Turner 1975; Patterson and Brattstrom 1972; Turner and Berry, in preparation); and (2) size is not necessarily an indication of age in animals over about 195-mm MCL. Furthermore, when turnover rates for the shell-wear classes ate examined (Table 5), it is obvious that rates are much higher for the Tortoises in Cla lower than the higher wear classe: 4, 5, and 6 show Ai=42rates of change into the next highest vear class of 1 per cent over one~ co three-year periods. While interpretation of these rates is difficult at this point, the potential is high for tortoises to remain many years in these classes. We suggest that wild tortoises may live 15 0 30 years in Class 4, another 15 to 30 years in Class 5, and lesser amounts of time (10 to 15 years each) in Classes 6 and 7. Longevity for wild tortoises my exceed 100 years, If the study plots in California and Nevada are revisited at regular Antervals over the next 20 years and data are collected on shell~wear in previously marked individuals, we will have a much better idea of shell-wear class duration and longevity in the wild and our estimates can be revised accordingly. Possible Factors Affecting Duration of Shell~vear Classes Diet, sex, and habitat may affect shell-wear classes and are discussed below. Diet, Jackson et al, (1976, 1978) described rapid growth and early maturation in captive desert tortoises; one male reached 236-am MCL and was xually mature four years after hatching, The authors believed that the rapid growth and early maturation were due to an excellent diet and a "stimulating environment." They described a "knobby" condition of the carapace in their specimens, a feature also recognized by captive fanciers as associated with rapid growth. Medica, Bury, and Turner (1975) reported nore rapid growth of wild d rt tortoises in years with higher winter rainfall and higher production of winter annuals, the principal tortoise food. Abnd3Relevant references are also available on growth and diet for other species of tortoises and turtles. Townsend (1931) studied growth rates in captive Galapagos tortoises (Geochelone elephantopus) and attributed differences in growth rates to climate and food quality and supply. Quinn and Christensen (1972) studied vestern painted turtles (Chrysenys picta belli) on different pond substrates, They found that turtles living on highly organic mud substrates grew faster than those on relatively inorganic sand substrates and suggested that the more nutritive detritus might contribute to more rapid growch. Gibbons et al. (1981) studied the slider turtle (Pseudemys scripta) at two sites, one of which had “nore optimal food and temperature." They found that while females matured at the same age at both sites, individuals at the "nore optimal" site matured at a large size. Males matured at an earlier age at the “wore ‘optinal” site. The above examples indicate that desert tortoises and other turtles are capable of groving more rapidly when a good diet is available; thus a fa growing and "vell-fed" tortoise can attain a larger size, particularly in Classes 2 and 3 than a tortoise on an “average” or restricted diet. Furthermore, an improved or excellent diet may contribute to increased longevity, e.g., lengthening of the middle~ and old-age periods (Classes 5, 6, and 7). The converse is true: a poor diet in captives often results in soft shells, low growth rates, disease, and death (Fowler 1976; Rosskopf et al. 1981; Rosskopf, Woerpel, and Yanoff 1982). The availability of critical nutrients such as minerals may be an important factor in condition of the bony carapace and the general well-being of desert tortoises. Minerals are known to be important to game Abbanimals; the use of salt licks and other types of licks has been documented for decades. However, the use of mineral deposits by turtles and tortoises to supplement dietary deficiencies has not been established. Recently Marlow and Tollestrup (1982) summarized the literature on use of mineral deposits by higher vertebrates and reported observations of mining of calcium carbonate deposits by female desert tortoises at the Desert Tortoise Natural Area in California. Six females were seen either excavating a deposit or using an existing mine. Ingested soils had significantly higher calcium levels than soils overlying or adjacent to the mineral deposits. The authors suggested that females use the calcium-rich soils during the nesting season to replenish body calcium reserves depleted during the development of calcareous egg JE mineral deposits of calcium carbonate are used as dietary supplements by tortoises and are important to their well-being, then the presence of such deposits may affect (1) tortoise distribution and abundance, (2) distribution of shell-vear classes within populations, (3) longevity in populations, and (4) duration of certain shell-vear classes within populations, For example, mineral content of soils is an important factor in distribution and abundance of pheasant populations (McCann 1961). Tortoises living on caleium-rich soils may live longer and age more slowly. Such populations might exhibit lower frequencies of the osteoporitic characteristics typical of wear Classes 6 and 7. Sex, Auffenberg and Iverson (1979) reported that many species of tortoises show pronounced differences between the sexes: In several species the female may possess a much thinner shell, with fontanelles common at wear points, such as the pelvis (Geochelone chilensis, anong others). Ab=65They offered no explanation for the thinner shell in some species, No Information was provided on age, diet, or reproductive status of such fenales. Adult females might age more rapidly and move more quickly through wear Classes 4, 5, 6, and 7 after reaching maturity than males if bone and bone-building materials are repeatedly depleted during egg production. Schneider (1961) suggested that bones of females might be nore porous because of calcium mobilization during reproduction, He found a large, old fenale which had recently died with 12 unlaid, shelled eggs. The bones of the carapace were very thin and porous and the pelvis vas a honeycomb. Four references are available on calcium mobilization in turtles and tortoises and may shed some Light on the topic. Edgren (1960) examined seasonal changes in bone density in female musk turtles (Sternothaerus edoratus). Bone density in the hind legs (the only bones tested) decreased prior to and during egg deposition but returned to normal two months late! Edgren (1960) suggested that decrease in bone density was due to loss of calcium for egg-shell material. Suzuki (1963) undertook detailed studies on morphological changes in bone associated with egg formation in the slider turtle (Pseudemys scripta elegans). The findings of Suzuki (1963) were similar to those of Edgren (1960) with an important addition, Suzuki also looked for changes in the carapace but found none. Simki: 961) examined femurs of the gopher tortoise (Gopherus polyphemus) and one tern painted turtle (Chrysemys picta picta) after injecting the specimens with estrogen. Histological examination of the bones revealed no medullary bone, a bone type which acts as a labile-calcium store in birds. Zangerl (1969), in a discussion of turtle shell anatomy, reported that the Ab-46shell, in contrast to the limb bones, does not appear to serve mineral reservoir for calcium or egg production. We know of no references suggesting calcium mobilization in the carapac. of turtles except for a note by Patterson (1970), This author suspected that ingestion of calcium might affect the flexibility of che transverse hinge at the femoral-abdominal joint in female tortoises. He made a single observation on an old captive female desert tortoise. When the individual was fed egg shells in the diet, the hinge became less flexible. Flexibility returned to the previous state 30 days after the last egg-shell feeding. Based on existing information on calcium mobilization, we suggest that the osteoporosis in the carapace of the female described by Schneider (1981) was associated with age, rather than reproductive activities. This tortoise was an advanced Class 7 and had the thinnest and most worn scutes and the thinnest bones of any carcass we observed. Furthermore, we think that desert tortoises in Classes 6 and 7 exhibit osteoporotic characteris- tics that are related to age rather than Co sex. The effects of courtship and combat behavior on shell wear were considered. Courtship activities may contribute to more rapid wear on the first and second costal scutes of females than on males, During mounting and copulation, males rake these scutes with the claws of their forelegs (Black 1976 and our observations). The action is probably more accentuated in captives where the sexes are constantly together (Glenn Stewart, personal communication). Combat behavior may cause chipping and wear on the nuchal and anterior marginals, We doubt that courtship and combat significantly affect shell-wear classes in wild tortoises, because these Ab-47activities are infrequent. In addition, most characters used in shell-vear classification are independent of the kinds of wear induced by these behaviors. Terrain and Soil Texture, We suspected that terrain and soil texture might play an important role in duration of wear for Classes 2, 3, 4, and 5, and that tortoises living in rocky terrain and on rocky soils might experience mre wear at early stages than those living in sandy or sandy loam-type soils. To test this hypothesis, linear regressions were made of wear class assignments on carapace lengths for several populations in california and Nevada (Berry, Turner, and Woodman, Appendix 5, this volume). Tortoises <180-mm MCL were selected for the test because age and size are more closely related in this group than in tortoises >180-mm MCL.. When slope coefficients for study areas with rocky habitats were compared with slope coefficients for study areas with sandy soils, there appeared to be no correlation. The habitat type may affect the wear patterns on the carapace to some extent through abrasion (Classes 2 through 5), but not necessarily the characteristics used to delineate wear classes. Wear Classes 6 and 7 obviously are not determined by abrasion. well-defined raised areas or ridges of thickened scute and bone remain on the edges of the vertebral scutes—typical contact areas for abrasion. Some owners of captives suggest that their pets wear more slowly than wild animals because their pets experience little abrasion. (Glenn Stewart, personal communication). Such captives live in dog houses or other shelters instead of burrows. However, captives euch as Tortoise No. 4, the Klein tortoise, and No. 6, the Nowak tortoise, have lived in burrows much A448of their lives and are still in the "young" adult class. Some captives have scratches, furrows, or gouges from abrasion experienced in captivity (e.g., from repeatedly crawling under a volcanic boulder in a backyard, using a dog-door to gain entry to a house, etc.). These scratches, gouges, or furrows usually can be distinguished from the kinds of wear patterns we have described above. SUMMARY Seven shell-wear classes are described for the desert tortoise. These wear classes are associated with changes in bones and scutes of the carapace (e.g-, vear patterns on the areolae and growth rings; chipping, flaking, and peeling of laminae; thickness of laminae; and osteoporotic changes in bone). Frequency of growth (using presence of new epidermal material at scute seans and/or a change of > 5 mm in length as criteria) is greater for tortoises in lover than in higher wear classes. Tortoises in shell-vear Classes 1, 2, and 3 show a frequency of 80.2 to 86.9 per cent growth, whereas those in Classes 4 and 5 have a frequency of 66.1 and 35.2 per cent, respectively. Tortoises in Classes 6 and 7 exhibit only 11.9 and 0.0 per cent, respectively. Changes in bones of the carapace can be related to both size and wear class. Tortoises in Class 1 and rly stages of Class 2 are in bone~ building phases and have incompletely ossified skeletons. Closure of fontanelles is completed in the latter stages of Class 2; growth in length and thickening of carapacial bones continues throughout Classes 3 and 4. Tortoises in Class 5 appear to be at the peak of bone development, whereas Ab-49those in Classes 6 and 7 are in the degenerative phases typical of old a Mean carapace length increases with increase in shell-vear class, with males consistently showing greater mean lengths than fenales, A general relationship exists between carapace length, shell-wear class, and size, with the larger tortoises being older. However, some tortoises apparently stop growing or only grow very slovly after reaching about 195-nm CL, and reach old age and a high wear class at a small size. Others can grow rapidly and reach a large size at a young age and low wear class. Tortoises in the seven wear classes can be assigned relative ages, using the data on frequency of growth, size, shell-vear patterns, and bone condition characteristics. Tortoises in Class 1 are hatchlings or very snall juveniles <60 mm CL. Classes 2 and 3 tortoises are generally Juvenile, immature, or subadults, and have little shell wear except on areolae and growth rings; they exhibit rapid growth and many are sexually immature, Class 4 tortoises are considered young adults. Class 5 tortoises are middle-aged adults, and Classes 6 and 7 individuals are aged animals, Diet may affect shell-wear condition and class, No relationship was found between wear class and such factors as sex, terrain, and soil type. ACKNOWLEDGMENTS ‘The U.S. Department of Interior, Bureau of Land Management, funded studies on most populations of tortois well as a mentioned herein, special contract to A. Peter Woodman to assign shell-wear classes to tortoises, The Desert Tortoise Council, through a contract with the U.S. Fish and Wildlife Service, supported preparation and development of a Ab=50computer tape for study of tortoise population data, including shell-vear. Frederick Turner assisted with shell vear and size analyses. Ross Hardy provided information on the old, marked tortoises at the Woodbury-Hardy study site in Utah, Mark Jennings and Rosemary Sage sent photographs and drawings of their captive tortoises. Mary Trotter located valuable data on Jong-rera captives and helped with editing. Lori Nicholson made useful additions to shell-wear descriptions, the key, and illustrations, Bayard Brattatrom, Betty L, Burge, Mark Jennings, Phil Medica, Lori Nicholson, Robert Patterson, Glenn Stewart, David Stevens, and Janes St. Anant, and Frederick Turner offered valuable suggestions on the manuscript, Alice Karl made sone initial attempts at dividing shell-vear classes. Mark Maley and Betty Burge loaned tortoise slides and shells from study plots in Nevada for examination; George Sheppard did likewise for Arizona. Stephanie Anderson prepared the illustrations of tortoise shell; Ab-SLREFERENCES CITED Auffenberg, W., and J. B. Iverson, 1979, Chap, 25. Demography of terrestrial turtles. Pages 541-569 in M. Hatless and H. Morlock (eds.), Turtles: perspectives and research. J. Wiley and Sons: New York. Berry, K. He, F, B. Turner, and A. P. Woodman. This volume. Appendix V. Distribution of shell-vear classes in desert tortoise (Gopherus agassizit) populations in California, Nevada, and Arizona. Black, J. H. 1976. Observations on the courtship behavior of the desert tortoise. Great Basin Nat. 36(4) 467-470. Edgren, R. A, 1960, A seasonal change in tone density in female musk turtles, Sternothaerus odoratus (Latreille), Comp. Biochem. Physiol. 1:213-217. Flower, S. $. 1925. Contributions to our knowledge of the duration of life in vertebrate animals. III. Reptiles, Proc. Zool. Soc. London: 911-981, Flower, S. S. 1937. Further notes on the duration of life in aninals. TIL, Reptiles. Proc. Zool. Soc. London, Ser. A(1):1-39, Fouler, M. E. 1976. Respiratory disease in captive tortoises. Pages 89-98 in N. J. Engberg, S. Allan, and R. L. Young (eds.), Proc. 1976 Symposium of the Desert Tortoise Council, Long Beach, Calif. Gibbons, J. W. 1976. Aging phenomena in reptiles, Pages 454-475 in M. F, Elias, 8. B. Eleftherious and P, K. Elias (eds.), Experimental aging research. EAR, Inc.: Bar Harbor, Maine. Gibbons, J. W., and R, D. Semlitsch. 1982. Survivorship and longevity of a long-lived vertebrate species: how long do turtles live? J, Anim, Ecol. 51(2):523-528. Gibbons, J. W., R. D. Semlitsch, J. L. Greene, and J. P. Schubauer. 1981. Variation in age and size at maturity of the slider turtle (Pseudenys scripta). Am. Nat, 117:861-845. Hardy, R. 1976, The Utah population ~ a look in the 1970's. Pages 84-89 in'N, J. Engberg, S. Allan, and R. L. Young (eds.), Proc. 1976 Symp. of the Desert Tortoise Council, Long Beach, Calif. Jackson, C. G., Jes, J. As Trotter, T. Hy Trotter, and M. W, Trotter. 1976. Accelerated growth rate and early maturity in Gopherus agassi2i (Reptilia: Testudines). Herpetologica 32:139-145, Jackson, C. G., Jr., T. H, Trotter, J. A. Trotter, and M. W. Trotter. 1978. Further observations of growth and sexual maturity in captive desert tortoises (Reptilia: Testudines). Herpetologica 34(2):225-227. Ab-52Jennings, M. R. 1981, Life history notes: Gopherus agassizi (Desert Tortoise) longevity. Herp. Review 12(3):81-82. Klein, H.R. 1977. Roxie and family (Gopherus agassizit). Turtle Hobbyist 2(4):6-8. Legler, J. M, 1960, Natural history of the ornate box turtle, Terrapene ornata ornata Agassiz. Univ. Kansas Publ. Mus. Nat. Hist. 11(10; 527-669, Legler, J, M. 1963, Tortoises (Geochel distribution and variation, Am. Midland Nat. 70(2) carbonaria) in Panama: 90-503. Loveridge, A. and E, E. Williams, 1957, Revision of the African tortoises and turtles of the suborder Cryptodira. Bull. Mus. Comp. Zool. 115:163-557. Marlow, R. W., and K. Tollestrup. 1982. Mining and exploitation of natural mineral deposits by the desert tortoise, Gopherus agassizii. Anim. Behav. 30(2):475-478. McCann, L. J. 1961. Grit as an ecological factor. Am. Midland Nat. 65(1) 2187-192. Medica, P. A., R. B. Bury, and F, B, Turner, 1975. Growth of the desert tortoise (Gopherus agassizi) in Nevada, Copeia 1975(4):639-643. Miller, L. 1932, Notes on the desert tortoise (Testudo agassizi). ‘Teans. San Diego Soc, Natur. Hist. 7:187-208. Miller, L, 1955. Further observations on the desert tortoise, Gopherus ‘agassizi, of California. Copeia 1955:113-118. Nigrelli, R. F, 1954, Some longevity records of vertebrates. Trans. New York Acad. Sci. 16(6):296-299, Patterson, R. 1970, A request for information on hinge flexibility and calcium metabolism in female tortoises. Inter. Turtle and Tortoise Soc. J. 4(5)239. Patterson, R. 1972. Age, please? Inter. Turtle and Tortoise Soc. Nov.-Dec. 1972:6-7. Patterson, R., and B. Brattstrom. 1972. Growth in captive Gopherus agassizi. Herpetologica 28:169-171. Quinn, J. A., and J. L. Christiansen. 1972, The relationship between pond bottom type and growth rate of western painted turtles Chrysemys picta belli in Iowa, a preliminary report, Proc. Iowa Acad. Sci. 78(3-4): 67-69. 4-53Rosskopf, W. Je, Jr., E, Howard, A. P. Gendron, E. Walder, and J. O.Britt, Jr. 1981, Mortality studies on Gopherus agassizi and Gopherus berlandieri tortoises. Pages 108-112 in K. A. Hashagen and £. St. Amant, Proc. 1981 Symposium of the Desert Tortoise Council, Long Beach, calif. Rosskopf, J. Wi, Jr.) Re W. Woerpel, and S, Yanoff, 1982, Severe shell deformity caused by a deficient diet in a California desert tortoise. Vet. Medicine/Snall anim. Clinician. April 1982:593-594, Schneider, P. agai’. a population analysis of the desert tortoise, Gopherus agassizii in Arizona. Report to U.S. Dept. of Interior, Bureau of Land Management, Phoenix District Office, Arizona, Contr. No, Az-950-CT9-0014, Simkiss, K. 1961, Influence of large doses of estrogens on the structure of the bones of some reptiles. Nature 190(4782):1217-1218, Suzuki, H. K. 1963. Studies on the osseous system of the slider turtle. Pages 351-410 in H. E. Whipple (ed.), Comparative biology of calcified tissue, Ann. New York Acad. Sci. 109, Townsend, C. H. 1931, Growth and age in the giant tortoise of the Galapagos. Zoologica: Sci. Contri. New York Zool. Soc. 9(13) 2459-474. Woodbury, A. and R, Hardy. 1948, Studies of the desert tortoise, Gopherus agassizii. Ecol. Monogr. 18:145-200, Zangerl, R. 1969. Chapter 6, The turtle shell. Pages 311-339 in C. Gans (ed.), Biology of the reptilia, Vol. 1, Morphology A. Academic Press: London and New York. 1. This report has been distributed in several drafts, each with the sane title and date, We refer to the draft distributed in February 1982. A-54Appendix © vu ve eo 6 o a on ot . ve s ot om a om 3 st mG prtn tet ost 95 oe wom og * ee a oo: a) i & sor “pron Gest vo “Forten paeppoas 6 o Ogre cH ote tute ect ter « ote 1% tot 2 & eo : ® 6 “pen owt yo ‘erin some es or Og ete cl ere eevee] . ° st eo 0 tT 9 & 8 3 : © 2 prin ont » zr oa Oe etn vac oe esl . 6 u rr A t t prin ust wo tyes womwsss§ rat oor lee oes ese aoe meeer tc . yo tr02u09 oaraead ose ee 5 8 ot : Baer eoay team oa t St Or oz “pntn erst serouon ame 00 pee el teree acta ct ae) . vo “(11 wor22%s) ou 3 eo om & a uo 3 i easy tena & eye et “prin est eeouer used € scat * 1 6 6 & @ ¢ . et a 1 8 om 8 8 i oe oy spin 1961 eu sn o 8 Tete) . oor oe 2M moa 6 i co peed ok aces cust vo *Korgeg auonazg 1 pur 9 seemers sree 88 ys meen suru oars “ou 9258 lvossay pur ‘epenoy ‘epusosttey ay Suoyretndod 98y02302 332699 om-AauaN UT S9580TS IOR-TTONS yo sopsngrasera * y o1ges asa6 zy ee ee * Sst 8 0 6 fo mH tO 1 o ° pron 6st wo WER TAOMPEEND — Oz ce 6 tom om om 6 . te tt eee 2 %8 o pron ‘oxst vo A0rTeR PIER 39MIN vee ra cow we oe eon . ve ia eis cr oe st a 1 * eg ren ovat one 0s sou om a ou 8 . ose 05 ee 1 ‘ no pron ust wo ‘sn09 st rs ss ee . ou rng D8 ot 8 6 8 2 rd cw ron ust es a eee eed . 3 ot te o 8 a 9 € € 1 Ea you pron lst yo “Soren uedueat ot ve on ° eee 8 be é 1 et 1 , « prin exst vo *4oTTeA mopeus fT ey w eee eee . oer & 0 6 t 6 2 8 1 ca er spy oper vo “Fortean voter 11 eu ® Pc or er 8 ost ° o ft ft 8 oO ot 3 w ot prin est wo *Aorten eus921 oF ae} poe 9 eoeser> x sree 69 6 me ee ws snug geeu anys cou org soeser 3098 T1945 suortay pue ‘epeany ‘eyusoszte9 uf suoyieinded seyors0% asoap om-Asuam) uy gasses sean-{foUe JO vorInaT sya °t MTAeL0-06 a ete « os et Oo oo F fg 3 a wot prin ont Borate 9 3 toy © 9 4 = oy zr Ce ee ee 3 e Pk eprtn over ae trim csery 9 gor a 0 6 f 9 OG « er a o € oO F ¢§ 5s 2 w 2% tcprin 6161 fw *Korten 938 ome o eee or ee . ve a eee : * Doh ek PRn i ‘spay oF voy o 8 © bt tog . 0°05 eee 2 toe pron eest——aw “eFeanon doous az. oe vz Ot eed or « o ie o 0 2 6 ra 5 cS * ee onet ‘1 Seren erremoni 97 eat 8 cou om «w a 6 = om 8 Mee cl (ote cree : 96 oe ere ree ree 60st oor os roy ee og . oot ey 0 8 mw a 6 ¢ 2 % w rin Lest Yo “Hou ATTEND ez 5 Geen ice oy . 6 6 69 0 6 € 2 3 5 5 prin U61 VO 880205 poomor3e9 zz uu a o + 6 £€ 6 4 " re " Oe eee fi 6 5 opin aust vo ‘ursen ota tz L pur 9 sasser> x stern eee ee cee vas sere ats cow ons euossay por ‘epenon ‘erusossreg ef suoraernded e2yo2s03 asosep om3-Aauens Uy SosseI> 2ean-TTaHP Jo oFIMaTSIETG {PTA asTable 2. Mean maximum carapace lengths and proportions of shells in wear classes 6 and 7 among male and fenale tortoises at 16 sites in california and Nevada Males Fenales Mean Percent Mean Percent carapace wear cerapace wear Areas length (am) clagses 6-7 length (nm) classes 6-7 2 Frenont Valley 250 Ft) 225 2 3 DIN 263 20 238 28 4 DrNa 261 8 233 26 5 Frenont Peak 268 5 234 ° 7 Kramer Hills 246 4 238 2% 9 Stoddard Valley 260 2 29 2 10 Lucerne Valley 256 a 240 16 11 Johnson Valley 262 5 243 13 14 Ivanpah Valley 246 6 225 v 15 Gofts 251 2% 220 2 16 Upper Ward Valley 250 3 221 2 20 Chenehuevi ash 240 10 226 16 23 Chuckwalla Bench 249 v 227 25 26 Chuclwalla Valley 250 4 232 ° 30 Arden 251 35 225 2 31 Piute Valley 236 2 218 u ASS4m 16 populations in California and Nevada (Table 3). As expected, slopes for snaller tortoises were generally low, simply reflecting less wear. Only 99 of the 976 (10.1%) tortoises <180mm in Table 1 were assigned to wear classes other than 1 and 2, Only 8 (1%) were assigned to classes higher than 3 (classes 4 and 5). As predicted, slope coefficients for larger females vere generally greater than for wales, and the overall mean for larger fenales (0.037) vas statistically significantly greater than that for males (0.030). A paired ¢~ test of the 32 values in Table 3 gave a E-value of 3.06 (ty 9) * 2-95)- Nevertheless, this pattern vas not expressed in almost half the samples compared (Fremont Peak, Stoddard Valley, Lucerne Valley, Ward Valley, Chemhuevi Wash, Chuckwalla Valley and Arden). The explanation lies in the fact that vhen wale slope coefficients exceeded those of fenales the differences were small. when slope coefficients for females were greater, they were often much greater (e.g., Piute Valley). Comparisons of slope coefficients of males in different populations (by E-tests of pairs of different population showed several "groups," e.g-, Chemehuevi Wash, Ward valley and Arden (with high slopes), the DINA, Frenont Valley, Kramer Hills, and Ivanpah Valley (with slopes £ 0.28), and Lucerne Valley, Stoddard Valley, Goffs, Chuckwalla Bench and Chuckwalla Valley (with internediate slopes of around 0.032). Fremont Peak and Johnson Valley formed another group with the lowest slopes. In a general vay, slopes were Jow (vith an tmplication of less wear) in many western Mojave areas, intermediate at e: tern Mojave and Southern Desert sites, and highest near the Colorado River and in southern Nevada. Although data from females could be grouped on the basis of t-tests of paire of populations, no intelligible geographic pattern emerged. As indicated earlier, the regression analyses just discussed are influenced not only by body size of males and females, but also by the 5-6Table 3. Slope coefficients for regressions of shell wear class on carapace length among tortoises at 16 sites in California and Nevada. Undetermined Males Females Areas Years <180 om 2180 mm 2180 mm 2 Fremont Valley 1981 0.0074 0.028 0.036 3 DTNA 1979 0.0107 0.026 0.038 4 DINA 1979 0.0089 0.027 0.045 5 Fremont Peak 1980 0.0267 0.020 0.021, 7 Kramer Hills 1980 0.0070 0.026 0.037 9 Stoddard Valley 1979 0.0140 0.033 0.026 10 Lucerne Valley 1980 0.0047 0.032 0.028 11 Johnson Valley 1980 0.0119 0.022 0.041, 14 Ivanpah Valley 1979 0.0066 0.026 0.046 15 Goffs 1980 0.0055 0.031, 0.043, 16 Upper Ward 1980 0.0097 0.035 0.034 Valley 20 Chemehuevi Wash 1979 0.0050 0.035 0.033, 23 Chuckwalla Bench 1979 0.0070 0.031, 0.047 26 Chuckwalla 1980 0.0028 0.031, 0.030 Valley 30 arden 1976-5 0.0118 0.037 0.034 31 Piute Valley 1979 0.0045 0.033, 0.056distributions of wear classes in samples. Slope can be increased by higher relative numbers of tortoises with shells assigned to high wear classes. We examined the joint effects of mean body size and relative frequency of highly'worn shells by stepwise multiple regression, using regression slope coefficients as dependent variables. The independent variables were 1) mean maximum carapace length, and 2) proportion of shell wear distribution composed of class 6 and 7 shells. This last independent variable was computed after adding half the number of tortoises of undetermined sex to male and female totals, The analysis was based cn samples set forth in Table 1. Table 3 give relevant values for the independent variables. The results of the tuo analyses are given in Table 4. Anong males analyzed, variations in slope coefficients were influenced principally by the composition of the samples--i.e., samples with relatively more highly worn shells had higher coefficients. However, mean body size also contributed substantially to the multiple R*, These two factors accounted for 68% of the observed variation in slopes, Among females, what Little variation was explained (ca. 37%) was largely owing to the composition of samples, and mean body size was of minor importance (increasing R” only about 132). The foregoing analyses have provided some understanding of the relationship between shell wear and body size, and the influence of two potential factors on this relationship. However, we have no better understanding of how shell wear is related to age. It is not encouraging to observe that the relative proportions of wear class 6 and 7 shells are not particularly similar among males and females of the same population. A rank correlation test based on figures in Table 3 gave r, = 0.327. The critical value (5% level) for rejecting the null hypothesis of no correlation between ranks of the sexes is 0.482 (Snedecor, 1956: 190). AS-8Table 4. Results of stepwise aultiple regression analyses of shell wear and body size among male and female tortoises from 16 sites. Males Fenales First independent variable entered (and multiple 8) Second indepedent variable entered (and multiple R) Percent variation explained by first variable Percent variation explained by both variables Percent of sample composed of wear class 6 and 7 shells (0.635) Mean carapace Length (0.838) 40.3 70.1. Percent of sample composed of wear clase 6 and 7 shells (0-533) Mean carapace length (0.604) 28.4 36.5 AS-9We can use the data on slopes to examine the relationships between shell wear and soil types. Do the shells of tortoises Living in gravelly and rocky soils wear faster than chose in sandy, loany soils? Tortoises < 180-nn in length are most suitable for such an examination. Soils at Sites 16, 23, and 26 are firmer, rockéer, and have extensive areas of desert pavenent. Slope coefficients are not aarkedly different from slopes of areas with the sandy, loany soil types. ‘Are populations with higher proportions of tortoises with highly worn shells (e.g., Coffs, Upper Ward Valley, Arden) really composed of more aged tortoises? And, 1£ this be true, why do some samples exhibit high proportions of worn shells anong one sex but not the other (e.g., Kramer Hills, Stoddard Valley, Plute Valley)? If there is some geographical pattern to shell wear distributions, why {s it not manifested in both sexes? one vay our understanding of wear of shells can be advanced is to clarify the relationship of wear co age, and to understand the variability of this relationship. Indeed, there may be so much variation in wear and age that only the most coarse correlations may be dravn. Until we know this co be true, the possibility remains that shell vear may be an effective way to estimate age composition of natural populations of desert tortoises. A second vay to inert our understanding of shell wear is to follow changes in composition of shell wear at a number of sites. Changes might be related to hunan uses, e.g., collecting of the larger (and often older) individuals. Differences in coaposition of shell-vear class in California in the 1970's and early 1980's might be due to past land uses, e.g, tank maneuvers of General Patton may have killed a large portion of the adults at many sites in the 1940's. aS-10Appendix 6 A DESCRIPTION OF CARCASS DETERIORATION FOR THE DESERT TORTOISE AND A PRELIMINARY ANALYSIS OF DISINTEGRATION RATES AT THO SITES IN THE MOJAVE DESERT, CALIFORNIA A, Peter Woodman and Kristin H. Berry The study of carcass deterioration in the desert tortoise (Gopherus agassizii) is a long-term and continuing project; it is an essential part o our analysis of the condition of tortoise populations in the Southwest. Mor specifically, the findings on carcass deterioration rates will contribute to our understanding of whether populations are declining, static, or increasing. Desert tortoise carcasses can persist for aany years in situ an have potential for providing information on causes and years since death, an population mortality rates. However, the stages of decay and time span for each of these stages aust be known to estimate years since death. Berry (1973) suggested that there was 2 well-defined sequence ranging from two to five years for deterioration of tortoise carcasses. The sequence was: (1) scutes ordinarily adhere eightly to the bony skeleton for six months to a year, and sonetines longer; (2) after one to two years scute gradually dry and Lighten in color, and laninae and grovth rings begin to peel; (3) scutes fall from the bone; and (4) the bone disarticulates and disintegrates, In 1975, we realized that the process was far aore complex than first belfeved and that chis was just one of many possible patterns of weathering. In 1978, we initiated a study to determine: patterns of and bone: decomposition of carcasses, including deterioration of scut decomposition patterns as related to tortoise size and age at time of death; 6-1environnental factors affecting decomposition rates; ranges in variation of carcass decomposition; and possible assignaent of tine scales to decomposition patterns, Ultinately, our goal vas to use the information to (1) estimate the age of cortofse renains; and (2) to estimate uortality rates for several populations. Between 1972 and 1982 over 1,500 carcasses were collected from pernanent study plots in California. This paper is a preliminary report of our findings. Several more years of study are needed for definitive conclusions about the number of years since the death of a tortoise and all of the factors that affect decay rates of carcasses. Several pertinent studies of decomposition rates of larger vetebrates have been conducted during the last five years. Coe (1978) studied seral phases of decomposition in three elephants whose dates of death were known. Bohcensmeyer (1978) identified six stages of weathering in bones of large African mammals and discussed tates and processes of deterioration. Hill (1980) deat vith major postnortes changes experienced by some large African vertebrates and prepared a classification of bone damage. Hare (1980) described the geochemistry of bone and the use of amino acid racemization data in determining age since death, primarily for use with fossils. The only paper on decomposition of tortoise remains is the study of decay and aortality im the Aldabran giant tortoise (Bourn and Coe 1979). The authors studies decomposition in 31 individuals over a period of 18 months. MATERIALS AND METHODS All desert tortoise carcasses involved in our experiments were in the “dry phase” of disintegration (Coe 1978; Bourn and Coe 1979; Payne 1965). The remains already had passed through the “wet” stages, described as A6-2“freshly dead” and “putrid” by Bourn and Coe (1979), “bloat” and “wet” and “collapse” phases by Coe (1978), and “Fresh, bloated, active and advanced decay stages” by Payne (1965). Under ideal circunstances ve would have selected carcasses for which both the time and mode of death were known and tortoises which had expired fumeciately prior co our experisents. Unfortunately, fresh tortoise carcasses are a rare find (except for badly damaged road kills) and even aore eifficult to locate than the elephant remains described by Coe (1978)+ Forty-two shell-skeletal remains vere selected froa the carcass collection for the experiment. They were placed in wire cages on tuo permanent desert tortoise study plots in California: Site 3, the Desert Tortoise Natural Area (Section 11), Kern County in the western ojave Desert; and Site 14, Ivanpah Valley, San Bernardino County in the eastern Mojave Desert (see Appendix 2 for site descriptions). Four wire shell cages were used at each sites each was 122 cm x 61 cm x 61 cm, Two vere covered on all sides with 13-ma-nesh hardware cloth. This size uesh excluded rodents and prevented saaller shells and all but the suallest scutes from blowing avay. The other two cages were enclosed with 38- am chicken wire, which allowed rodents access to the carcasses. All four cages vere placed on sandy-Loan or sandy-gravel substrata with good drainage and in full sunlight. The wire floors were buried in so{l go that the shells rested completely on the substrate. Most shell remains were collected in spring and fall of 1978 from six 6-3permanent study plots (see Appendix 2 for site descriptions). Three sites were in San Bernardino County: Site 8, Calico (2 carcasses); Site % Stoddard Valley (4 carcasses); and Site 13, Shadow Valley (1 carcass). One collection site was in Kern County at Site 3, the Desert Tortoise Natural Area (26 carcasses); one in Riverside County at Site 22, Cottonwood Springs (7 carcasses); and one in Imperial County at Site 27, Cargo Muchacho (1 carcass). One carcass was collected in the fall of 1979 at Site 14, Ivanpah valley; the tortotse probably had died between June 1 and mid-October. Tortoise carcasses of a variety of sizes and in various stages of decoupositton--from recently dead to obviously advanced stages of decay and weathering--were selected for placement in the cages (see Table 1 for shell sizes and their distribution anong cages). Fourteen juvenile and tanature tortoises with shells under 179m maxiaus carapace length (MCL) vere included; they were believed to have died within a year of collection. 411 shells had unfaded scutes (Figs 1a) with the luster typical of live tortoises and unweathered bones (Fig. 2a). Three adults estimated to have died within one year of collection were included. They had a putrid smell and dried flesh remaining within the shell. The remaining 21 carcasses were estimated to be older than one year. They were selected to represent various degrees of bone disarticulation, sloughing of scutes from bone, and bone and scute conditions. We hoped to develop estimated time periods for each bone and scute condition class and the stages of scute sloughing and bone aisarticulation by analyzing data from numerous individuals. A6~4Table 1, Size classes of desert tortoises in and near the shell disintegration cages in Ivanpah Valley, San Bernardino County, and in the Desert Tortoise Natural Area, Kern County, California. numbers of tortoises in cages Maxisun carapace Chicken Hardvare-ciotn Outside Total Length (sa) wire cages cages cages Adule 2: > 240 5 3 2 10 Adule 1: 208-239 4 8 4 16 Sub-adult: 180-207 i 1 2 Ienature 2: 140-179 1 7 Iapature 1: 100-139 1 . Juvenile 2: 60-99 4 2 6 Juvenile 1: <60 2 3 1 Total v Fry 7 2 AS-5hatchling plate or areola growth rings —S hatchling plate or areola growth rings paar eopeeeeee e hatchling plate or areola growth rings oa “1 0 10 20 centimeters 0 10 20 centimeters Fig. 1, Classes for conditions of scutes attached to or associated with a de Class I, scutes appear in cross- section as on a live tortoise; b: Class 2, growth laminae on scutes begin to peel; c: Class 3, peeling of laminae on utes {s more pro- nounced; scute curls downward and inward; d: Class 4, laminae peel and scute breaks apart. 26-6Fig. 2. Classes for conditions of bones of desert tortoises during the disinte- gration process. a: Class 1, bone appears smooth, as on a live tortoise; b: Class 2a (for tortoises "100-mm carapace length), generally some porousness; c: same as b, but shown at 40 X magnification; d: Class 25 (for tortoises with fully ossified bones), bones rough, may be peeling and cracking; e: Class 3a (for tortoises 100-mm carapace length), very porous, shown at 40 X magnification; £: Class 3b, subsurface layers of bone are peeling or cracking; g: Class 4, mosaic surface cracking. The example of bone depicted is a costal or rib bone. 6-7Eight carcasses had one or aore legs remaining. These vere distributed as follows: two in the hardware-cloth cages (1 in each) and eix in the chicken-wire cages (three in each). Six of the eight were shells of juvenile tortoises (<100-nm CL), one was an immature (121-am MCL), and the last was 235-nm MCL adult male. The inmature carcass had tvo rear legs remaining, but the anterior half of the shell was chewed avay. Four other carcasses, all adults, had plastrons partially or completely cheved off, presumably before death by a coyote of other predator or after death by a scavenger. Only the carcass of the adult male still had visceral contents within the shell. Most shells were placed with the plastron on the substratum, but one adult shell in each of the Site 3 cages and two adult and tvo juvenile shells at Site 14 were placed on their carapaces. Of the six shells outside cages, two (one at each site) were placed at the eastern bases of creosote bushes (Larrea tridentata). There they were exposed to direct sunlight 35 to 45 per cent of the day. One shell at each site was placed under the canopy of the western base of creosote bushes. These two carcasses were exposed to direct sunlight about 75 per cent of the day. The other two shells (one at each plot) were placed in the open in small, sandy washes and exposed to direct sunlight throughout the day. The carcasses were first placed in the cages in mid-November of 1978 and revisited in mid-November of 1980 and early November of 1981. On these visite the renains were photographed and their conditions recorded. Classes or stages of scute and bone disintegration were identified and defined with the help of: (1) data and nomenclature from the Desert Tortoise 6-8Shell Remains Card developed by K.H. Berry and L. Nicholson; (2) weathering stages described by Bebrensmeyer (1978); (3) the two visits to the cages} and (4) observations of approximately 1,000 shell-skeletal remains from the 27 permanent desert tortoise study plots in California. RESULTS Processes of Carcass Detertoration and Disintegration In describing the different phases of deterioration, care was taken co separate predator damage and congenital abnormalities in scutes and bones from changes associated with death. ‘The dry phase of deterioration and disintegration involves at least four distinct but interrelated processes: (1) weathering of the scutes theaselves, (2) sloughing of scutes from bones, (3) weathering of bones in the skeleton, and (4) disarticulation of the bony skeleton, Weathering of scutes and bones is described in Tables 2 and 3, respectively. Deterioration or weathering {s a gradual, continuow and generally sequential process, but has been broken into stages of condition classes for convenience. The condition classes of scutes and bones are based on examination of 42 shell-skeletal remains in various states of weathering over a period of three years. The descriptions are Limited to bones of the carapace and plastron because the skull, lisb, pelvic, and pectoral girdle bones often disappear before collection during "wet" phases of decay. More than one stage of bone or scute weathering can be found on a carcass, In some cases, gore than one condition class can be categorized for single piece of bone. However, the individual scutes generally can be assigned a single condition class. The sloughing of scutes from the skeleton vas divided into eight stages, 6-9Table 2. Classes for conditions of scutes attached to or associated with a desert tortoise carcass. Clase Description of condition No fading of scutes; scutes appear as on a live tortoise (Fig. la). Growth laminae (scute material laid down during successive growth periods) are not peeling. Scutes of large, old tortoises, either live or recently dead, have a luster, even though browth laminae may be peeling naturally.’ when a luster 1s present on such a dead tortoise, scutes are considered Class 1+ Either scutes exhibit fading or growth laminae are beginning to peel or both (Fig. 1b). During the fading process, scutes generally become Lighter and lose all luster. When growth laminae peel, the thickened portion or edge of the laminae generally begin curling upward toward the exposed hatchling plate. Laminae at this stage are brittle and break easily when touched. Scutes faded, peeling, and possibly curling. In curling, scute edges may turn downvard and invard 90° (Fig. 1c). Scutes are brittle; most have sloughed from bone. Laminae either will be partially or completely broken away. Scutes have characteristics of Class 3, but are also breaking apart (Fig. 1d). Scutes generally torn laterally at thin areas between thickened laminae; almost alvays are sloughed from bone. (Note: sometimes predators or scavengers will break scutes when gnawing on the shell. Such breaks do act warrant Class 4 designation.) Class 4 scutes probably have more age variation than the other three classes owing to various causes of tearing. The previous classes appear to be dependent priaarily on amount of exposure. Tearing of scutes may depend on the stresses from curling as well. Scutes disappear. 6-10Table 3. Classes for conditions of non-limb bones of desert tortoises during disintegration. class Description of condition a Bone solid and smooth with no roughness or porousness (at least visible to the naked eye) on the surface (Fig. 2a). Generally bleached shiny white if exposed, brownish if scutes have just sloughed. 2a Tortoises <100-mm MCL where bones are thin and ossification incomplete: generally sone porosity (small, round or oval holes 1mm in diameter and depth visible to the unaided eye on the bone surface) (Figs. 2b and 2c). Bone usually white, sometimes brownish. 2» Tortoises with fully ossified shells (generally >120-mm MCL): bones rough or traversely ridged and brownish in color (Fig. 2d. Roughness generally occurs when scutes remain attached either tightly or loosely to bone. Bones may peel and crack when carcass is exposed in direct sunlight most of the day. When peeling of crackling begins, the outer bone surface chips off in small, thin rectangles. Bone is a dullish white color. 3a Tortoises <100 mm MCL: bones are very porous with over 75 per cent of the surface pitted. Concavities of pits generally are not round but are more elongated, larger, and deeper than concavities in Class 2a. Bone is generally more dull than in Class 2a. (Fig. 2e)- 3b Tortoises with fully ossified shells (generally >120 um MCL: subsurface Layers of bone peel and crack (Fig. 2£). Bone appears the same ac Class 2, except that subsurface layers peel. Bone color is chalky white. 4 Mosaic surface cracks (after Behrensmeyer 1978); bone has many small, generally shallow, interconnected cracks (Fig. 2g). The mosaic patterns tend to appear when bones decompose in shade. Bone crumbles, breaking into many small pieces when crushed between thumb and fingers. Bone usually chalky white, sometimes pinkish. 5 Bones disappear. 6-11and the disarticulation of bones was divided seven stages. The stages of scute sloughing from bone were based on the percentage of scutes loose, attached, or sloughed from the bony skeleton: (1) scutes adhere tightly to bone; (2) 10 per cent or fewer of the scutes are loose; (3) more than 10 percent and up to 50 per cent of the scutes are loose; (4) gore than 50 per cent of scutes are loose and/or fever than or equal to 10 per cent may have sloughed; (5) more than 10 but fewer than or equal to 50 per cent of scutes are sloughed; (6) more than 50 but fewer than 70 per cent of the scutes are sloughed; (7) all scutes are sloughed; and (8) the scuted disappear. After scutes have sloughed, they aay remain near the bony skeleton for some time. Disarticulation of the bony skeleton of the plastron and carapace is described in similar stages: (1) bones are fully articulated, and there is no evidence of separation at the sutures; (2) bones start to separate at the sutures or seams; (3) bones separate at the sutures or seams but are still in places (4) fewer than or 10 per cent of the bones disarticulate; (5) between 10 and up to 50 per cent of the bones disarticulate; (6) more than 50 per cent of the bones disarticulate; and (7) bones disappear. There is no single, distinct sequence or pattern of deterioration or weathering, particularly for scute sloughing or bone disarticulation. For example, over 50 per cent of the scutes of one adult, dead for perhaps up to two months, had sloughed and several skeletal seans had separated. The scutes of another adult, whom we estimated to have been dead over four years, still were adhering tightly to the skeleton, and seams between the bony plates showed no signs of separating. No appreciable change occurred after the carcass had been in the cage two years (total time since death estimated at A612six years). Estinating Tice Since Death for the Carcasses ‘The dour cistinet processes of weathering—degree of looseness o} sloughing of scutes, disarticulation of the bony skeleton, weathering rates 0 scutes, end veatkering rates of bone--are shown in Tables 4, 5, 6, and 7 respectively, for each of the 42 remains, The condition of carcasses fcr each of the four processes is depicted in the tables at the start of the expericent and two, three, ana four years later. ‘The 42 carcasses are arrayed both by size and by estinated tine since death at the start of the experinent. The size classes and estinated tines since death for the carcasses are presente in Table 8. In all cases, the nost weathered portion of the carcass was used for assigning a condition class. That portion of the carcass had to occupy a least an area cf 1 ca“, Sehrensmeyer (1978) used this sane criterion in assigning veathering stages to bones of large African uammals. At the start of the experiment, tortoise carcasses were grouped into four categories for estimated time since death: (1) dead less than one year; (2) dead one to two years; (3) dead gore than two but less than or equal to four years; and (4) dead wore than four years (Tables 4, 5, 6, and 7). These estinates vere based on the rates of weathering and deterioration for carcasses shown in Tables 4, 5, 6, and 7; scute and bone condition classe: were the key factors in these groupings. Classification of the condition of scutes, when scutes are present, A6-13,appears to be the most reliable neans of deternining the age of tortoise carcasses, because tortoise remains are exposed to the elements inuediately after death. There are obvious problens with again skeletal remains using only tone. If scutes remain on one carcass for several years but are 50 per cent sloughed from another within two weeks to six sonths, the bones will not be exposed to equivalent amounts of sunlight, soisture, and changes {n temperature. they may not weather equally in a given period of tine. However, if only bones are present, then the only choice is to use the There is too muck variation in the sloughing of scutes snd disarticulation cf the bony skeleton to use these features in aging. The data points on Tables 4 and 5 are scattered in all four estisated carcass age classifications. ‘Those carcasses with scutes in Class 1 at the start of the experiment were placed in either the “estimated dead less than one year” or the “estimated dead one to two years” categories. The deci! ion for splitting the Class 1 carcasses into these two categories was based on changes in scutes and on the rapidity with which the three carcasses in the one-to-tworyear category had moved from bone condition Class 1 to bone condition Class 2 or 3 during the three-year study period (Table 7). Note that tortoises DINA-GW 7 and DINA- HC-6 only changed one seute condition class in three years. Two others, DINA- CW-1 and IV-CH-9, changes two classes in the sane tine period, but had th very thin, flaking scutes typical of tortoises of advanced age. In contrast, the three tortoises in the estinated dead one-to two-years period all change: two to four condition cl: ‘Those carcasses in scute condition Class 2 at the beginning of the project were placed in the “estimated dead tvo to four years" category, while A614