Herpetologists' League

Ecology, Reproduction, and Reproductive Effort of the Iguanid Lizard Urosaurus graciosus on the Lower Colorado River Author(s): Laurie J. Vitt and Robert D. Ohmart Source: Herpetologica, Vol. 31, No. 1 (Mar., 1975), pp. 56-65 Published by: Herpetologists' League Stable URL: http://www.jstor.org/stable/3891986 Accessed: 22/09/2008 16:37
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ECOLOGY, REPRODUCTION,

AND REPRODUCTIVE

EFFORT

OF THE IGUANID LIZARD UROSAURUS GRACIOSUS

ON THE LOWER COLORADO RIVER
LAURIE J. VITT AND ROBERT D. OHMART
ABSTRACT: Urosaurusgraciosus was studied from March through November 1973 on the lower Colorado River, primarily Yuma and Mohave counties, Arizona. Limited reproductive informationfrom the same population in 1974 was also evaluated. The species is strictly arboreal,inhabiting primarilymesquite, ironwood, and smoke trees. Foraging occurred in the outer canopy of trees and stomach contents showed a high incidence of arboreal insects. Seasonal shifts were apparent in the diet, probably reflecting seasonal differences in insect abundance. Reproductive season extended from May through August. Clutch size for 23 9 9 averaged 5.3 (range, 3-10) and there was the potential for two clutches per season. In d 8, spermiogenesiscommenced in April and sperm were present in the testes and epididymis from April through July. Fat body cycle in 8 8 appeared to be a function of food supply. Fat body cycle in 9 9 was inversely related to reproductive cycle early in the season but did not show this trend later in the season. Tail break incidence was low as compared to other species and observations on foraging and climbing suggest that the tail serves a balance function, with autotomyfor predatorescape and intrasexualfighting secondary. Reproductive effort, defined as the ratio of total clutch calories of ash-free dry weight (AFDW): total body calories (AFDW), averaged 0.640 (range, 0.474-0.725). Mean caloric content per gram of egg (AFDW) was 6,612 compared to 5,492 calories per gram of body (AFDW). The wet ratio of clutch weight: body weight was 0.313 (range, 0.271-0.399) whereas the dry ratio was 0.531 (range, 0.4030.604). In terms of reproductiveand other attributes, U. graciosus fits the r strategy.

UROSAURUS graciosus inhabits small shrubs in sparsely vegetated desert regions of western North America (Stebbins, 1954), but it is common in larger vegetation in riparian situations in Arizona and southeastern California. Thermal ecology of this species was studied by Gates (1963) on the Hassayampa River near Wickenburg, Arizona, and Brattstrom (1965) included this species in a study of reptile body temperatures. Norris and Lowe (1964) studied color matching in U. graciosus from the Mohave Desert, and Pianka (1965, 1967) included this species in studies of lizard species diversity in flatland deserts of North America. Our study presents new information on reproduction and ecology of U. graciosus in riparian habitats on the lower Colorado River, in combination with estimates of reproductive effort based on caloric content of eggs and bodies.
MATERIALS AND METHODS

Description of Study Area.-The study area was located between 320 30' and 350 15'
HERPETOLOGICA

N and 114?-115? W, extending from near Topock, Mohave County, Arizona in the north, to the Mexican-American International Boundary in the south. The desert in this region falls into the "Lower Colorado Desert Subregion" (Shreve, 1951) and a general description of the habitat appears elsewhere (Vitt and Ohmart, 1974). Samples of U. graciosus were collected from March through November of 1973. In addition, data from 9 9 collected in the same localities during June, July, and August of 1974 were included. Lizards were collected by hand, noose, and BB rifle. At the time of collection, snout-vent length (SVL), total tail length, and length of regenerated tail portion (if any) were recorded to the nearest millimeter. Data on microhabitat included vegetation type where lizard was first sighted, time of day, position of lizard, and whether the animal was basking, foraging, or attempting escape. For analysis of diet, contents of stomachs were identified. Volume of excised stomach was determined by liquid volume displacement in a narrow graduated cylinder. The

31:56-65. March 1975

March 1975]

HERPETOLOGICA

57

stomach was then emptied and the volume of the lining was determined in the same manner. The difference between the two values yielded stomach content volume. The relative proportion of each type of food item was determined visually with the aid of a gridded petri dish, taking into account differences in thickness of prey items. The volume of any given food item could then be determined by multiplying the proportion taken up by that item by the stomach content volume. All food items were identified to at least family where possible. Incidence of parasites and non-food material such as rocks, sand, and miscellaneous vegetation was also recorded. Food items were then grouped into broad categories related to probable spatial location within the habitat in order to gain insight into foraging patterns of U. graciosus. These data were then coupled with observations on foraging and feeding behavior. Lizards used for reproductive information were fixed in 10% buffered Formalin within one hour of collection. After 24-48 hours fixation, lizards were stored in 50% isopropyl alcohol. Laboratory measurements on testes and fat bodies of 8 S included length, width, and weight. Since shape of fat bodies was variable, only weight data were used. Fat bodies were apparently not affected by storage in isopropanol since they were solid and intact when measured. Left testis and attached epididymis of each 8 were embedded in paraffin and sectioned at 10 jum after standard dehydration. Sections were doubly stained with Harris'hematoxylin and eosin. For each Y, clutch size, egg length and width (ovarian and oviducal), and weight of fat bodies were recorded. Weights were to the nearest 0.001 g and linear measurements were to the nearest 0.1 mm. Lizards used for estimating reproductive effort were fresh frozen in the field using dry ice (solid CO2). Only 9 9 with freshly ovulated (oviducal) eggs were used. Wet weights of body and total clutch were recorded to the nearest 0.1 mg. In addition,

number, size, and location of oviducal eggs were recorded. Bodies were then oven dried at 700 C and eggs freeze-dried to constant weights (unsatisfactory drying of bodies resulted from attempts to freeze-dry them). These weights were also recorded to the nearest 0.1 mg. Bodies were ground in a Wiley Mill and eggs were ground by mortar and pestle. Samples of each body and clutch were ashed at 5500 C for 24 h for correction of caloric determinations. This was necessary since ash content of bodies and eggs are quite different. Three aliquant samples (9-14 mg) per lizard body and per individual (eggs pooled) clutch were burned in a Phillipson microbomb calorimeter (Phillipson, 1965). Calories of ash free dry weight (AFDW) were then calculated in the standard fashion, utilizing known calories per unit weight of benzoic acid as a standard. The ratio of total clutch calories to total body calories was utilized as the best estimate of reproductive effort.
RESULTSAND DIsCUSSION

On the lower Colorado River, U. graciosus was strictly arboreal, inhabiting mesquite (Prosopis juliflora and P. pubescens), smoke trees (Dalea spinosa), and ironwood trees (Olneya tesota). A few adults were found in cottonwood trees (Populus fremontii) and several juveniles were found on creosote (Larrea divaricata) and other small shrubs. During spring and fall, lizards were active from late morning to late afternoon. During late spring, summer, and early fall there was a shift toward early morning (05300800) and late afternoon (1800-2000) activity. Few were seen during midday at this time, and those observed were not foraging but rather positioned on tree trunks in the shade. During warmer months, individuals were found at night, apparently sleeping in horizontal positions on branches. During spring and late fall, lizards did not spend the nights in trees, but could be found early in the morning beneath bark and in packrat nests. Feeding and Foraging.-Foraging activity

58

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[Vol. 31, No. 1

TABLE 1.-Summary of stomach contents of 87 Urosaurus graciosus collected between March and November, 1973 from the lower Colorado River. Thirty families of insects were present in lizard stomachs in addition to mixed larvae, spiders, and miscellaneous matter. Insects were grouped according to the place they occur in the habitat in order to gain insight into foraging place for U. graciosus. Prey category Number % Total number Volume (cm3) % Total volume Frequency*

Arboreal invertebrates Homopterans Hymenopterans (not including ants) Hemipterans Orthopterans Dipterans Neuropterans Coleopterans Trichopterans Odonata Lepidopterans (parts) Terrestrialinvertebrates Orthopterans Neuropterans

36 12 219 3 43 3 79 6 2

2.64 0.88 16.08 0.22 3.15 0.22 5.79 0.44 0.15

0.93 0.33 1.11 0.59 0.62 0.13 1.20 0.07 0.32 0.07 0.50 0.01

9.51 3.30 11.36 5.98 6.44 1.29 12.11 0.86 3.22 0.71 5.14 0.15

15 10 28 3 11 3 30 5 2 1 5 1

5 1

0.36 0.07

Mixed invertebrates (including families with both arboreal and terrestrial species and larval forms) Coleopterans (Family 18 9 Tenebrionidae) 1.32 0.32 3.24 842 Ants (Formicidae) 61.82 1.15 11.75 37 11 Unidentified insect parts 0.27 2.80 7 31 2.28 0.15 Insect eggs 1.55 Larvae 21 1.54 14 Eruciform 0.52 5.26 14 1.03 1.31 1 Scarabaeiform 0.13 7 0.55 2 0.51 Elateriform 0.05 6 0.44 0.15 Unidentified 1.53 6 13 0.95 0.34 3.47 Spiders 10 Miscellaneous Lizard (shed skin) Immature U. graciosus Sand, gravel, and vegetation parts TOTALS 0.05 0.19 0.58 1362 99.96% 9.78 0.47 1.94 5.93 99.87% 1 1 55 268

0.07

* Number of stomachs containing the specific item indicated.

occurred in the morning with seasonal shifts as mentioned earlier. Lizards foraged in the outer canopy of the vegetation. One individual descended a smoke tree, crossed 10 m of a sandy wash, ascended a mesquite, climbed to an outer branch tip, and devoured a small vespid wasp. The actions were so deliberate that it appeared the lizard observed the wasp from its original perch in the smoke tree. Both the "sit and wait" and "widely foraging" strategies

(Pianka, 1974) were observed. Here, "widely foraging" is somewhat restricted, referring to active searching from branch tip to branch tip. Individual lizards, while apparently basking, would take any insects that moved in front of them. Information on food habits (Table 1) indicates that U. graciosus is restricted in diet only by the microhabitat that it utilizes. The wide variety (over 30 insect families in addition to other arthropods) suggests

Alarch 1975]

HERPETOLOGICA

59

50
12 A

w8

~~~40
1
*I

o
#14

Ants
~~~~~~~~A
.4
2 20 z

> 30 30
T/

~~~~~'00

rtotrn

O,tntopterans
Z

20

1~"

/~~~~~~

A 10 At

~ ~
-

~
-

~ ~

Homopterans
'A,

AAll Larvae
b .... ..A

Hemipterans

MAR-APR1MAY

JUN 'JUL

AUG

SEP

OCT-NOV

MONTH
FIG. 1.-Seasonal variation in the diet of Urosaurus graciosus from the lower Colorado River. Only the most common items in terms of % volume (see Table 1) are shown. Seasonal differences in the utilization of prey items suggest differences in seasonal relative abundance of prey in view of observations on foraging (see text).

that this species is a very generalized feeder. Seasonal shifts were apparent in the composition of the diet (Fig. 1) and probably reflect seasonal abundance of potential prey items. Overall, arboreal families of the following orders comprised a majority of the diet; hemiptera, coleoptera, homoptera, orthoptera, and hymenoptera. Compared to the sympatric semiarboreal species Sceloporus magister (Parker and Pianka, 1973; Vitt and Ohmart, 1974), U. graciosus utilizes more insects occurring in the outer canopy of the vegetation than S. magister and the latter utilizes more terrestrial insects. Barnes (1974) found that these two species even utilize different ant species. Since the foraging patterns of the different ant species were similar to the foraging patterns of the lizard species feeding on them, the data of Barnes (1974) is in full support of the above discussion. Body form of U. graciosus is slender as compared to either S. magister or U. ornatus, both of which occur in sympatry with U.

graciosus in areas along the lower Colorado River (Table 2). The slender body form coupled with small body size of U. graciosus is advantageous in terms of agility and ability to forage in the outer canopy of vegetation where branches are thin. In addition, this body form is adaptive in terms of predator escape in that the lizards become cryptic in form and color (Norris and Lowe, 1964) when pressed against a thin branch. The long tail appears to serve a balancing function, similar to the lizard Anolis carolinensis (Ballinger, 1973). Lizards observed foraging extended the tail, apparently for balance, while positioned in a perpendicular stance on small limbs. Since tail retention would be at a premium if in fact balance were important, incidence of tail break as evidenced by regenerated tails would be expected to be low as compared to species with tail adaptations primarily for predator escape. In fact, tail break in U. graciosus (Fig. 2) is quite low as compared to either U. ornatus (Vitt, 1974a; Barnes, 1974) or S.

60
TABLE

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[Vol. 31, No. 1

in 2.-Comparison of body proportions three arboreal or semiarboreal lizards from the lower Colorado River. All figures represent mean ratios of adult d 8 (N = 10/species). Data on 9 9 were also collected but were not different from those of d . In terms of tail length and body width, U. graciosus was significantly different from U. ornatus. U. graciosus was significantly different in all ratios from S. magister. Overall, U. graciosus is more slender in body form than the two other arboreal forms.
Tail length/ SVL Head width/ SVL Head length/ SVL Body width/ SVL

.02

Species

.00 -_
C))

__
3

Urosaurus graciosus 2.24*

0.18**

0.21**

0.22*

F-.03-

Urosaurusornatus 1.69
Sceloporus magister 1.30
ornatus and S. magister.

0.19
0.23

0.22
0.18
.05)

0.25
0.32
from U from S.

(D 02
F-)

* Indicates significant difference (P < .05) ** Indicates significant difference (P < magister, but not from U. ornatus.

ol
r OC-

f3
Urosaurus graciosus MAY JUN JUL

--

magister (Parker and Pianka, 1973; Vitt et al., 1974; Vitt and Ohmart, 1974). It must be kept in mind that U. graciosus is capable of tail autotomy and some tails are undoubtedly lost from intrasexual fighting and predation. In view of work by Congdon et al. (1974), the tail adaptation in U. graciosus most likely represents an adaptive com-

MAR APR

AUG SEP

NOV

FIG. 3.-Seasonal changes in weights (in grams) of testes and fat bodies of adult Urosaurusgraciosus d d from the lower Colorado River. Horizontal line representsthe mean and bars represent one standard error of the mean. * Sample size for testes was 2 here since one lizard had been shot through the testes.

u 0300 \
---,*

L20
-

0 0 - --

810 20

00 *--e--@

-0---0

15

Urosaurus graciosus N =88m

n 05-

promisewhere retentionfor balancingfunc~~~~~~~~~~~0 tion is important,but autotomyfor predator escape and escape in intrasexualencounters is also important. MaleReproduction Fat Body Cycling. and -The breeding season for d d extended from April through July. Mating was observed and spermwere presentin the testes and epididymis during this time. Testes remainedat maximumsize (Fig. 3) during this period. Testes in Marchwere inactive and no spermiogenesis was observed. April testes were much largerand spermatogonial
5155

cell proliferation, spermatogenesis, and spermiogenesis were occurring. The epiFiG. 2.-Composition of samples of Urosaurus didymal lining was regressed during March, graciosus from the lower Colorado River showing but by mid April, the epididymal epithesexual dimorphism in size and relative tail break. lium increased in height with the cell nuclei In view of other studies on tail breaks in iguanid lizards, the high incidence noted in 20-40 mm S 8 migrating toward the basal membrane. Sperm were present in the epididymis at is probably a result of sampling error.
20-35 51 5 556 35-40 141-45 145-50 Snout vent Length (mm) 50

March 1975]

HERPETOLOGICA

61

E 2
z
.

u
U

400
30-

w
20F-

10-

0 JAN FEB MAR APR MAY JUN JUL MONTH AUG SEP OCT NOV DEC

FIG. 4.-Seasonal high (upper line) and low (lower line) temperatures and rainfall data from the Yuma substation of the National Weather Service. The study area is characterizedby little fluctuation between daytime high and nighttime low temperaturesin addition to little rainfall.

this time. Spermiogenesis was not occurring in July testes but sperm remained in both the testes and epididymis. Regression was not yet apparent. By August, regression was apparent and cellular debris was present in the testes and epididymis. At this time, the epididymal epithelium was reduced in height and the nuclei were positioned away from the basal membrane. September and November testes were very regressed and sperm were absent from the testes and epididymis. In terms of testes size, Gates (1963) reported a similar cycle for both U. graciosus and U. ornatus near Wickenburg, Maricopa County, Arizona. Near Phoenix, however, Parker (1973) noted that U. ornatus showed increases in testes size during February and March but in southern New Mexico and west Texas, U. ornatus displayed a testicular cycle similar to that reported here for U. graciosus on the lower Colorado River. Fat bodies of g U. graciosus were large

upon emergence from overwintering and displayed a marked size decrease with the onset of the reproductive season (Fig. 3). Shortly after breeding season began, and during a period when insect abundance appeared high (indirect observation based on vegetation blooms in the area), weights of fat bodies increased. During July, fat bodies showed a distinct decrease in size, possibly reflecting a food shortage. Since summer rains did not occur at this time (Fig. 4), food shortage as the result of localized drought could have occurred. Following rains in August, fat bodies of M S increased in size and remained large throughout the remainder of the season, a time when orthopterans became abundant in the stomachs (Fig. 1). Although the fat body cycle of 8 S appears to be inversely related to the testicular cycle (see Fig. 3), correlation of testes and fat bodies was low (r =-.035) and insignificant (F = 0.05, df = 1/40).

62
TABLE

HERPETOLOGICA

[Vol. 31, No. 1

3.-Female reproductive data for three arboreal or semiarboreal lizard species from Arizona. Figures in some instances are interpolations from the data presented by the cited authors. Also, the information on Sceloporus magister is not limited to Arizonasince the study included data from throughout the species' range.
Species x Clutch size No. clutches per season Reproductive season Minimum SVL at maturity Source

Urosaurusgraciosus Urosauruis graciosus Urosaurus ornatus Urosaurus ornatus Sceloporus magister

5.3 4.6 7.6 4.8 8.4

(3-10) (2-7) (6-10) (2-7) (3-12)

1-2 1-2 1-2 1 or more 1-2

May-Aug May-Aug May-Aug Mar-Aug May-Aug

47

44 80

This paper Gates (1963) Gates (1963) Parker (1973) Parker and Pianka (1973)

The fat body cycle of U. graciosus was quite different from that of a sympatric semiarboreal species, S. magister (Vitt and Ohmart, 1974) even though collections were made simultaneously. S. magister had relatively small fat bodies from March through June with a marked increase during July and August, followed by a decrease in September. Perhaps differences in the relative seasonal abundance of insects within the microhabitats where foraging occurs accounts for the observed fat body cycle differences. Current studies by the authors on the same species in another riparian habitat, utilizing large samples, may bear this out. Although there are no fat body data on U. ornatus, differences in the foraging micro-

10w8
N

6D

0 *0 * 0 2n =23 r =0.39

U 4-

I~~~~~ I 50 55 SNOUT-VENT LENGTH (mm)

(SVL) for 23 Urosaurus graciosus

60

FIG. 5.-Clutch size data as related to body size

9 9 collected

along the lower Colorado River during 1973-74.

habitat (Barnes, 1974) between this species and U. graciosus and S. magister would be expected to influence fat body cycles relative to insect abundance as discussed above. Female Reproduction and Fat Body Cycling.-Deposition of lipids in follicles was apparent by the first of May during 1973. The first 9 containing oviducal eggs was collected on 28 May. 9 9 with oviducal eggs were found from the end of May through the end of August, suggesting an extended breeding season. In addition, two different size groups of juveniles were observed in late fall indicating success of early and late clutches. Shaw (1952) reported incubation times of 62 and 78 days for eggs of this species under laboratory conditions. 9 9 producing early clutches had the potential of producing a second clutch in view of the above information, even though we do not have mark recapture data to verify this. It is not uncommon for deserticolous insectivorous lizards to produce multiple clutches within a season (Table 3). Mean clutch size for 23 9 9 was 5.3 (range, 3-10) and there was very little difference between clutch size of 12 9 9 in spring (X = 5.1, range = 4-9) and that of 11 9 9 in late summer (X = 5.6, range = 3-10). There was a low and insignificant correlation (r = 0.39, F = 3.783, df = 1/21) between clutch size and SVL, an indicator of relative age (Fig. 5). Gates (1963) reported a mean clutch size in 12 9 9 from the Hassayampa River of 4.6 (range, 2-7). The breeding season for U. graciosus firomthe lower Colo-

AMarch 1975]

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63

TABLE 4.-Fat body weights of adult 9 9 of Urosaurusgraciosus from the lower Colorado River. Included are only those 9 9 collected during 1973. Reproductiveand nonreproductive 9 9 have been separated to facilitate comparisons. Reproductive 9 9 include only those with enlarged yolked follicles or oviducal eggs, and those with large corpora lutea, indicative of having just laid. Fat body weights (FBW) are in grams and snout-vent lengths (SVL) are in millimeters. Month N x SVL (range) Reproductive condition xi FBW (range)

April May June July August September

1 3 1 6 1 5 4 6

48 51.7 52 50.7 57 52.6 53.3 50.7

(49-54) (47-54) (49-57) (51-55) (47-55)

nonreproductive reproductive nonreproductive reproductive reproductive nonreproductive reproductive nonreproductive

0.051 0.000 0.037 0.006 (0-0.034) 0.028 0.012 (0-0.044) 0.055 (0-0.119) 0.091 (0-0.300)

rado River was shorter but clutch size was larger than for U. ornatus from near Phoenix (Parker, 1973). Gates (1963) indicated no difference in length of breeding season between the two Urosaurus species but noted significant differences (although no actual statistics were presented) in clutch size between the two (see Table 3 for actual data). Comparisons of reproductive attributes of 9 9 of three arboreal desert lizards of Arizona appear in Table 3. Although sample size is too small for statistical comparisons, fat body weights of 9 9 (Table 4) do exhibit certain trends. Reproductive 9 9 from spring and early summer had smaller fat bodies than nonreproductive 9 9 from the same season. In view of Derickson's (1974) work on lipid deposition and utilization in Sceloporus graciosus, it is likely that the differences noted in U. graciosus result from fat body mobilization for egg production. Late summer 9 9 were much more variable in fat body weights (Table 4) most likely resulting from differences in number of clutches produced in that particular season by each individual 9 (for example, one 9 may have produced a spring clutch, and then a fall clutch, whereas another may have only produced a fall clutch). efReproductive Effort.-Reproductive fort, defined by Williams (1966) as the total amount of energy that is available to organisms that goes into reproductive re-

lated activities (mating, additionalcost associated with carrying eggs, and egg and shell production,etc.), is a central concern of evolutionaryecologists. The conceptwas initially discussedby Fisher (1930). In the past, the measurementof reproductiveeffort has been crude, involving simple ratios of clutch weights to body weights (Clark, 1970;Parkerand Pianka,1973;and others). Recent studies by Tinkle and Hadley (1973) and Vitt (1974b) indicate that interspecific differences in the amount of calories per unit weight of egg may exist. Since distinct differencesare apparentbetween egg calories per unit weight and body calories per unit weight (eggs being the energy richest life history stage), simple use of wet or dry weight ratios as suggested by Ballingerand Clark (1973) underestimates the energetic cost of reproduction. Caloric content of eggs and bodies was determinedon four U. graciosus 9 9 which containedoviducaleggs. Althoughthe sample is small, all studies thus far utilizing microbomb calorimetry for reproductive effort estimationshave shown no significant intraclutch variation in calories per unit weight, and no significant interclutch (within a population) variationin calories per unit weight (see Tinkle and Hadley, 1973; Ballinger and Clark, 1973; and Vitt, 1974b). Table 5 presents information on calories,percent ash, and water content for this sample. Mean calories per gram egg

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TABLE

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[Vol. 31, No. 1

5.-Energy (calories of ash-free dry weight, AFDW), ash content, and water content of bodies and eggs of the iguanid lizard Urosaurusgraciosus from the lower Colorado River. Only lizards with oviducal eggs were utilized for caloric determinations (see text). F-tests represent comparisonsof caloric determinationswithin each sample (3 determinations/pooled sample) to comparisonsbetween each sample (4 samples, 3 determinations/sample). As shown, no significant differences were found in caloriesper gram within and between samples.
Category N No. of caloric determinations x % ash
X

calories per gram

Range of individual means

F-test (.05)

x % water content

Bodies Eggs

4 4

12 12

19.38 6.95

5492 6612

5384-5620 6347-6770

F=

F = 2.10, 3/8 df 2.06, 3/8 df

70.50 56.20

weight (AFDW) of U. graciosus were higher than any species studied by Ballinger and Clark (1973), higher than the viviparous high latitude species Gerrhonotus coeruleus principis (Vitt, 1974b), and lower than the high elevation viviparous montane species Sceloporus jarrovi (Tinkle and Hadley, 1973). Reproductive effort (total clutch calories total body calories ratio) averaged 0.640 (range, 0.474-0.725) in U. graciosus and the total calories per egg averaged 638 (range, 438-771). Of the four experimental clutches, two contained four eggs and two contained five. The two larger clutches contained smaller eggs (in weight and total calories) than the two smaller clutches. Although these data were very limited, there may be a negative feedback between clutch size and egg size in this species. This could account for the lack in correlation between 9 size and clutch size. The wet ratio of clutch: body weight averaged 0.313 (range, 0.271-0.399) whereas the dry ratio was 0.531 (range, 0.403-0.604). In view of the differences in body and egg energy, body and egg ash, and caloric ratios and weight ratios, the ratio of clutch calories to body calories corrected for ash content serves to best estimate reproductive effort in species exhibiting no post-ovulatory contribution to eggs and no parental care. Reproductive Strategy.-Reproductive attributes of U. graciosus and growth rate information (Gates, 1963) suggest that U. graciosus conforms to the early maturing, short lived species presented by Tinkle et al. (1970). The production of many (3-10)

small eggs once or more per season, small adult body size (as compared to many iguanids) adds support to this and places U. graciosus in the r category of Pianka (1970) and MacArthur and Wilson (1967). Clutch weight to body weight (wet and dry) ratios are high as is the clutch calories to body calories ratio (reproductive effort) especially when number of clutches per season is considered, suggesting an r strategy. Data on survivorship would probably also indicate an r strategy but such data are not yet available.
Acknowledgments.-This study was supported by Bureau of Reclamation Grant No. 4-01-01-10310. The Colorado River Indian Reservation,California Department of Fish and Game, Arizona Department of Game and Fish, and the U. S. Fish and Wildlife Service supplied scientific collecting permits necessaryfor completion of this study. We thank Mike Walker and Harry Shankemanof Arizona State University for technical assistance and Justin D. Congdon (ASU) for critically reviewing the manuscript. In addition, criticisms by W. S. Parker,Mississippi University for Women, and two anonymous reviewers enabled us to clarify many aspects of the discussion of our data.
LITERATURE CITED

R. E. 1973. Experimental evidence of the tail as a balancing organ in the lizard Anolis carolinensis. Herpetologica 29:65-66. , AND D. R. CLARK. 1973. Energy content of lizard eggs and the measurement of reproductive effort. J. Herpetol. 7:129-132. BARNES, R. A. 1974. Competitive co-existence of two species of lizards, Genus Urosaurus. M.S. thesis. Arizona State Univ., Tempe. 92 p. B. H. 1965. Body temperaturesof BRATTSTROM, reptiles. Am. Midl. Nat. 73:376-422. CLARK, D. R. 1970. Age specific "reproductive effort" in the worm snake Carphophis vermis (Kennicott). Trans. KansasAcad. Sci. 73:20--24.
BALLINGER,

AMarch 1975]
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(Full page charges borne by authors)

Received: 21 October 1974 Accepted: 4 December 1974

Department of Zoology, Arizona State University, Tempe, Arizona 85281, USA

REPRODUCTION IN GUATEMALAN ANOLIS BIPORCATUS (SAURIA: IGUANIDAE) C. J. McCoy

HENDERSON (1972) noted the scarcity of information on reproduction in larger species of Anolis, and reported observations on egg-laying by a captive Anolis biporcatus. Henderson's anole laid two eggs within 7

hours, suggesting the unlikely possibility that A. biporcatus may have a normal clutch size of two eggs in parts of its range. A Carnegie Museum (CM) series of Anolis biporcatus from southeastern Guatemala

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