Comparative ecomorphology of the sandstone night lizard (Xantusia gracilis) and the granite night lizard (Xantusia henshawi)
expand article infoL. Lee Grismer
‡ La Sierra University, Riverside, United States of America
Open Access


It is hypothesized that shape differences between the closely related sandstone night lizard (Xantusia gracilis) and the granite night lizard (X. henshawi) may be correlated with structual differences in their respective microhabitats. Multivariate and univariate analyses of 22 morphometric characters taken from the head, body, and limbs of both saxicolus specialists recovered statistically significant differences between them with X. gracilis having a wider head, longer snout, larger eyes, wider sternum, higher and wider pelvis, thinner limbs, longer forearms and thighs, and longer hind limbs. Many of these same proportional differences have been reported among very closely related saxicolus species in other lizard families (i.e. Eublepharidae, Gekkonidae, Phrynosomatidae) that also live on different rocky substrates. This supports the inference that morphometric differences between X. gracilis and X. henshawi are ecomorphological adaptations for navigating the substantially different substrates of their respective microhabitats. Xantusia gracilis is restricted to a loose, heterogeneous, sandstone microhabitat composed of large boulders, small rocks, and cliff faces where cracks, crevices, holes, and exfoliations are used as retreats versus the compact, more homogeneous, granite boulder microhabitat of X. henshawi where narrow spaces beneath exfoliations and cap-rocks are the preferred retreats.

Key words

Anza-Borrego, ecomorph, microhabitat, scansorial, saxicolus, Xantusiidae


The concept that an animal’s form has evolved in response to the way it navigates its habitat underpins the study of ecomorphology—the intersection of organismal morphology, life history, and adaptation (Van der Klaauw 1948; Wainwright and Reilly 1994). Many studies have shown that locomotor performance is linked to morphology and strongly associated with structural features of the environment (i.e. ecomorphological specialization; Losos 1990, 2009, Luxbacher and Knouft 2009, Kaliontzopoulou et al. 2010a). Within diurnal lizards, ecomorphology has been well-studied in a number of ecologically diverse lineages of skinks, anoles, tropidurines, and wall lizards (Melville and Swain 2000; Bergmann and Irschick 2010; Kaliontzopoulou et al. 2010a; Losos 2010; Lee et al. 2013; Pincheira-Donoso and Meiri 2013; Pincheira-Donoso et al. 2015; Grismer et al. 2018; Toyama 2017) but comparatively little has been done on nocturnal lizards—and most of those involved only a single gekkonid genus (Grismer et al. 2015; 2017; 2020; Grismer and Grismer 2017; Nielson and Oliver 2017; Nguyen et al. 2021; Kaatz et al. 2021).

The Night Lizard family Xantusiidae is a New World lineage composed of three genera and 35 generally nocturnal species that collectively range from Southwestern North America to southern Central America (Noonan et al. 2013; Uetz et al. 2021). The family is dominated by highly cryptic, reclusive species with low vagility (Zweifel and Lowe 1966; Fellers and Drost, 1991; Mautz 1993) where high degrees of microhabitat specialization drive their differing ecomorphologies (Bezy 1989a, 1989b; Noonan et al. 2013). Within southern California, the Xantusiidae is represented by four, possibly five species of Xantusia bearing three trenchantly different ecomorphs and four different microhabitat preferences (Fig. 1): X. riversiana, an insular habitat generalist (Mautz 1993; Fellers and Drost 1991; Fellers et al. 1998); species of the X. vigilis complex (sec. Leavitt et al. 2007), vegetative specialists (Zweifel and Lowe, 1966); and X. gracilis and X. hensahwi, “rock-crevice-dwelling” (sec. Leavitt et al. 2007) specialists (Grismer and Galvin 1986; Lee 1975, respectively). Nonetheless, there have been only limited discussions extolling the ecomorphological differences between rock-crevice species and vegetation specialists within the genera Lepidophyma and Xantusia (Bezy 1967; Ramírez-Bautista et al. 2008; Adams et al. 2018) and only limited data have been generated to quantitatively evince the degree of morphological differences between the rock-crevice species X. henshawi—a granite exfoliation specialists (Lee 1975) and X. gracilis—a more generalized sandstone inhabitant (Grismer and Galvin 1986). Using a limited number of metrics, Grismer and Galvan (1986) noted differences in head proportions and the growth trajectories of body width and limb length, suggesting these were correlated with the more “terrestrial” life-style of X. gracilis. In the analyses herein, multivariate and univariate tests are used to quantify morphometric differences between X. gracilis and X. henshawi using an expanded data set comprised of 22 characters that have proven useful as ecomorphological indicators across a wide taxonomic range of lizard groups (Losos 1990, 2009, 2010; Melville and Swain 2000; Luxbacher and Knouft 2009, Bergmann and Irschick 2010; Kaliontzopoulou et al. 2010a; Lee et al. 2013; Pincheira-Donoso and Meiri 2013; Pincheira-Donoso et al. 2015; Grismer et al. 2015, 2018, 2020; Grismer and Grismer 2017; Nielson and Oliver 2017; Toyama 2017; Kaatz et al. 2021). These data are compared to similar data generated from distantly related lizard groups as leverage to generate preliminary hypotheses as to the adaptive significance of how the different ecomorphologies between X. gracilis and X. henshawi may correlate with their structurally different microhabitats.

Figure 1. 

A. The Island Night Lizard, Xantusia riversiana, from Santa Barbara Island, Santa Barbara County, California—a habitat generalist. Photo by Gary Nafis. B. Heterogeneous habitat of X. riversiana on Santa Barbara Island. Photo by Gary Nafis. C. The Yucca Night Lizard, X. cf. vigilis, from Anza-Borrego, San Diego County, California–a vegetative specialist. D. Yucca habitat of X. cf. vigilis at Blair Valley, San Diego County. E. The granite night lizard, X. henshawi, from Deep Creek, Riverside County, California– a granite exfoliation specialist. F. Granite boulder habitat of X. henshawi at Lost Valley, San Diego County. G. The sandstone night lizard, X. gracilis, from the Truckhaven Rocks, San Diego County—a sandstone specialist. H. Sandstone habitat of X. gracilis at the Truckhaven Rocks, San Diego County.

Materials and methods


Xantusia gracilis. Grismer and Galvin (1986) found X. gracilis to be restricted to a small (11.7 km2), circumscribed region known as the Truckhaven Rocks that fringe the eastern flanks of the Santa Rosa Mountains in the Anza-Borrego Desert State Park, San Diego County, California. The area is composed of crumbling sandstone and siltstone sediments that have undergone tremendous erosion, resulting in a number of slot canyons and massive, heavily weathered, titled outcroppings that form a complex microhabitat of boulders, rocks, and cliff faces bearing cracks, crevices, holes, and exfoliations (Fig. 2)—all of which are utilized by X. gracilis. This species is not found in the adjacent granitic habitat of the Santa Rosa Mountains that abut the Truckhaven Rocks (Grismer and Galvin 1986). As such, X. gracilis is considered a sandstone specialist.

Figure 2. 

A. The sandstone boulder and cliff face habitat of Xantusia gracilis, Truckhaven Rocks, San Diego County, California. B. Slot canyon microhabitat of X. gracilis, Truckhaven Rocks. C. Crumbling, exfoliation-like microhabitat of X. gracilis, Truckhaven Rocks. D. Relatively smooth, compact granite boulder microhabitat of X. henshawi showing a cap-rock (at top) and an exfoliation (in lower right), Mountain Meadows, San Diego County, California. E. Granite boulder outcrop habitat of X. henshawi, Aguanga, Riverside, County, California.

Xantusia henshawi. Lee (1975) indicated that the distribution and density of X. henshawi is closely tied to suitable rock crevices and exfoliations of granitic boulders throughout coastal and ecotonal regions in Riverside, San Diego, and Imperial counties, California and northern Baja California. Granite boulders lacking such features are uninhabitable, and thus the distribution of this species is fragmented in places (Lee 1975). For example, some hilly areas on the border of Riverside and San Diego counties bear extensive granitic outcroppings that lack exfoliations and cracks and do not harbor X. henshawi. Nor do areas where the granite boulders are extremely weathered and eroding, producing a course crumbling substrate, and crevices filled with decomposed granite. Xantusia henshawi is far more common in areas where the more compact, granitic surface of the boulders is relatively smooth and subject to cracking and exfoliating (Fig. 2). Specimens used in this analysis are listed in Table 1.

Table 1.

Locality of specimens of Xantusia henshawi used in this analysis. Based on distribution, all are presumed to belong to haploclade A of Lovich (2001) along with X. gracilis.

Catalog number County Locality
LSUHC 1530 Riverside Aguanga
LSUHC 1531 Riverside Aguanga
LSUHC 1998 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 1999 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2000 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2001 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2002 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2004 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2005 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2006 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2007 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2009 San Diego Intersection of Deer Springs Road and Interstate 15
LSUHC 2010 San Diego Ranchita
LSUHC 2011 San Diego Ranchita
LSUHC 2013 San Diego Ranchita
LSUHC 2015 San Diego Ranchita
LSUHC 2016 San Diego Ranchita
LSUHC 2017 Riverside Hemet
LSUHC 2018 Riverside Hemet
LSUHC 2019 Riverside Hemet
LSUHC 2020 Riverside Aguanga
LSUHC 2021 Riverside Aguanga

Mensural data

The following 22 measurements were taken from the left side of the body when possible to the nearest 0.1 mm using Mitutoyo dial calipers under a Nikon SMZ 1500 dissecting microscope:

SVL (snout-vent length) – measured from the tip of the snout to the cloacal opening.

HL (head length) – measured from the tip of the snout to the posterior margin of the retroarticular process of the mandible.

HW (head width) – measured across the widest portion of the head, posterior to the eyes.

HD (head depth) – measured from top of the head above the center of the orbit to the ventral surface of the mandibles.

SL (snout length) – measured from the anterior margin of the bony orbit to the tip of the snout.

SW (sternal width) – measured from the articulation point of the humeri and the glenoid fossae across the sternum from one side to the other.

ED (eyeball diameter) – measured in a horizontal plane from the posterior to the anterior margins of the eyeball.

FLW (forelimb width) – measured from the anterior and posterior insertion points of the forelimb on the body.

FL1 (brachial length) – measured from the articulation point of the humerus and the glenoid fossa to the distal margin of the elbow joint while flexed 90°.

FL2 (forearm length) – measured from the proximal margin of the elbow joint while flexed 90° to the distal margin of the ulna while the wrist joint is extended.

FL3 (manus length) – measured from the distal margin of the ulna to the base of the 4th finger.

FL4 (fourth finger length) – measured from the base of the fourth finger to the tip of the claw.

FLL (forelimb length) – the sum of FL1FL4. These data were used in separate analyses so as not to overleverage the analysis with forelimb dimensions.

AG (axilla-groin length) – measured from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body.

PW (pelvic width) – measured from the lateral margins of the ilia across of the body.

PH (pelvic height) – measured from top of the illium to the ventral surface of the pubic bone.

HLW (hind limb width) – measured from the anterior and posterior insertion points of the thigh on the body.

HL1 (thigh length) – measured from the articulation point of the femur and the acetabulum to the distal margin of the knee joint while flexed 90°.

HL2 (tibia length) – measured from the proximal margin of the knee joint while flexed 90° to the distal margin of the tibia while the ankle joint is extended.

HL3 (pes length) – measured from the distal margin of the tibia to the base of the 4th toe.

HL4 (fourth toe length) – measured from the base of the fourth toe to the tip of the claw.

HLL (hind limb length) – the sum of HL1HL4. These data were used in separate analyses so as not to overleverage the analysis with hind limb dimensions.

Data analysis

To ensure that allometric biases in the raw data were appropriately removed prior to analysis, hatchlings were omitted from the data set and the raw data were adjusted using the following equation: Xadj=log(X)-β[log(SVL)-log(SVLmean)], where Xadj=adjusted value; X=measured value; β=unstandardized regression coefficient for each population; and SVLmean=overall average SVL of all populations (Thorpe 1975, 1983; Turan 1999; Lleonart et al. 2000)—accessible in the R package GroupStruct (available at The morphometrics of each species were adjusted separately and then concatenated so as not to conflate intra- with interspecific variation (Reist 1985; McCoy et al. 2006). Small sample sizes for X. gracilis (females n=5, males n=6) recovered no sexual dimorphism in data that were non-parametric (Welch’s t-test; p>0.05). Lee (1975) also, noted no sexual dimorphism in X. henshawi for head and body dimensions. Although this may change with larger sample sizes bearing normally distributed data (Grismer in prep), data from both sexes were combined herein for each species. The raw and adjusted data are presented in Tables S1 and S2.

A principal component analysis (PCA) of the adjusted data was employed to visualize and assess the degree of difference in morphospatial clustering among Xantusia gracilis and X. henshawi. Principal component analysis is an unsupervised analysis that does not group individuals a priori according to species. All adjusted data were scaled to their standard deviation to ensure they were analyzed on the basis of correlation and not covariance. A subsequent supervised analysis, discriminant analysis of principal components (DAPC) from the ADEGENET package in R (Jombart and Collins 2015), that does group individuals a priori according to species, was also performed. A DAPC relies on data calculated from its own PCA as a prior step to ensure that variables analyzed are not correlated and number fewer than the sample size. Dimension reduction of the DAPC prior to plotting is accomplished by retaining the first set of principal components that account for 90–95% of the variation as determined from a scree plot generated as part of the analysis.

To search for and quantify significant morphological differences between Xantusia gracilis (n=11) and X. henshawi (n=22), all data were first tested for normality with an F-test. Those with homogeneous variances (p<0.05) were subjected to a Student’s t-test and those with unequal variances (p>0.05) were subjected to a Welch’s t-test. Both tests search for statistically significant different mean values (p<0.05) in all the characters between both species. Violin plots, embedded with boxplots, were generated for the characters bearing statistically different means in order to visualize their range and frequency of variation, mean, 50% quartile, and the degree of difference between the species. All data analyses were performed in R [v3.4.3] (R Core Team 2018).


The PCA demonstrates that Xantusia gracilis and X. hensahwi occupy non-overlapping positions in morphospace among the collective ordination of the first two principal components (Fig. 3A). Principal component 1 accounts for 24.3% of the variation and loads most heavily for limb width (FLW and HLW), thigh length (HL1), head width (HW) and snout width (SW) (Fig. 3C; Table 2). Principal component 2 accounts for an additional 15.3% of the variation and loads most heavily for head depth (HD), brachial length (FL1), fourth finger length (FL4), axilla-groin length (AG), pelvic height (PH), tibia length (HL2) and fourth toe length (HL4). Principal components 3 and 4 account for an additional 10.0% and 8.4%, of the variation, respectively (Fig. 3C; Table 2). Retention of the first seven eigenvalues of the DAPC, accounting for 93.3% of the variation, further recovered complete separation of X. gracilis and X. henshawi along the first discriminant function (Fig. 3B).

Figure 3. 

A. PCA and B. DAPC of Xantusia gracilis and X. henshawi. C. Bar plots of PCA loading scores of PC1–PC4.

Table 2.

Summary statistics and principal component analysis scores for the adjusted morphometric characters of Xantusia gracilis and C. X. henshawi. Abbreviations are listed in the Materials and methods.

PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9 PC10 PC11 PC12 PC13 PC14 PC15 PC16 PC17 PC18 PC19
Standard deviation 2.2054 1.7470 1.4168 1.2978 1.1505 1.0746 0.9953 0.9130 0.8209 0.8062 0.7670 0.7392 0.6512 0.5647 0.5257 0.4700 0.3956 0.3585 0.2336
Proportion of variance 0.2432 0.1526 0.1004 0.0842 0.0662 0.0577 0.0495 0.0417 0.0337 0.0325 0.0294 0.0273 0.0212 0.0159 0.0138 0.0111 0.0078 0.0064 0.0027
Cumulative proportion 0.2432 0.3958 0.4962 0.5804 0.6466 0.7043 0.7538 0.7955 0.8292 0.8617 0.8911 0.9184 0.9396 0.9556 0.9694 0.9804 0.9883 0.9947 0.9974
Eigenvalue 4.8639 3.0519 2.0074 1.6842 1.3237 1.1547 0.9907 0.8337 0.6739 0.6500 0.5883 0.5464 0.4241 0.3189 0.2763 0.2209 0.1565 0.1285 0.0546
SVL –0.0460 0.0228 –0.0248 –0.0261 0.0684 –0.1519 0.9766 –0.0130 0.0754 –0.0590 0.0090 –0.0255 0.0430 –0.0102 0.0425 –0.0083 0.0136 0.0200 –0.0063
SW –0.2892 –0.1317 –0.1023 0.0910 0.2057 0.4154 0.0675 –0.1512 –0.3248 –0.0858 –0.1846 –0.1978 –0.4010 0.0830 –0.0303 0.2506 0.2321 0.3123 0.2526
PW –0.2512 –0.0802 –0.1529 0.3721 0.1465 –0.0176 –0.0677 –0.3703 0.0745 –0.0421 0.1929 –0.4709 0.3354 –0.3291 0.1055 –0.0410 –0.2292 0.1113 –0.2007
PH –0.1867 –0.3630 –0.1312 –0.1903 –0.1488 0.0572 0.0208 0.4195 –0.0990 –0.0684 0.0696 0.1754 0.2863 –0.2852 –0.2235 0.3139 –0.3085 0.2742 0.1269
AG –0.1108 0.2409 0.3821 0.0879 –0.3953 0.0238 0.0179 0.1533 –0.1344 –0.0526 0.3443 –0.3094 0.2248 0.4206 0.1415 0.0975 –0.0471 0.1767 0.2479
HL –0.2116 0.0075 –0.4318 –0.0909 0.3720 0.0187 –0.0681 0.1262 0.1352 –0.2869 0.0702 0.0078 0.2013 0.5023 0.0509 0.1113 –0.1491 –0.3707 0.1272
HW –0.3053 –0.1526 0.0302 –0.1844 –0.1926 0.2856 0.0578 –0.1830 –0.3340 0.1723 –0.0038 0.1887 0.4262 –0.0951 0.0999 –0.1738 0.3573 –0.3721 –0.0292
HD 0.0081 –0.2928 –0.3123 –0.0469 –0.3221 –0.1252 0.0199 0.2661 0.0584 0.3856 –0.2227 –0.5662 –0.1361 0.1123 0.0306 –0.0414 0.0891 –0.2157 –0.0212
SL –0.2577 –0.1572 –0.2075 –0.0470 –0.0632 –0.5067 –0.0868 0.1029 –0.3064 –0.1576 0.1161 0.1626 –0.2036 0.0379 0.3259 –0.3725 0.0382 0.2865 –0.0889
ED –0.2577 –0.0667 0.2509 –0.2691 0.1566 0.2469 –0.0406 0.1892 0.4907 0.1218 –0.1264 0.0079 –0.0145 –0.0273 0.5796 0.0034 0.0185 0.1936 –0.1562
HLW 0.3574 0.0377 –0.2416 –0.2525 0.0224 0.1556 0.0336 –0.0038 –0.0648 0.2479 0.2246 –0.0562 0.0488 0.0310 –0.0119 0.2396 –0.0296 0.2114 –0.3282
HL1 –0.3529 0.1922 –0.0454 –0.0006 –0.0421 –0.0825 –0.0040 –0.0578 0.2445 0.4127 0.2082 0.1154 –0.2648 –0.2445 –0.1273 –0.0254 –0.2040 –0.1569 0.4699
HL2 –0.1740 0.3774 –0.1458 –0.1107 0.1231 0.3465 0.0597 0.2172 –0.1894 0.1996 0.1832 –0.0433 –0.1015 0.0880 –0.2176 –0.4541 –0.1989 0.1086 –0.2820
HL3 –0.2522 0.1909 0.0408 0.2413 0.1977 –0.3155 –0.0467 0.0682 –0.0081 0.4223 –0.3749 0.1487 0.3156 0.2341 –0.2189 0.1763 0.1736 0.2612 –0.1430
HL4 0.0042 0.3999 –0.3671 –0.0151 –0.2082 –0.0619 –0.0041 –0.1709 –0.1529 0.1224 0.0656 0.1736 –0.0619 –0.0824 0.4443 0.4465 –0.0003 –0.0407 –0.0312
FLW 0.3487 0.0654 –0.1820 –0.2334 0.2797 –0.0473 –0.0435 –0.0528 –0.0168 0.1550 0.0096 –0.1226 0.3234 –0.0815 0.1305 –0.2782 0.1859 0.2877 0.5633
FL1 –0.0687 0.3241 0.1363 –0.4109 –0.0156 –0.1196 –0.0281 –0.0762 –0.2560 –0.1797 –0.5469 –0.2552 0.0564 –0.1626 0.0184 0.0051 –0.3853 –0.0866 0.0239
FL2 –0.2166 0.1027 0.1193 –0.4759 0.1490 –0.2987 –0.0930 –0.0532 0.0827 –0.1257 0.3310 –0.2550 –0.0767 –0.1240 –0.2801 0.2112 0.4371 –0.0345 –0.1439
FL3 –0.1165 0.1300 –0.3238 –0.1213 –0.4945 0.1219 –0.0175 –0.2852 0.4354 –0.2624 –0.1847 0.0797 0.0758 0.1193 –0.2311 –0.1652 0.1268 0.3024 –0.0040
FL4 0.0129 0.3657 –0.1499 0.3011 –0.0147 0.0993 –0.0150 0.5446 0.0443 –0.2714 –0.0791 –0.0847 0.0657 –0.3969 0.0477 –0.0165 0.3661 –0.0863 0.0181

Although Xantusia gracilis and X. henshawi do not differ significantly in SVL, they do differ in a number of other proportions. The Student’s and Welch’s t-tests of the adjusted data recovered 11 characters bearing statistically significant mean differences between them (Fig. 4; Table 3), illustrating that X. gracilis has a significantly wider head (HW), longer snout (SL), larger eyes (ED), wider sternum (SW), thinner limbs (FLW and HLW, i.e. reduced limb mass), longer forearms (FL2), longer thighs (HL1), and a wider and higher pelvis (PW and PH, respectively). Welch’s t-test also demonstrated that X. gracilis has significantly longer hind limbs (HLL) than X. henshawi but not significantly longer forelimbs (FLL). Grismer and Galvin (1986) also noted that X. gracilis had a thinner body (inter-forelimb and inter-hind limb distances) than X. henshawi. Although X. gracilis has a longer snout, it does not have a significantly longer head and similarly, it has a significantly longer forearm, but does not have a significantly longer foreleg. These metrics indicate that not all components of the head and forelimb scale isometrically (i.e. they do not grow at the same rate).

Figure 4. 

Violin plots overlain with box plots of Xantusia gracilis and X. henshawi showing the range, frequency, mean (white dot), and 50% quartile (black rectangle) of the significantly different size-adjusted morphometric characters. The means of FLL are not significantly different.

Table 3.

Summary statistics of Student and Welch’s t-tests. Shaded cells are those characters bearing significantly different means between Xantusia gracilis and X. henshawi. Character abbreviations are in the Materials and methods.

character t-test t value p value
HL Welch’s 0.7763 0.4453
HW Welch’s 4.6329 6.828e-05
HD Student’s 0.3646 0.7170
SL Welch’s 3.1815 0.0033
SW Welch’s 4.6399 6.938e-05
ED Student’s 3.3674 0.002
FLW Welch’s -13.362 3.48e-12
FL1 Welch’s -0.1597 0.0742
FL2 Welch’s 2.3969 0.023
FL3 Welch’s 1.4885 0.1551
FL4 Welch’s -0.5856 0.5674
FLL Welch’s 1.0322 0.3137
AG Welch’s 1.7186 0.1078
PW Welch’s 0.0271 0.0056
PH Welch’s 2.8392 0.0083
HLW Welch’s -7.1422 9.989e-07
HL1 Student’s 4.2352 0.0002
HL2 Welch’s 0.3227 0.7504
HL3 Welch’s 1.9674 0.0646
HL4 Welch’s -0.9722 0.3424
HLL Welch’s 2.2286 0.0341


The quantification of ecomorphological differences among species is important to many downstream analyses based on body size and shape. These types of data are not just necessary for comparative methods that combine phylogenies with phenotypic data to understand evolutionary processes such as the mode and tempo of trait evolution (Mahler et al. 2010). They also apply to a multitude of ecological principles where phenotypic differences within and among populations can influence the rate and direction of evolution, population dynamics, and the outcome of several other community interactions (Bolker et al. 2003; Werner and Peacor 2003; Krohne 2018).

These data clearly demonstrate that Xantusia gracilis and X. henshawi are proportioned differently and do not scale equally across various components of their bodies. Overall, X. gracilis is more slender and less robust than X. henshawi (Fig. 5) as was first reported by Grismer and Galvin (1986) using a limited data set. They inaccurately suggested these differences correlated with a more “terrestrial” life style. Using an expanded, more highly partitioned data set, however, reveals precisely where such differences occur and aptly justifies functional ecomorphological comparisons and inferences across a wide range of distantly related lizard groups bearing the same traits. It is logical to posit that the morphological differences between X. gracilis and X. henshawi are somehow related to their abilities to navigate their respective microhabitats. It has been well-established in a number of small clades of very closely related lizard species, that longer and thinner limb components increase climbing performance and are ecomorphological adaptations for a scansorial life style, as opposed to shorter, more robust limbs that occur in their less scansorial or strictly terrestrial (non-climbing) close relatives (Grismer and Grismer 2017; Neilson and Oliver 2017; Grismer et al. 2021; Kaatz et al. 2021). We hypothesize here, that these same adaptations in X. gracilis help it to navigate a more complex heterogeneous and less firm, topographically complex sandstone substrate (i.e. sandstone is not necessarily a flat, smooth, hardened surface; Fig. 2C) as compared to a more homogeneous, relatively flat, hardened granite boulder substrate (Fig. 2D). The smaller eye diameter (and obligatorily flatter head) and shorter pelvic height of X. henshawi would be more adaptive for taking refuge within narrow, restrictive retreats beneath granitic exfoliations and cap rocks (Fig. 2D). Although both species share adaptions for a scansorial, saxicolus life style—trenchantly different from that of strictly terrestrial species (e.g. X. riversiana)—differences in the complexity of their associated microhabitats are likely the driver of their ecomorphologicial differences.

Figure 5. 

A. Adult male Xantusia gracilis, Truckhaven Rocks, San Diego County, California. B. Adult female X. gracilis, Truckhaven Rocks. C. Juvenile X. gracilis, Truckhaven Rocks. D. Adult male X. henshawi, Deep Canyon, Riverside County, California. E. Adult female X. henshawi, Ramona, San Diego County, California. F. Adult female X. henshawi, Deep Canyon.

What little work has been done looking at morphological adaptations related to climbing and clinging ability among closely related species living on different rock substrates is illustrative. In the eublepharid gecko genus Goniurosaurus, Grismer et al. (2021) demonstrated that the highly scansorial G. catbaensis, G. huuliensis, and G. luii, which are adapted to topographically complex karstic substrates, have significantly longer limbs, wider heads, longer snouts, and larger eyes than does the less scansorial G. lichtenfelderi that is adapted to a less complex granite substrate. Similarly, Cobos et al. (submitted), has shown experimentally, that reduced limb mass (i.e. thinner limbs), longer limbs, smaller hands and feet, and shorter digits, underpin the superior clinging ability of karst-adapted gekkonid species in the Cyrtodactylus pulchellus group as opposed to their closely related granite-adapted species. The point is, that very closely related saxicolus species navigating different rocky substrates are proportioned differently and that there may be a general convergence in relative limb and head proportions shared among those species that lead a more scansorial life style on topographically complex substrates.

The above research opens up several avenues of inquiry as to the locomotor performance between Xantusia gracilis and X. henshawi on their respective substrates as compared to that on one another’s substrate. Experimental trials such as those done by Cobos et al. (submitted) where climbing and clinging ability are measured on different substrates among different ecomorphs, could be used to test the morphology-based inference here that a complex sandstone habitat and a more homogenous granitic substrate require different ecomorphologies. Additionally, similar experiments can be designed to test the effects of unequal scaling in the various components (e.g. different limb and head dimensions) within functional systems (e.g. the limbs and skull) in order to ascertain what adaptive value different allometric trajectories may have during the course of ontogeny. Kaliontzopoulou et al. (2010a, 2010b) have shown that plasticity in limb trait variation in the saxicolus Podacris bocagei (Lacertidae) is under the influence of total body size. Whereas habitat-related differentiation is amplified for other limb characters when size effects are removed, and for others, it is completely eliminated or even reversed. Studying the ontogeny of limb traits in X. gracilis and X. henshawi (Grismer in prep.) may help to understand the putatively adaptive mechanisms underlying their ecomorphological differences.


I wish to thank Raquel Barbosa for taking the raw data and Amanda Kaatz, Jesse L. Grismer, and Evan S. H. Quah for several helpful discussions and comments concerning Night Lizards and ecomorphology. Bradford D. Hollingsworth (SDSNHM) provided helpful comments on the manuscript, sent me data on, and granted the loan of, specimens of Xantusia gracilis.


  • Adams NE, Dean MD, Pauly GB (2018) Morphological divergence among populations of Xantusia riversiana, a night lizard endemic to the Channel Island of California. Copeia 106: 550–562.
  • Bergmann PJ, Irschick DJ (2010) Alternate pathways of body shape evolution translate into common patterns of locomotor evolution in two clades of lizards: Body shape and locomotor evolution. Evolution 64: 1569–1582.
  • Bezy RL (1967) Variation, distribution, and taxonomic status of the Arizona Night Lizard (Xantusia arizonae). Copeia 3: 653–661.
  • Bezy RL (1989a) Night Lizards. The evolution of habitat specialists. Terra 28: 29–34.
  • Bezy RL (1989b) Morphological differentiation in unisexual and bisexual xantusiid lizards of the genus Lepidophyma in Central America. Herpetological Monographs 3: 61–80.
  • Chan KO, Grismer LL (2021) Correcting for ontogenetic size variation in morphometric analysis. Biological Journal of the Linnean Society in press.
  • Cobos JC, Grismer LL, Quah ESH, Anuar S, Ezpinoza REE (in prep) An introduction to Gollum’s Rule with an assessment of convergent evolution for cave-dwelling Malaysian Bent-toed Geckos.
  • Fellers GM, Drost CA (1991) Ecology of the island night lizard, Xantusia riversiana, on Santa Barbara Island, California. Herpetological Monographs 5; 28–78.
  • Fellers GM, Drost CA, Mautz WJ, Murphey T (1998) Ecology of the island night lizard, Xantusia riversiana, on San Nicolas Island, California. Prepared for the US Navy by the USGS, Western Ecological Research Center, Pt. Reyes Field Station, Pt. Reyes National Seashore, CA.
  • Grismer LL, Galvin MA (1986) A new night lizard (Xantusia henshawi) from a sandstone habitat in San Diego County, California. Transactions of the San Diego Society of Natural History 21: 155–165.
  • Grismer LL, Grismer JL (2017) A re-evaluation of the phylogenetic relationships of the Cyrtodactylus condorensis group (Squamata; Gekkonidae) and a suggested protocol for the characterization of rock-dwelling ecomorphology in Cyrtodactylus. Zootaxa 4300: 486–504.
  • Grismer LL, Ngo HN, Qi S, Ziegler T (2021) The phylogeny and taxonomy of Goniurosaurus (Squamata: Eublepharidae) and the correlation between habitat preference and ecomorphology in two species groups from karstic landscapes in northern Vietnam. Vertebrate Zoology 71: 335–352.
  • Grismer LL, Wood PL Jr, Quah ESH, Anuar S, Ngadi EB, Izam NAM, Ahmad N (2018) Systematics, ecomorphology, cryptic speciation and biogeography of the lizard genus Tytthoscincus Linkem, Diesmos & Brown (Squamata: Scincidae) from the sky-island archipelago of Peninsular Malaysia. Zoological Journal of the Linnean Society 183: 635–671.
  • Grismer LL, Wood PL Jr, Ngo VT, Murdoch ML (2015) The systematics and independent evolution of cave ecomorphology in distantly related clades of Bent-toed Geckos (Genus Cyrtodactylus Gray, 1827) from the Mekong Delta and islands in the Gulf of Thailand, Zootaxa. 3980: 106–126.
  • Grismer LL, Chan KO, Oaks JR, Neang T, Sokun L, Murdoch ML, Stuart BL, Grismer JL (2020) A new insular species of the Cyrtodactylus intermedius (Squamata: Gekkonidae) group from Cambodia with a discussion of habitat preference and ecomorphology. Zootaxa 4830: 75–102.
  • Kaatz A, Grismer JL, Grismer LL (2021) Convergent evolution of karst habitat preference and its ecomorphological correlation in three species of Bent-toed Geckos (Cyrtodactylus) from Peninsular Malaysia. Vertebrate Zoology in press.
  • Kaliontzopoulou A, Carretero MA, Llorente GA (2010a) Intraspecific ecomorphological variation: linear and geometric morphometrics reveal habitat-related patterns within Podarcis bocagei wall lizards. Journal of Evolutionary Biology 23: 1234–1244.
  • Kaliontzopoulou A, Carretero MA, Llorente GA (2010b) Sexual dimorphism in traits related to locomotion: ontogenetic patterns of variation in Podarcis wall lizards. Biological Journal of the Linnean Society 99: 530–543.
  • Krohne DT (2018) Ecology. Evolution, Application, Integration. Oxford Univsertity Press, New York. 552 pp.
  • Lee JC (1975) The autecology of Xantusia henshawi henshawi (Sauria: Xantusiidae). San Diego Society of Natural History Transactions 17: 259–278.
  • Lee MSY, Skinner A, Camacho A (2013) The relationship between limb reduction, body elongation and geographical range in lizards (Lerista, Scincidae). Journal of Biogeography 40: 1290–1297.
  • Leavitt DH, Bezy RL, Crandall KA, Sites Jr (2007) Multi-locus DNA sequence data reveal a history of deep cryptic vicariance and habitat-driven convergence in the desert night lizard Xantusia vigilis species complex (Squamata: Xantusiidae). Molecular Ecology 16: 4455–4481.
  • Lleonart J, Salat J, Torres GJ (2000) Removing allometric effects of body size in morphological analysis. Journal of Theoretical Biology 205: 85–93.
  • Losos JB (1990) Ecomorphology, performance capability, and scaling of west Indian Anolis lizards: an evolutionary analysis. Ecological Monographs 60: 369–388.
  • Losos JB (2010) Adaptive radiation, ecological opportunity, and evolutionary determinism: American Society of Naturalists E. O. Wilson Award Address. The American Naturalist 175: 623–639.
  • Luxbacher AM, Knouft JH (2009) Assessing concurrent patterns of environmental niche and morphological evolution among species ofhorned lizards (Phrynosoma). Journal of Evolutionary Biology 22: 1669–1678.
  • Mahler DL, Revell LJ, Glor RE, Losos JB (2010) Ecological opportunity and the rate of morphological evolution in the diversification of greater Antillean anoles. Evolution: 64: 2731–2745.
  • Mautz WJ (1993) Ecology and energetics of the Island Night Lizard, Xantusia riversiana, on San Clemente Island. In: Hochberg, FG (Ed), Third California Islands Symposium: Recent Advances in Research on the California Islands. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 417–428.
  • McCoy MW, Bolker BM, Osenberg CW, Miner BG, Vonesh JR (2006) Size correction: Comparing morphological traits among populations and environments. Oecologia 148: 547–554.
  • Melville J, Swain R (2000) Evolutionary relationships between morphology, performance, and habitat openness in the lizard genus Niveoscincus (Scincidae: Lygosominae). Biological Journal of the Linnean Society 70: 667–683.
  • Nguyen AT, Duong TA, Grismer LL, Poyarkov NA (2021) A new granite cave-dwelling Bent-toed Gecko from Vietnam of the Cyrtodactylus irregularis group (Squamata; Gekkonidae) and a discussion on cave ecomorphology. Vertebrate Zoology 71: 155–174.
  • Nielsen SV, Oliver PM (2017) Morphological and genetic evidence for a new karst specialist lizard from New Guinea (Cyrtodactylus: Gekkonidae). Royal Society Open Science 4: 170781.
  • Noonan BP, Pramuk JB, Bezy RL, Sinclair EA, de Queiroz K, Sites Jr (2013) Phylogenetic relationships within the lizard clade Xantusiidae: using trees and divergence times to address evolutionary questions at multiple levels. Molecular Phylogenetics and Evolution 69: 109–122.
  • Pincheira-Donoso D, Meiri S (2013) An intercontinental analysis of climate-driven body size clines in reptiles: no support for patterns, no signals of processes. Evolutionary Biology 40: 562–578.
  • Pincheira-Donoso D, Harvey LP, Ruta M (2015) What defines an adaptive radiation? Macroevolutionary diversification dynamics of an exceptionally species-rich continental lizard radiation. BMC Evolutionary Biology 15: 1–13.
  • R Core Team (2018) R: A language and environment for statistical computing. R Foundation for Statistical Computing. Vienna. Available from: (accessed 29 December 2018)
  • Ramírez-Bautista A, Vitt LJ, Ramírez-Hernández A, Quijano FM, Smith G (2008) Reproduction and sexual dimporphism of Lepidophyma sylvaticum (Squamata: Xantusiidae), a tropical lizard from Tlanchinol, Hidalgo, Mexico. Amphibia-Reptilia 29: 207–216.
  • Reist JD (1985) An empirical evaluation of several univariate methods that adjust for size variation in morphometric data. Canadian Journal of Zoology 63: 1429–1439.
  • Thorpe RS (1975) Quantitative handling of characters useful in snake systematics with particular reference to intraspecific variation in the Ringed Snake Natrix natrix (L.). Biological Journal of the Linnean Society 7: 27–43.
  • Thorpe RS (1983) A review of the numerical methods for recognising and analysing racial differentiation. In: Felsenstein J, ed. Numerical Taxonomy. NATO ASI Series (Series G: Ecological Sciences), vol. 1. Springer-Verlag, Berlin, 404–423.
  • Turan C (1999) A note on the examination of morphometric differentiation among fish populations: The Truss System, Turkish Journal of Zoology 23: 259–263.
  • Van der Klaauw C.J. (1948) Ecological studies and reviews. IV. Ecological morphology. Bibliotheca Biotheoretica 4, 27–111.
  • Wainwright PC, Reilly SM (1994) Ecological Morphology, Integrative Organismal Biology. The University of Chicago Press, Chicago, pp. 1–367.
  • Zweifel R.G., Lowe C.H. (1966) The ecology of a population of Xantusia vigilis, the Desert Night Lizard. American Museum Novitates 2247: 1–57.