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US ISSN 0006-9698
CAMBRIDGE,MASS.SEPTEMBER 16, 2013 NUMBER 536
FOUR NEW SPECIES OF CALIFORNIA LEGLESS LIZARDS (ANNIELLA)
THEODORE J. PAPENFUSS
1
AND JAMES F. PARHAM
2
ABSTRACT. A previous genetic study of the California legless lizard (Anniella pulchra) revealed five deep genetic
lineages and alluded to morphological differences among them. Here we show that three of these genetic lineages can
be readily diagnosed from topotypic A. pulchra through a combination of coloration, scalation, and skeletal
characters (trunk vertebra number). A fourth lineage is cryptic, but can be diagnosed from A. pulchra by its
karyotype. We argue that these genetic clades of A. pulchra are strong candidates for species recognition because they
exhibit properties that corroborate the DNA evidence for lineage separation. We therefore hypothesize that each of
the five genetic clades of A. pulchra (‘‘Anniella clades A–E’’) are distinct species and so describe four new species
(Anniella alexanderae, sp. nov., Anniella campi, sp. nov., Anniella grinnelli, sp. nov., and Anniella stebbinsi, sp. nov.).
In naming these new species we have chosen to honor four natural historians whose contributions to the study of
California’s vertebrate biodiversity are an ongoing inspiration for students of natural history and natural history
museum curators. Two of these new species have small and poorly characterized ranges in the San Joaquin Valley
and Carrizo Plain (A. alexanderae and A. grinnelli). A third restricted-range species (A. campi) is known from just
three sites in the eastern Sierra Nevada. The fourth new species (A. stebbinsi) is a wide-ranging cryptic lineage that
occurs throughout Southern California and into Baja California, Mexico. The limited distribution and fragile
habitats occupied by the new species of Anniella warrant additional scientific research and conservation attention.
KEY WORDS:Anniella pulchra; California; conservation; lizard; new species
INTRODUCTION
The genus Anniella Gray, 1852, includes
limbless lizards endemic to western North
America. Anniella is the last survivor of an
anguimorph lineage that first appeared in the
Eocene of the western interior (Gauthier,
1982; Smith, 2011). From the Miocene on,
the fossil record is restricted to within the
known range of extant Anniella in California
and Baja California, Mexico (Gauthier,
1980; Bell et al., 1995; Hunt, 2008a). The
populations traditionally assigned to the type
species, Anniella pulchra Gray, 1852, occur
1
Museum of Vertebrate Zoology, 3101 Valley Life
Sciences Building, University of California, Berkeley,
California 94720, U.S.A.; e-mail: asiaherp@berkeley.edu
2
John D. Cooper Archaeology and Paleontology
Center, Department of Geological Sciences, California
State University, Fullerton, California 92834, U.S.A.
EThe President and Fellows of Harvard College 2013.
from northwest Baja California, to the
eastern San Francisco Bay Area (Stebbins,
2003; Hunt, 2008c; Fig. 1). A second species,
Anniella geronimensis Shaw, 1940, is restricted
to coastal sand dunes in northwest Baja
California, Mexico. There is a limited area of
sympatry in Baja California in the vicinity of
San Quintin Bay (Shaw, 1953; Hunt, 2008b).
Like other fossorial taxa with reduced or
missing limbs, Anniella species are morpho-
logically conservative. Hunt (1984) per-
formed a study of morphological variation
among A. pulchra populations. His detailed
study, combined with our own observations,
confirms that there are few significant
differences in scalation among populations
of A. pulchra (Table 1). The major morpho-
logical variation within A. pulchra is color-
ation, with some coastal populations show-
ing dorsal melanism (Hunt, 1984, 2008a,c).
Previous genetic studies of A. pulchra show
that melanistic populations do not form a
monophyletic group or correspond to the
deepest genetics divergences (Bezy and
Wright, 1971; Bezy et al., 1977; Pearse and
Pogson, 2000; Parham and Papenfuss, 2009).
We would expect that the reduced vagility
associated with a subterranean ecology
would facilitate speciation. Indeed, numer-
ous genetic studies have shown that other
morphologically conservative, fossorial, rep-
tile taxa harbor cryptic species (e.g., Daniels
Figure 1. Map showing the traditional (inset) distribution of Anniella pulchra and a detail (main) showing the
hypothesized distribution of the newly described species. White stars indicate type localities, black dots show referred
specimens used in this study. Color-shaded areas are speculated based on the distribution of museum specimens and
genetic clades (Parham and Papenfuss, 2009) updated through the addition of MVZ 172784 from the southern Sierra
Nevada (A. campi, referred by morphology) and two genetically characterized specimens of A. pulchra sensu stricto:1)
SBMNH HE-2448 from within the Santa Barbara city limits; 2) MVZ 13376 (A. pulchra sensu stricto) based on data
from Pearse and Pogson (2000). Contact zones between A. pulchra and the newly described species in the Carrizo
Plain, San Joaquin Valley, Transverse Ranges, and Sierra Nevada are uncertain and so are not shaded. Scale
bars 5100 km.
2BREVIORA No. 536
et al., 2009; Mott and Vietes, 2009; Heide-
man et al., 2011). A similar pattern was not
revealed in Anniella until our range-wide
study of genetic variation among popula-
tions of Anniella (Parham and Papenfuss,
2009). In that paper, we reported mitochon-
drial and nuclear DNA sequences from 45
localities spanning the range of A. pulchra.
Our data revealed five genetic clades that can
be diagnosed with mitochondrial and nuclear
DNA markers. The maximum uncorrected
mitochondrial sequence divergence among the
five genetic lineages of A. pulchra ranges from
4.3%to 9.2%(Parham and Papenfuss, 2009).
This amount of divergence corresponds to
species level differences of other lizard genera
(reviewed by Papenfuss et al., 2001). But more
significantly, the genetic clades we uncovered
can be diagnosed with morphological charac-
ters, including previously unreported colora-
tion, vertebral counts from x-rays, or pub-
lished karyotypic differences. Thus, unlike the
aforementioned melanistic populations, the
deep genetic clades of A. pulchra are strong
candidates for species recognition because
they exhibit properties that corroborate the
genetic evidence for lineage separation (de
Queiroz, 2007). We therefore hypothesize that
each of the five genetic clades of A. pulchra
(Anniella clades A–E of Parham and Papen-
fuss [2009]) are distinct species.
Before describing four new species, it is
necessary to establish which of the five clades
will remain as A. pulchra. Murphy and Smith
(1991) designated a neotype for A. pulchra
because of problems associated with the
original description of the genus and species
noticed by Hunt (1983). The neotype of A.
pulchra, MVZ 64656, is from Pinnacles
National Monument, San Benito County,
California (Figs. 1, 2). A topotypic speci-
men, MVZ 247489, belongs to clade A,
which is distributed throughout Northern
California. Therefore, A. pulchra is now
TABLE 1. SUMMARY OF NINE SCALE CHARACTERS FOR THE ANNIELLA PULCHRA COMPLEX.
Anniella spp.
No. of Scales or Scale Rows
a
Supralabials Infralabials Dorsal Ant. Mid. Post. A. clear M. clear P. clear
Hunt (1984)
b
A. pulchra 5–7 5–8 198–250 31–36 28–32 22–26 4–8 4–7 3–5
A. stebbinsi 6–8 4–9 188–249 28–36 24–30 22–26 4–7 4–6 3–6
Holotypes
c
A. alexanderae 6 5 257 32 26 24 6 4 4
A. campi 5 5 244 35 32 27 5 4 4
A. grinnelli 6 5 239 30 25 23 7 5 6
A. stebbinsi 6 4 215 30 28 24 6 5 5
a
Supralabials: number of supralabials; Infralabials: number of infralabials; Dorsal: number of dorsal scales counted
along the midline; Ant.: number of anterior scale rows counted at two head lengths posterior to the interoccipital;
Mid.: number of scale rows at mid-body; Post: number of posterior scale rows counted at 10 scales anterior to vent;
A. clear: number of anterior clear scale rows (i.e., lacking dark pigmentation) between the dorsal and lateral stripes
counted at two head lengths posterior to the interoccipital; M. clear: number of clear scale rows (i.e., lacking dark
pigmentation) between the dorsal and lateral stripes at mid-body; P. clear: number of posterior clear scale rows (i.e.,
lacking dark pigmentation) between the dorsal and lateral stripes rows counted at 10 scales anterior to vent.
b
Summary of data from Hunt (1984) based on 102 A. pulchra and 614 A. stebbinsi. Data included are from the
clearly designated groups in that study that do not include more than one species. For A. pulchra, the groups included
are: 1–3, 4, 7, 10. For A. stebbinsi, the groups included are: 12, 14–27. The following groups were excluded because
they span geographic regions that may contain more than one species: 5, 6, 8, 9, 11, 13.
c
Data from the holotypes of the new species described in this paper.
2013 FOUR NEW SPECIES OF ANNIELLA 3
limited to populations in clade A of Parham
and Papenfuss (2009). As such, genetic clades
B–E are described as new species below. In
naming these new species we have chosen to
honor four natural historians whose contri-
butions to the study of California’s vertebrate
biodiversity are an ongoing inspiration for
students of natural history and natural history
museum curators.
MATERIALS AND METHODS
Museum abbreviations: CAS, California Acad-
emy of Sciences, San Francisco, California;
LACM, Natural History Museum of Los
Figure 2. A topotypic Anniella pulchra (MVZ 247488) in dorsal and ventral views, and the type locality for the
species at Pinnacles National Monument, San Benito County, California, U.S.A.
4BREVIORA No. 536
Angeles, Los Angeles, California; MCZ,
Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts; MVZ,
Museum of Vertebrate Zoology, University of
California, Berkeley, California; SBMNH,
Santa Barbara Museum of Natural History,
Santa Barbara, California; SDNHM, San
Diego Natural History Museum, San Diego,
California; UCMP, University of California
Museum of Paleontology, University of Cali-
fornia, Berkeley, California. Locality data for
all referenced material have been standardized
into metric units and for format. Original
locality information is available from the
repositories.
Specimens used in the study were collected
over a 14-year period. Anniella species are
fossorial and are rarely active on the surface.
Many of the sites sampled have no cover to
search such as logs, stones, or leaf litter. More
than 2,000 cover objects (flattened cardboard
boxes and pieces of plywood) were placed at
localities throughout the range of Anniella in
California. Nearly all of the new species
described here from sites north of the
Transverse Ranges were found by raking in
sandy soil under cover objects.
Molecular data used to delineate species
here are taken from Parham and Papenfuss
(2009). The samples from the Parham and
Papenfuss (2009) study include all of the new
holotypes and a topotype for A. pulchra.We
refer closely related samples of each species by
clade in the section called Referred Specimens
(see below). We refer to the molecular marker
sequenced by Parham and Papenfuss (2009),
NADH dehydrogenase subunit 2 and five
adjacent tRNAs (trnWANCY), as well as
parts of cytochrome oxidase 1 and trnM,as
‘‘ND2’’ hereafter. Coloration data were taken
from living specimens (Appendix) observed
under a sunlight spectrum Bell & Howell lamp
using a Munsell Book of Color (Munsell
Color Company, 1976) and RGB Hexadeci-
mal color conversions (Kelly and Judd, 1955)
listed as ‘‘(Munsell, RGB).’’ Ventral colora-
tion characters can only be observed in fresh
specimens, whereas characters relating to the
lateral stripes can be observed in preserved
specimens. Vertebral counts were taken from
x-rays (Appendix). Morphological abbrevia-
tions: SVL, snout-vent length; TL, tail length.
Scale names in the descriptions are based on
Smith (1946, p. 466). Scalation of the new
holotypes was compared with a large survey
of A. pulchra complex populations (Hunt,
1984; Table 1). Diagnostic characters are
presented in Table 2.
TABLE 2. DIAGNOSTIC CHARACTERS FOR THE ANNIELLA PULCHRA COMPLEX.
BOLD CHARACTER STATES DIAGNOSE THE NEW SPECIES FROM A. PULCHRA.
A. pulchra A. alexanderae A. campi A. grinnelli A. stebbinsi
Ventral color Yellow Grey Yellow Purple Yellow
Lateral stripe ,Single Single Double Single ,Single
Mean vertebral count ,77 (76.0) .81 (82.2) ,77 (76.5) .81 (81.2) ,77 (75.4)
Dorsal scales #250 (198–250) .250 (252–278) ,250 (219–244) ,250 (234–249) ,250 (188–249)
Mean dorsal scales 222.3 261.2 227.6 242.4 213.7
Chromosomes 2n 520 ? ? ? 2n =22
Maximum ND2
divergence from A.
pulchra (%)
a
—8.0 8.4 9.2 8.7
a
Max. ND2 divergence from A. pulchra is the maximum sequence divergence from A. pulchra based on the
mitochondrial DNA marker used by Parham and Papenfuss (2009). See descriptions and diagnoses of each species
for more details.
2013 FOUR NEW SPECIES OF ANNIELLA 5
Anniella alexanderae, new species
Temblor Legless Lizard
Figure 3
Anniella pulchra lineage B—Parham and
Papenfuss, 2009.
Holotype. MVZ 250570, an adult male
from 35.20906N, 119.56726W (380 m eleva-
tion [elev.]; Figs. 1, 4), Shale Rd., 1.3 km S
(by road) junction with Hwy. 33, Kern
County, California, U.S.A., collected on
February 21, 2005, by Theodore J. Papenfuss
and James F. Parham.
Paratypes. CAS 238588, an adult male
from 35.21016N, 119.56706W (375 m elev.;
Figs. 1, 4), Shale Rd., 1.3 km S by road of
the junction with Hwy. 33, Kern County,
California, U.S.A., collected on October 2,
2007, by Theodore J. Papenfuss; MCZ R-
189386 and MVZ 267237, both adult males
from 35.20926N, 119.56716W (413 m elev.;
Figs. 1, 4), Shale Rd. 1.3. km S by road of
the junction with Hwy. 33 (Figs. 1, 4), Kern
County, California, U.S.A., collected on
April 18, 2010, by Theodore J. Papenfuss;
Figure 3. Four new species of Anniella and their diagnostic characters. Upper left, Anniella alexanderae: dorsal
(MVZ 250549, paratype); ventral view showing the diagnostic gray coloration (MVZ 257720, paratype). Upper right,
Anniella campi: dorsal (MCZ-R-189380, paratype); detail (MVZ 257277, holotype) showing diagnostic double dark
lateral stripes. Lower left, Anniella grinnelli: ventral (MVZ 247487, paratype) showing diagnostic purple coloration;
dorsal (MVZ 267228, paratype). Lower right, Anniella stebbinsi: dorsal (MVZ 250558, paratype); ventral (MVZ
267248). Center: comparison of ventral coloration from three of the new species. Left, A. grinnelli (MVZ 250546,
paratype); center, A. alexanderae (MVZ 250549, paratype); right, A. stebbinsi (MVZ 250558, paratype).
6BREVIORA No. 536
MVZ 250549 (Fig. 3), an adult male from
35.20906N, 119.56666W (380 m elev.;
Figs. 1, 4), Shale Rd. 1.3. km S by road of
the junction with Hwy. 33, Kern County,
California, U.S.A., collected on October 21,
2005, by Theodore J. Papenfuss; MVZ
257720 (Fig. 3), an adult, not sexed, from
35.20906N, 119.56666W, (380 m elev.;
Figs. 1, 4), Shale Rd., 1.3 km S by road of
the junction with Hwy. 33, Kern County,
California, U.S.A., collected April 2, 2007,
by Theodore J. Papenfuss.
Referred Specimens. Additional specimens
listed in the Appendix of this study and from
clade B of Parham and Papenfuss (2009;
localities 22 and 23 in the appendix of that
study).
Diagnosis. Distinguished from all other
species of the A. pulchra complex by a unique
ventral coloration of Light Gray (5Y 7/1,
RGB #D3D3D3) that is continuous from
the insertion of the lower jaw to the end of
the tail. This coloration is present in all
paratypes and referred specimens. It is
further distinguished from A. pulchra,An-
niella stebbinsi, and Anniella campi by its
higher vertebral count (Fig. 5) and from all
species of the complex by its higher dorsal
scale count (Tables 1, 2). Anniella alexan-
derae shows a maximum mitochondrial
Figure 4. Type localities of the four new species of Anniella. Upper left, Anniella alexanderae: Shale Rd., 1.3 km
S (by road) junction with Hwy. 33, Kern County, California, U.S.A. Upper right, Anniella campi: Big Spring, 5.8 km
NW junction Hwy. 14 (by Hwy. 178) Kern County, California, U.S.A. Lower left, Anniella grinnelli: Jack Zaninovich
Memorial Nature Trail, Sand Ridge Preserve, Kern County, California, U.S.A. Lower right, Anniella stebbinsi:El
Segundo Dunes, Los Angeles International Airport, Los Angeles County, California, U.S.A.
2013 FOUR NEW SPECIES OF ANNIELLA 7
sequence divergence (for ND2, see Materials
and Methods) from A. pulchra of 8.0%, from
A. grinnelli of 6.0%, from A. campi of 4.9%,
and from A. stebbinsi of 4.9%(Parham and
Papenfuss, 2009).
Description (Based on Holotype). Adult
male, 158 mm SVL, 81 mm TL; 81 trunk
vertebrae. Coloration in life: dorsal color
Pale Olive (5Y 6/4, RGB #A79367), lateral
color Strong Orange (5YR 7/12, RGB
#A85400), ventral color ventral Light Gray
(5Y 7/1, RGB #D3D3D3); a mid-dorsal
black stripe one-third scale wide is present
from the parietals to the tip of the tail; lateral
black stripes one-third scale wide are present
from the eye to the tip of the tail.
Rostral large, visible from above; posteri-
or tip pointed and in contact with prefrontals
in a slight groove at anterior suture of
prefrontals; supraoculars 3-3; preoculars
1-1; postoculars 2-2; occipitals 2-2; supralabials
6-6, first small and located directly beneath
nasal with posterior edge in contact with
second supralabial, second largest, third and
fourth half the length of second and in contact
with eye, fifth equal in size to third and fourth
and in contact with postoculars; anterior two-
thirds of mental contacts first pair of infra-
labials, posterior pointed one-third of men-
tal inserts in a groove between postmentals;
infralabials 5-5; 32 scale rows two head lengths
posterior to the interoccipital, 26 scale rows at
mid-body; 24 scale rows counted 10 scales
anterior to vent; six clear (no dark pigment
from stripes) scale rows between dorsal and
lateral stripe on right side of body two head
lengths posterior to the interoccipital; four clear
scale rows at mid-body, four clear scale rows at
a point 10 scales anterior to vent; 257 dorsal
body scales counted along right side of mid-
dorsal line from posterior border of interoccip-
ital to a point above the vent.
Distribution. This species is known from
two sites separated by continuous suitable
habitat west of Hwy. 33. The known sites are
in areas of sandy soil at the southeast base of
the Temblor Range between McKittrick and
Taft on the west side of the Southern San
Joaquin Valley in Kern County, California
(Fig. 1). All specimens have been found
between California State Highway 33 and
the Temblor Range. Detailed searches, in-
cluding multi-year use of cover boards, have
failed to yield Anniella in apparent suitable
habitat on the floor of the San Joaquin
Valley east of Highway 33.
Natural History. All specimens were found
under cover boards and flattened cardboard
boxes placed on sandy soil. This species is
most easily found between February and
March when the soil is damp. The known
range is in an arid part of California (average
annual rainfall at nearby McKittrick is just
184 mm).
Etymology. This species is named after the
naturalist Annie Montague Alexander (1867–
1950; Fig. 6), who collected thousands of
botanical, paleontological, and zoological
specimens from western North America and
provided intellectual support and crucial en-
dowments for both the Museum of Vertebrate
Zoology and the Museum of Paleontology at
Figure 5. Box and whisker plots (range, sample
standard deviation, mean) of trunk vertebral counts of
Anniella pulchra and the four new species based on x-ray
images of museum specimens. Anniella alexanderae and
Anniella grinnelli show higher vertebral counts than the
other three species of the A. pulchra complex. Note that
a single outlier with a trunk vertebral count of 74 (the
rest are 79–84) affects the range of A. grinnelli.
8BREVIORA No. 536
the University of California at Berkeley (Stein,
2001).
Anniella campi, new species
Southern Sierra Legless Lizard
Figure 3
Anniella pulchra lineage D—Parham and
Papenfuss, 2009.
Holotype. MVZ 257727 (Fig. 3) from
35.62516N, 117.95816W (1,230 m elev.;
Figs. 1, 4), Big Spring, 5.8 km NW Junction
Hwy. 14 (by Hwy. 178) Kern County,
California, U.S.A., collected on March 31,
2006, by Theodore J. Papenfuss.
Paratypes. CAS 233827, an adult male,
233828, an adult female, from 35.62526N,
117.95816W (1,240 m elev.; Figs. 1, 4), Big
Spring, 5.8 km NW junction Hwy. 14 (by Hwy.
178) Kern County, California, U.S.A., collected
on March 31, 2006, by Theodore J. Papenfuss;
MCZ R-189380 (Fig. 3), 189381, 189382, adults
not sexed from 35.62526N, 117.95816W
(1,240 m elev.; Figs. 1, 4), Big Spring, 5.8 km
NW junction Hwy. 14 (by Hwy. 178) Kern
County, California, U.S.A., collected on May 7,
2011, by Theodore J. Papenfuss.
Referred Specimens. MVZ 172784 (Kern
County, California, U.S.A.), additional spec-
imens listed in the Appendix of this study
and from clade D of Parham and Papenfuss
(2009; localities 28 and 29 in the appendix of
that study).
Diagnosis. Distinguished from all other
species of the Anniella pulchra complex by a
unique color pattern consisting of continuous,
double, dark lateral stripes from the side of the
head to the tip of the tail. This character is
present in all paratypes and referred specimens.
Anniella campi shows a maximum mitochon-
drial sequence divergence (for ND2, see Mate-
rials and Methods) from A. pulchra of 8.4%,
from A. grinnelli of 5.8%,fromA. alexanderae
of 4.9%, and from A. stebbinsi of 4.3%.
Description (Based on Holotype). Adult
male, SVL 152 mm, regenerated TL 60 mm,
74 trunk vertebrae. Coloration in life: dorsal
color Yellowish Gray (2.5Y 7/2, RGB
#C1AE96), lateral color Vivid Yellow (5Y
7/12, #DBA600), ventral color Vivid Yellow
(5Y 8/14, RGB #FBCE00).
Rostral large, visible from above. Posteri-
or tip pointed and in contact with prefrontals
Figure 6. Four natural historians whose contributions to the study of California’s vertebrate biodiversity are
honored by the new species of Anniella described here. Left, Annie Montague Alexander (1867–1950) on expedition
collecting Pleistocene fossils at Fossil Lake, Lake County, Oregon, in 1901 (from the UCMP archives). Center left,
Charles Lewis Camp (1893–1974) on expedition in San Juan County, Utah, in 1942 (from the UCMP archives) next
to the holotype of the Permian tetrapod Tseajaia campi Vaughn, 1964. Center right, Joseph Grinnell (1877–1939) on
expedition collecting vertebrates in Imperial County, California, in 1910 (from the MVZ archives). Robert Cyril
Stebbins (1915–) with an Ensatina salamander on the University of California at Berkeley campus in 1951 (from the
MVZ archives).
2013 FOUR NEW SPECIES OF ANNIELLA 9
in a slight groove at anterior suture of
prefrontals; supraoculars 3-3; preoculars
1-1; postoculars 2-2; occipitals 2-2; suprala-
bials 5-5, first small and located directly
beneath nasal with posterior edge in contact
with second supralabial, second largest, third,
fourth, and fifth half the length of second,
third and fourth in contact with eye; anterior
two-thirds of mental contacts first pair of
infralabials, posterior pointed one-third of
mental inserts in a groove between postmen-
tals; infralabials 5-5; 35 scale rows two head
lengths posterior to the interoccipital, 32 scale
rows at mid-body; 27 scale rows counted 10
scales anterior to vent; five clear scale rows
between dorsal and lateral stripe on right side
of body two head lengths posterior to the
interoccipital; four clear (no dark pigment
from stripes) scale rows at mid-body, four clear
scale rows at a point 10 scales anterior to vent;
244 dorsal body scales counted along right side
of mid-dorsal line from posterior border of
interoccipital to a point above the vent.
Distribution. Anniella campi is only known
from thee localities along the western edge of
the Mojave Desert in Kern and Inyo counties
(Fig. 1). The Big Spring locality is a perma-
nent spring that supports a small area of
suitable habitat, estimated at less than
2 hectares, in an otherwise desert environ-
ment. The Anniella population here is clearly
relictual since there is no other suitable
habitat in the area. Parham and Papenfuss
(2009) reported a second along Nine Mile
Canyon Road in southern Inyo County,
north of Big Spring. A third locality, south
of Big Spring in Kern County is represented
by a museum specimen that shows the
diagnostic character of the complete double
lateral stripes (MVZ 172784). Specimens
have been found crossing the road at night
(Robert W. Hansen, personal communica-
tion). It is likely that this species will be
found in canyons between Big Spring and
Nine Mile Canyon.
Natural History. This species is locally
common at Big Spring, where specimens
have been collected by raking under debris
that has accumulated at the base of Chamisa
(Ericameria nauseosa) that grow adjacent to
the spring. Specimens have been found in
April and May.
Etymology. This species is named after
Charles Lewis Camp (1893–1974; Fig. 6),
former student at the Museum of Vertebrate
Zoology and later director of the University
of California Museum of Paleontology. On
a 1915 collecting expedition to Yosemite
National Park with Joseph Grinnell, he
discovered the Mt. Lyell salamander, Hydro-
mantes platycephalus (Camp, 1916), part of
a lineage that is otherwise restricted to the
Old World and therefore one of the more
significant herpetological discoveries in
North America. Charles Camp also partici-
pated in successful paleontological expedi-
tions throughout western North America, as
well as Africa, Australia, and South Amer-
ica. Camp’s (1923) influential ‘‘Classification
of the lizards’’ formed the foundation for
modern taxonomy of squamates (Estes and
Pregill, 1988).
Anniella grinnelli, new species
Bakersfield Legless Lizard
Figure 3
Anniella pulchra lineage C—Parham and
Papenfuss, 2009.
Holotype. MVZ 257714, from 35.30546N,
118.80136W (254 m elev.; Figs. 1, 4), Jack
Zaninovich Memorial Nature Trail, Sand
Ridge Preserve, Kern County, California,
U.S.A., collected on April 11, 2007, by James
F. Parham and Theodore J. Papenfuss.
Paratypes. CAS 234253, an adult male,
CAS 234254 and 234255, both adult females,
from 35.38946N, 119.06976W (130 m elev.), in
a field 0.8 km N of Rosedale Hwy. by Fruitvale
Ave then 0.3 km E of the end of Price
Way, Bakersfield, Kern County, California,
10 BREVIORA No. 536
collected on February 1, 2006, by James F.
Parham and Theodore J. Papenfuss; MCZ R-
189378, R-189379, adult males from
35.39006N, 119.06086W (125 m elev.),
0.65 km N of Rosedale Hwy. by Landco Dr.,
then 0.15 km W at end of Gilmore Ave.,
Bakersfield, Kern County, California, col-
lected on April 21, 2010, by Theodore J.
Papenfuss; MVZ 247487 (Fig. 3), an adult, not
sexed, from 35.38946N, 119.06976W(120m
elev.) in a field 0.8 km N of Rosedale Hwy. by
Fruitvale Ave then 0.3 km E of the end of Price
Way, Bakersfield, Kern County, California,
U.S.A., collected on April 27, 2002, by
Theodore J. Papenfuss; MVZ 250546
(Fig. 3), an adult female from 35.39006N,
119.06086W(125melev.),0.65kmNof
Rosedale Hwy. by Landco Dr., then 0.15 km
W at end of Gilmore Ave., Bakersfield, Kern
County, California, collected on April 21,
2010, by Theodore J. Papenfuss; MVZ
267228 (Fig. 3), an adult, not sexed, from
35.38946N, 119.06976W (120 m elev.) in a field
0.8 km N of Rosedale Hwy. by Fruitvale Ave.
then 0.3 km E of the end of Price Way,
Bakersfield, Kern County, California, U.S.A.,
collected on March 17, 2005, by Theodore J.
Papenfuss.
Referred Specimens. Additional specimens
listed in the Appendix of this study and from
clade C of Parham and Papenfuss (2009;
localities 24 through 27 in the appendix of
that study).
Diagnosis. Distinguished from all other
species of Anniella by a unique ventral
coloration of Grayish Red (2.5R 4/2, RGB
#755A61). This coloration is continuous
from the anterior end of the lower jaw to
the end of the tail and is present in all
paratypes and known specimens. It is further
distinguished from A. pulchra,A. stebbinsi,
and A. campi by its higher vertebral count
(Fig. 5). Anniella grinnelli shows a maximum
mitochondrial sequence divergence (for
ND2, see Materials and Methods) from A.
pulchra of 9.2%, from A. stebbinsi of 6.4%,
from A. alexanderae of 6.0%, and from A.
campi of 5.8%.
Description (Based on Holotype): Adult
male, 148 mm SVL, 93 mm TL; 79 trunk
vertebrae. Coloration in life: dorsal color
Light Olive Gray (7.5Y 5/2, RGB #837A67),
lateral color Strong Orange (5YR 7/12, RGB
#A85400), ventral color Grayish Red (2.5R
4/2, RGB #755A61 [appears purple]); a mid-
dorsal black stripe one-half scale wide is
present from the parietals to the tip of the
tail; lateral black stripes one scale wide are
present from the eye to the tip of the tail.
Rostral large, visible from above, posteri-
or side flat and in contact with prefrontals;
supraoculars 3-3; preoculars 1-1; postoculars
2-2; occipitals 2-2; supralabials 6-6, first
small and located directly beneath nasal,
second largest, third and fourth half the
length of second and in contact with eye;
mental rounded and broadly in contact with
first pair of infralabials and postmentals;
infralabials 5-5; 30 scale rows two head
lengths posterior to the interoccipital, 25
scale rows at mid-body; 23 scale rows
counted 10 scales anterior to vent; seven
clear (no dark pigment from stripes) scale
rows between dorsal and lateral stripe on
right side of body two head lengths posterior
to the interoccipital; five clear scale rows at
mid-body, six clear scale rows at a point 10
scales anterior to vent; 239 dorsal body scales
counted along right side of mid-dorsal line
from posterior border of interoccipital to a
point above the vent.
Distribution. This known range of A.
grinnelli is restricted to the southern San
Joaquin Valley and the east side of the
Carrizo Plain (Fig. 1). Specimens have been
collected within the city limits of Bakersfield.
During the last 10 years, two of the three
known Bakersfield populations were de-
stroyed by housing development. A protected
population is located at the type locality, the
2013 FOUR NEW SPECIES OF ANNIELLA 11
Sand Ridge Preserve. Individuals from the
Carrizo Plain are similar in coloration and
mitochondrial sequences to San Joaquin
samples but have a nuclear genotype known
only from lineage B. Parham and Papenfuss
(2009) speculated that this population may be
a hybrid or intergrade population, but we
include Carrizo specimens in our concept of
A. grinnelli (Fig. 1).
Natural History. All specimens were found
under cover objects (plywood scraps and
flattened cardboard boxes) that had been
placed on sandy soil. The type locality is a
stable sand dune of Pleistocene origin
(Fig. 4).
Etymology. This species is named after
Joseph Grinnell (1877–1939; Fig. 6), the first
director of the Museum of Vertebrate
Zoology at the University of California at
Berkeley. Joseph Grinnell published hun-
dreds of scientific papers based on his
extensive collecting and surveys in western
North America, and developed the Grinnell
Method of note taking that has become the
standard for natural history observations.
Anniella stebbinsi, new species
Southern California Legless Lizard
Figure 3
Anniella pulchra lineage E—Parham and
Papenfuss, 2009
Holotype. MVZ 267246, from 33.95006N,
118.44156W (24 m elev.; Figs. 1, 4), El
Segundo Dunes, Los Angeles International
Airport, Los Angeles County, California,
U.S.A., collected on April 20, 2010, by
Theodore J. Papenfuss.
Paratypes. MVZ 267247, a subadult male
collected with the holotype; MVZ 250558
(Fig. 3), a subadult male from 34.00426N,
118.81006W (5 m elev.), Point Dume, Los
Angeles County, California, U.S.A., col-
lected on November 24, 2005, by Theodore
J. Papenfuss. MVZ 267248 (Fig. 3), from
33.90156N, 116.74476W (470 m elev.), 4.0 km
SE (airline) of Cabazon, Riverside County,
California, U.S.A., collected on March 19,
2005, by Theodore J. Papenfuss.
Referred Specimens: LACM 64583 (kar-
yotyped specimen [Bezy et al., 1977] from the
type locality in Los Angeles County, Cali-
fornia, U.S.A.), SDNHM 42040, 42041,
42876, 42878, 42879 (San Diego County,
California, U.S.A.), additional specimens
listed in the Appendix of this study, and
from clade E of Parham and Papenfuss
(2009; localities 30–45 in the appendix of
that study, but excluding the A. geronimensis
from locality 41 [Colonia Guerrero]).
Diagnosis. Distinguished by its yellow
ventral coloration from A. grinnelli, which
has a purple (grayish-red) ventral coloration
and from A. alexanderae, which has a light
gray ventral coloration. Distinguished from
A. pulchra which also has a yellow ventral
coloration by a somatic chromosome num-
ber of 2n 520 rather than 2n 522 (Bezy
et al., 1977). Distinguished from A. campi,
which also has a yellow ventral coloration by
a single dark lateral stripe on each side rather
than a double lateral stripe. Some specimens
of A. stebbinsi have a double lateral stripe,
but it is never continuous or exceeds 50%of
the combined body and tail length, whereas
in A. campi it is continuous and extends to
the tip of the tail. Anniella stebbinsi shows a
maximum mitochondrial sequence diver-
gence (for ND2, see Materials and Methods)
from A. pulchra of 8.7%, from A. grinnelli of
6.4%, from A. alexanderae of 4.9%, and
from A. campi of 4.3%.
Description (Based on Holotype). Adult
female, SVL 132 mm, regenerated TL 81 mm,
79 trunk vertebrae. Coloration in life: dorsal
color Light Olive Brown (2.5Y 5/2, RGB
#8B7863), lateral color Strong Yellow (RGB
#E1A129), ventral color Moderate Yellow
(5Y 7/8, RGB #CFA639); mid-dorsal black
stripe less than one scale wide is present from
the parietals to the tip of the tail; lateral
12 BREVIORA No. 536
black stripes one scale wide are present from
the eye to the tip of the tail.
Rostral large, visible from above, posteri-
or tip pointed and in contact with prefrontals
in a slight groove at anterior suture of
prefrontals; supraoculars 3-3; preoculars 1-
1; postoculars 2-2; occipitals 2-2; suprala-
bials 6-6, first small and located directly
beneath nasal with posterior edge in contact
with second supralabial, second largest, third
and fourth two-thirds the length of second
and in contact with eye; mental rounded,
broadly in contact with first pair of infra-
labials and postmentals; infralabials 4-4; 30
scale rows two head lengths posterior to the
interoccipital, 28 scale rows at mid-body; 24
scale rows counted 10 scales anterior to vent;
six clear (no dark pigment from stripes) scale
rows between dorsal and lateral stripe on
right side of body two head lengths posterior
to the interoccipital; five clear scale rows at
mid-body, five clear scale rows at a point 10
scales anterior to vent; 215 dorsal body scales
counted on right side of mid-dorsal line from
posterior border of interoccipital to a point
above the vent; 215 dorsal body scales
counted along right side of mid-dorsal line
from posterior border of interoccipital to a
point above the vent.
Distribution. Throughout Southern Cali-
fornia south of the Transverse Ranges into
northern Baja California, Mexico (Fig. 1).
Populations in the Tehachapi and Piute
mountains of Kern County are disjunct from
the main distribution of this species to the
south. Therefore, the distribution of A. steb-
binsi is presumably bisected by southern
populations of A. pulchra ranging from the
Santa Barbara region into the Antelope Valley
of the western Mojave Desert (Fig. 1; Parham
and Papenfuss, 2009). Based on the bulk of
their hypothesized range, we recommend the
common names of ‘‘Northern California
legless lizard’’ for A. pulchra and ‘‘Southern
California legless lizard’’ for A. stebbinsi.
Natural History. Anniella stebbinsi is found
in a broader range of habitats that any of the
other species in the genus. Often locally
abundant, specimens are found in coastal
sand dunes and a variety of interior habitats,
including sandy washes and alluvial fans
(Stebbins and McGinnis, 2012). Much of the
coastal dune habitat has been destroyed by
coastal development between Ventura Coun-
ty and the Mexican Border. Fortunately, a
large protected population persists in the
remnant of the once extensive El Segundo
Dunes at Los Angeles International Airport
(Fig. 4).
Anniella stebbinsi is common at the west-
ern margin of the Colorado Desert under
trash dumped at the base of Mt. San Jacinto
in the vicinity of Cabazon, Riverside County.
Here the only large shrub is Creosote (Larrea
tridentata). The seasonal Whitewater River
provides sufficient moisture near the surface.
The disjunct northern populations occur in
sandy soils in the Piute and Tehachapi
mountains at elevations of 400–900 m in
both Oak Woodland and Mixed Conifer
Forest. In the lower drainage of Caliente
Creek at Caliente Post Office, individuals
have been collected beneath cardboard cover
placed under Scalebroom bushes (Lepidos-
partum squamatum). There is continuous
sandy habitat along Caliente Creek between
Caliente Post Office and Sand Ridge Pre-
serve, the type locality for A. grinnelli.
Additional fieldwork is needed to document
the location of an almost certain contact
between these two species. Contact between
A. stebbinsi and A. pulchra is likely along the
coast of California between the cities of
Santa Barbara and Oxnard and along the
southeastern slope of the Tehachapi Moun-
tains, where A. pulchra is common in Joshua/
Juniper woodland.
Etymology. This species is named after
Robert Cyril Stebbins (1915–; Fig. 6) who
was appointed the first Curator of Herpetol-
2013 FOUR NEW SPECIES OF ANNIELLA 13
ogy at the Museum of Vertebrate Zoology in
1945. Robert Stebbins’ contribution to west-
ern North American herpetology includes
many scientific publications, but especially
his classic, comprehensive, beautifully self-
illustrated, and influential field guides (Steb-
bins 1951, 1954, 1960, 1966, 1972, 1985,
2003; Stebbins and McGinnis, 2012).
CONSERVATION IMPLICATIONS
The former A. pulchra,aspeciesofspecial
concern (Jennings and Hayes, 1994), is now
divided into five species. This means A.
pulchra has a smaller distribution than
previously recognized, thereby enhancing
concern about its conservation status. The
remaining four species have even smaller
ranges, some of which are degraded or
threatened by human activities. Whereas
much of the range of A. stebbinsi is already
compromised by urban development, the
conservation implications for the other three
new species are even more striking because
of their very limited distributions. Anniella
grinnelli is known from a few sites in the
southern San Joaquin Valley, an area that has
been greatly modified by urban and agricul-
tural development (PPIC, 2006; Great Valley
Center, 2007). Anniella grinnelli persists in
small patches within the Bakersfield city
limits, but some of the populations we
collected were extirpated by development
during the course of this study. The type
locality at the Sand Ridge Preserve is a secure
site that will help ensure the species survival.
Anniella alexanderae is known from two sites
at the base of the Temblor Mountains, and
should be considered rare pending further
study. Finally, Anniella campi is known from
just three sites. This species may be restricted
to the vicinity of potentially fragile springs in
canyons that open into the Mojave Desert
and so warrants careful monitoring. Addi-
tional research into the distribution, contact
zones, and diversity of Anniella is clearly
needed.
ACKNOWLEDGMENTS
Scientific collection permits were issued
by the California Department of Fish and
Wildlife. Assistance with fieldwork was
provided by David Germano (California
State University, Bakersfield), Chris R. Feld-
man (University of Nevada, Reno), Sarah
Rieboldt (LSA Associates, Inc.), Jonathan
Richmond (USGS Western Ecological Re-
search Center, San Diego), Greg Pauly
(Natural History Museum of Los Angeles),
Paul Collins (Santa Barbara Natural History
Museum), Barbara Bradford (Riverbank,
California), and Robert W. Hansen (Clovis,
California). Carol S. Spencer (MVZ), Jens
Vindum (CAS), Jose´ Rosado (MCZ), Brad-
ford Hollingsworth (SDNHM), and Laura
Williams (SDNHM) are thanked for han-
dling the specimen accessions and/or fielding
numerous data inquiries.
Fieldwork was facilitated by Kathy
Sharum (Carrizo Plain National Monument)
and Greg Warrick (Sand Ridge Preserve).
Fieldwork at El Segundo Dunes at Los
Angeles International Airport was facilitated
by Nebu John (Environmental Services
Division, Los Angeles World Airports),
along with Jose U. Alvarenga, Karin Chris-
tie, and Peggy Nguyen.
Plant identifications were provided by
Denis Kearns (Bakersfield Field Office,
Bureau of Land Management), Kathy
Sharum (Carrizo Plain National Monu-
ment), and L. Maynard Moe (California
State University, Bakersfield). Aaron Bauer
(Villanova), Miguel Vences (Technical Uni-
versity of Braunschweig), and David B.
Wake (MVZ) are thanked for discussions
about describing cryptic species.
Sarah Rieboldt (LSA Associates, Inc.)
provided assistance with the figures. Jere H.
14 BREVIORA No. 536
Lipps (John D. Cooper Archaeology and
Paleontology Center), David K. Smith (Uni-
versity of California, Berkeley), Barbara R.
Stein (Fred Hutchinson Cancer Research
Center), Karen Klitz (MVZ), and Michelle
Koo (MVZ) provided assistance with photos
of Annie Alexander, Charles Camp, Joseph
Grinnell, and Robert Stebbins. Robert B.
Jensen and Richard A. Arnold (Entomolog-
ical Consulting Services Ltd.) provided the
photo of El Segundo Dunes. Jon Fong
(CAS) and Jonathan Woodward (MCZ)
provided some of the x-rays.
Lawrence E. Hunt (Santa Barbara Natural
History Museum), Samuel S. Sweet (Univer-
sity of California, Santa Barbara) provided
insightful discussion on this research.
APPENDIX 1
The following specimens were used for the
vertebral counts (shown in parentheses)
summarized in Table 1 and 2: A. alexan-
derae:CAS 238588 (82); MCZ R-189383
(84); MCZ R-189384 (83); MCZ R-189385
(82); MCZ R-189386 (82); MVZ 250528 (81);
MVZ 250549 (83); MVZ 250550 (80); MVZ
250570 (81); MVZ 250573 (81); MVZ
250574(82); MVZ 250575 (83); MVZ
250576 (82); MVZ 257082 (84); MVZ
257717(84); MVZ 257718 (81); MVZ
257720 (82); MVZ 257741 (82). A. campi:
MCZ R-189380 (78); MCZ R-189381 (76);
MCZ R-189382 (78); MVZ 104771 (76);
MVZ 228817 (77); MVZ 228818 (77); MVZ
228819 (77); MVZ 228829 (78); MVZ 232844
(76); MVZ 257727 (74); MVZ 257728 (75);
MVZ 257729 (78); MVZ 257730 (75). A.
grinnelli:MCZ R-189378 (79); MCZ R-
189379 (83); MVZ 230663 (83); MVZ
230665 (74); MVZ 247487 (82); MVZ
250527 (84); MVZ 250534 (81); MVZ
250541 (81); MVZ 250542 (79); MVZ
250543 (86); MVZ 250545 (84); MVZ
250546 (83); MVZ 250547 (83); MVZ
250548 (82); MVZ 257714 (79); MVZ
257716 (80); MVZ 257724 (80); MVZ
257725 (80); MVZ 257726 (80); MVZ
257737 (82); MVZ 257738 (81). A. pulchra:
MVZ 27300 (73); MVZ 33793 (78); MVZ
33795 (76); MVZ 33796 (75); MVZ 33860
(73); MVZ 45612 (74); MVZ 58106 (75);
MVZ 58410 (78); MVZ 60216 (74); MVZ
60292 (75); MVZ 64105 (76); MVZ 64106
(76); MVZ 71919 (75); MVZ 71920 (75);
MVZ 71921 (74); MVZ 71922 (75); MVZ
84593 (73); MVZ 83594 (75); MVZ 117600
(75); MVZ 223384 (74); MVZ 227778 (73);
MVZ 228815 (74); MVZ 228816 (75); MVZ
228832 (78); MVZ 247488 (72); MVZ 247489
(74); MVZ 250536 (78); MVZ 250537 (80);
MVZ 250538 (79); MVZ 250539 (79); MVZ
250540 (78); MVZ 250562 (79); MVZ 250563
(78); MVZ 250564 (78); MVZ 250566 (79);
MVZ 250567 (79); MVZ 250569 (79). A.
stebbinsi:MVZ 226854 (75); MVZ 226855
(75); MVZ 226856 (74); MVZ 226857 (77);
MVZ 226859 (75); MVZ 226860 (76); MVZ
226863 (75); MVZ 228844 (75); MVZ 228861
(74); MVZ 230554 (77); MVZ 230556 (75);
MVZ 230666 (76); MVZ 230667 (75); MVZ
230668 (77); MVZ 230669 (78); MVZ 230673
(81); MVZ 230674 (74); MVZ 230675 (73);
MVZ 230676 (74); MVZ 230677 (74); MVZ
230678 (74); MVZ 232618 (77); MVZ 232619
(77); MVZ 232621 (76); MVZ 250552 (75);
MVZ 250553 (73); MVZ 250577 (75); MVZ
250731 (74); MVZ 250732 (77); MVZ 250733
(76); MVZ 257743 (74); MVZ 257744 (73);
MVZ 257745 (76); MVZ 274645 (78); MVZ
267246 (71); MVZ 267247 (71). The follow-
ing specimens were used for the dorsal scale
counts (shown in parentheses) summarized
in Table 1 and 2: A. alexanderae:MVZ
250528 (257); MVZ 250550 (255); MVZ
250570 (257); MVZ 250574 (263); MVZ
250576 (252); MVZ 257718 (268); MVZ
257720 (265); MVZ 257739 (256); MVZ
257741 (261); MVZ 267236 (278). A. campi:
MVZ 104771 (223); MVZ 172784 (227);
2013 FOUR NEW SPECIES OF ANNIELLA 15
MVZ 228817 (230); MVZ 228818 (224);
MVZ 228819 (222); MVZ 228820 (235);
MVZ 228821 (223); MVZ 257727 (244);
MVZ 257728 (219); MVZ 267231 (229). A.
grinnelli:MVZ 230663 (234); MVZ 250527
(246); MVZ 250527 (246); MVZ 250534
(248); MVZ 250543 (234); MVZ 250546
(243); MVZ 250547 (249); MVZ 257714
(239); MVZ 257726 (247); MVZ 257742
(238). The following specimens were used to
evaluate ventral coloration (see Materials and
Methods): A. alexanderae:CAS 238588; MVZ
250570; MVZ 250549; MVZ 257739. A. campi:
MVZ 257727; MVZ 257728. A. grinnelli:CAS
234252; MVZ 250546; MVZ 257718; MVZ
257737; MVZ 257738. A. pulchra:MVZ
257098; MVZ 257731; MVZ 257732. A.
stebbinsi:MVZ 250552; MVZ 250553; MVZ
250556; MVZ 257723; MVZ 257735.
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