BREVIORA

Museum of Comparative Zoology

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on aan

US ISSN 0006-9698

CAMBRIDGE, MaAss.

4 SEPTEMBER 2014

NUMBER 538

THE TRAVELS OF THOMAS BARBOUR ON THE SHIP UTOWANA IN 1931 AND
THE TAXONOMIC STATUS OF ANOLIS UTOWANAE

STEVEN POE

ABSTRACT.

Anolis utowanae was described by Thomas Barbour in 1932 with an accompanying backstory

reviewing the collection of the holotype and only specimen in detail. Subsequent workers have been unable to procure
additional representatives of this species near the purported type locality of Mazatlan, Mexico, and it has remained a
taxonomic enigma since its description. Here I show that A. wtowanae is a junior synonym of A. conspersus, a species
endemic to Grand Cayman in the West Indies. The type specimen appears to have been part of a series collected by
Barbour on Grand Cayman during his 1931 trip from Miami to the Panama Canal and then North to Pacific Mexico,
including Mazatlan, on the yacht named Utowana. How the specimen came to be associated with Mazatlan remains a

mystery.

KEY WORDS:

On a day last spring, April 10, 1931, while
driving with Mrs. Barbour and my daughter,
Mary, to a finca some miles north of
Mazatlan, we stopped in a dusty lane to let a
herd of calves pass by. The herd was followed
by a barefooted Indian who trudged wearily
behind them through the deep dust. He
carried in his hand a long lashed whip and
from time to time he snapped it viciously and
in so doing killed the lizards on rocks or fence
posts by his way with most extraordinary skill.

Museum of Southwestern Biology and Department of
Biology, University of New Mexico, Albuquerque, New
Mexico 87131, U.S.A.; e-mail: anolis@unm.edu.

Anolis utowanae; Mexico; Grand Cayman; taxonomy; nomenclature; Thomas Barbour

We watched him some time quite fascinated.
I asked him what on earth he was pocketing
these lizards for. He looked at me with
surprise and then added, “I am taking them
’ 1 bought what he had

for a few cents. It was obvious that he felt

home to feed my cats.’

quite certain that he had been dealing with a
person of unsound mind as he walked on
looking at the coins, for it surely had never
occurred to him that such small game had a
cash value. Among these lizards one, I feel
quite certain, is unknown.

Barbour (1932), On anew Anolis from western
Mexico (description of Anolis utowanae).

© The President and Fellows of Harvard College 2014.

2 BREVIORA

I am convinced that the important part of
Thomas Barbour’s story—that Barbour ob-
tained a new species of lizard on an April day
in 1931 near Mazatlan—is false. I experi-
enced a creeping realization regarding the
status of Anolis utowanae on the basis of
personal experience collecting anoles in
Mexico, reading Henderson and Powell
(2004), and having discussions with col-
leagues more skeptical than I of the reliabil-
ity of a Barbour account. Comparison of
specimens and examination of historical
accounts validated this concern. Below I
present evidence that the name A. utowanae
is a Junior synonym.

HISTORY

As has been well chronicled by Henderson
and Powell (2004) and Barbour himself
(1943, 1945), Thomas Barbour and _ his
family traveled extensively in the West Indies
and Central America on the yacht named
Utowana in the 1920s and 1930s. Henderson
and Powell (2004) provided maps and/or
itineraries for three of the four Utowana trips
taken by Barbour during 1929-34 but
offered only a brief summary of the 1931
trip, where Barbour presumably would have
obtained the A. utowanae holotype specimen.
Fortunately, Barbour (1945) and Barbour’s
daughter Mary (Barbour, 1932) recount this
voyage in some detail.

Barbour left Miami on 30 January 1931
with daughter Mary, wife Rosamond, and
yachtmaster Allison Armour. The group
traveled to the Bahamas and Cuba before
heading west to the mainland coast near
Tela, Honduras, and thence south to the
Panama Canal for traversal to the Pacific
Ocean and the bulk of the voyage. Barbour
(1945) does not mention Grand Cayman, but
a visit to this island on 8 February was
described by Mary Barbour (1932: 23-24).
She recounted the collection of four lizards

No. 538

and a snake by her father. Harvard’s
Museum of Comparative Zoology (MCZ)
possesses specimens of Alsophis caymanus
from Grand Cayman with date of collection
10 February 1931 (MCZ 31499) from Bar-
bour, and of Anolis conspersus from Grand
Cayman with date of collection 8 February
1931 (MCZ 31500-31504). These five speci-
mens represent all but one of Barbour’s listed
MCZ Anolis collections for 1931. The other
is the holotype specimen of A. utowanae
(MCZ 31035; Mary Barbour’s diary notes on
T. Barbour’s collections mostly reflect what
is present in the MCZ collections, but she is
frequently off by one or two specimens; see
additional comparisons below).

Their Pacific travels continued north to
the Honduran Golfo de Fonseca and Gua-
temala before skirting Mexico up to the Gulf
of California. This Pacific Mexico part of the
trip was lovingly recounted by Barbour
(1945:32) for its beauty, life experience, and
outstanding bugling:

I can close my eyes and see myself now, sitting
on a rock beside Rosamond and Mary B. It
was at Agua Grande on Carmen Island, where
we rested a little while in the shade after a
search for wild cotton plants. A tiny snake
crawled out from under the rock we were
perched upon and right at my feet. I could
hardly believe my eyes, for it was something
which I had long hoped to see in life. We
sailed on northward as far as Santa Rosalia,
and Ros and I still often recall the music of a
bugle played by a Mexican soldier in the old
fort overlooking the town. He was by all odds
the best bugler we have ever heard anywhere.
This visit to Lower California was the
highlight of all our experiences on board the
Utowana.

This snake observation is the only herpe-
tological mention in Thomas Barbour’s
(1945) chronicle of the 1931 trip. In addition

2014

to the A. utowanae holotype, two herpeto-
logical specimens were deposited in MCZ by
Barbour with date of collection 10 April
1931 and locality Mazatlan (MCZ 31589:
Ctenosaura pectinata; MCZ 31588: Cnemido-
phorus hyperythrus).

Mary Barbour (1932:100) gave an account
of the acquisition of lizards on 11 April
outside of Mazatlan that is strikingly similar
to Thomas Barbour’s (1932) description of
the obtainment of A. utowanae purported to
have occurred on 10 April:

Saturday, April 11, 1931

MAZATLAN, MEXICO. AT ANCHOR.
Daddy woke us up early this morning, and he
and Mother and I went ashore at 8:30. First
we went to the market, and then took our car
and motored way out into the country. We
stopped at different fincas and bought some
corn, eggplants, cabbages, tomatoes, and limes
to take on board. While we were waiting for
the limes to be picked, a man came down the
lane driving a herd of cows. Suddenly we
heard his whiplash crack and saw him pick up
something on a rock near us. On asking him
what it was he replied “Lizards for my cat!” He
was flabbergasted when Daddy gave him 10
cents for them, and he told his friend who was
standing near that Senor Americano must be
crazy.

The Mary and Thomas Barbour accounts
seem unlikely to be independent, as Thomas
Barbour would have been familiar with
Mary’s account as he oversaw the publica-
tion of her diary. It is possible that Thomas
Barbour (1932) took his treatment of the
acquisition of A. utowanae directly from his
daughter’s pages, although I know of no
direct evidence that this happened. Mary
Barbour (1932) occasionally commented on
collected specimens (e.g., p. 83: ‘Daddy
found some rare lizards.’ my italics), but
she did not mention that any unusual species

STATUS OF ANOLIS UTOWANAE 3

was obtained from the cattle farmer in
Mazatlan.

COMPARISONS

The type specimen of A. utowanae is
nondescript, even for an anole. It is a
subadult female, long preserved, with no
strong markings save for elongate striations
on the underside of the head (Fig. 1). I
showed the specimen to an Anolis-knowl-
edgeable undergraduate in my lab, who
remarked that it “looks like every other
anole.” I concur broadly with this sentiment,
but would narrow it to say that the specimen
looks like every other female trunk crown
Anolis. Numerous Northern Lesser Antillean
and Greater Antillean species come to mind.

Herpetologists often rely heavily on local-
ity and live male dewlap coloration for
identification of anoles. When neither is
available, identification can be difficult. I
first used a new electronic key for anoles
(Poe, in preparation) to narrow the choices.
This key operates on the Lucid platform and
allows comparison of attributes (color,
scales, size, geography) scored for an un-
known specimen to a database of attributes
for all species of Anolis. Candidate species
may be ranked on the basis of their degree of
difference from the unknown specimen. This
procedure reduced the possible species as-
signments for A. utowanae to a few Greater
and Lesser Antillean forms. Mainland spe-
cies generally were poor matches, and as
realized by previous workers (e.g., Hardy
and Mc Dhiarmid, 1969; Lieb, 2001), no
Mexican species was similar. After narrow-
ing the choices, I spent a day comparing the
A. utowanae specimen with descriptions in
pertinent literature (e.g., Underwood and
Williams, 1959; Lazell, 1972; Schwartz and
Henderson, 1991) and MCZ material with a
focus on areas known to have been visited or
near to areas visited by Barbour. Once

BREVIORA No. 538

Figure 1.
and holotype specimen of A. utowanae (right).

convinced of the true species identity of the
A. utowanae type specimen, I scored five
individuals of this species of comparable size
and sex for demonstrative comparison.
Characters scored were snout-to-vent
length (SVL), height of the ear, length of
head from anterior of the ear opening to tip
of snout, width of head between postero-
ventral corners of the jugals, femoral length
from the longitudinal midline of the body
laterally to the knee, length of fourth toe,
length of tail, condition of nasal scale,

Undersides of heads showing characteristic striated markings in Anolis conspersus (left; MCZ 45042)

number of superciliaries, minimum number
of scales across the snout at the level of the
second canthals, number of expanded lamel-
lae under the fourth toe (counted in the
manner of Williams et a/. [1995]), minimum
number of scales separating the supraorbital
semicircles, minimum number of scales
separating the interparietal and the supraor-
bital semicircles, number of scales in contact
with the mental between the infralabials,
number of scales in contact with the rostral
between the supralabials, number of longi-

2014 STATUS OF ANOLIS UTOWANAE 5
TABLE 1. SCALE COUNTS AND MEASUREMENTS FOR THE TYPE SPECIMEN OF ANOLIS UTOWANAE AND FIVE COMPARABLY SIZED
FEMALE SPECIMENS OF A. CONSPERSUS.
Trait Anolis conspersus Mean (Range) Anolis utowanae MCZ 31035

SVL 41.7 (36.5-46.8) 39.6
Scales across the snout at second 6.8 (6-7) 6

canthals
Lamellae on fourth toe 23.1 (22-25) 23/24
Scale rows separating supraorbital 0.8 (0-1) ]

semicircles
Scales from interparietal to 2.4 (2-3) 2

supraorbital semicircles
Postmentals 5.8 (5-6) 5
Postrostrals 8 (6-10) 9
Ear height/SVL 0.03 (0.02—0.03) 0.04
Head length/SVL 0.27 (0.26—0.28) 0.27
Head width/SVL 0.17 (0.16—0.17) 0.17
Femoral length/SVL 0.27 (0.26—0.28) 0.26
Toe length/SVL 0.19 (0.18—0.20) 0.18
Dorsal scales in 5% SVL 9.6 (8-11) 8
Ventral scales in 5% SVL 6.6 (5-8) 6
Supralabials to eye 6 (5-7) 7

Nasal

supralabial
Interparietal length/SVL
Tail length/SVL
Superciliaries

elongate, contacts rostral anterior to
sulcus between rostral and first

0.03 (0.03—0.04)
1.76 (1.51-1.94)
1 (1)

elongate, contacts rostral anterior to
sulcus between rostral and first
supralabial
0.03
1.36
]

Specimens examined Anolis conspersus: Museum of Comparative Zoology 31035, 45042, 87372, 87388, 87396,

174146.

tudinal dorsal scales in 5% of SVL, number
of longitudinal ventral scales in 5% of SVL.
Scoring for these characters follows Williams
et al. (1995) and Poe (2004).

RESULTS

Character states of the holotype of 4A.
utowanae fall within the range of scores for
A. conspersus (Table 1). This species further
possesses the distinctive chin striations of A.
utowanae (Fig. 1), which nevertheless are
not unique to A. conspersus, even among
trunk-crown anoles (pers. obs.). Perhaps
equally compelling, this result makes sense
in light of Barbour’s activities and MCZ
accessions of the time. Barbour’s only West
Indian collections of 1931 are from Grand

Cayman, and his only anole specimens are
A. conspersus.

I cannot determine whether Barbour’s
specimens, including A. utowanae, are A.
conspersus conspersus (from western Grand
Cayman) or A. conspersus lewisi (from
eastern Grand Cayman); the locality for the
specimens is simply “Grand Cayman,” and
the two subspecies are separated on the basis
of color. Anolis conspersus conspersus may be
more likely, as Mary Barbour’s treatment
(1932:24) suggests that the specimens were
collected on an excursion to the west. If
future workers suggest species status for the
eastern Grand Cayman conspersus and if
Barbour’s specimens are determined to be 4.
c. lewisi, it is noteworthy that the name 4.
utowanae (Barbour, 1932) predates Grant’s

6 BREVIORA

A. c. lewisi (Grant, 1941). That is, the proper
name for eastern Grand Cayman Anolis
would then be A. utowanae rather than
A. lewisi. Those who recognize subspecies
would call the eastern form A. conspersus
utowanae.

DISCUSSION

Several authors have included A. utowanae
in species lists without comment (e.g.,
Enderson et al., 2009), and a few have
discussed the taxonomic affinities of A.
utowanae. Barbour (1932) in the original
description noted similarity to A. baccatus,
which was deemed a synonym of A. caroli-
nensis by Kohler (2011). Thus Barbour
(1932) unwittingly associated A. utowanae
with a species outside of Mexico that shares
some potentially convergent features with A.
conspersus (they are both “trunk-crown”
species). Smith (1939) suggested close rela-
tionship with A. schmidti (= nebulosus;
Nieto-Montes de Oca et al., 2013), and the
final couplet regarding A. utowanae in the
lizard key of Smith and Taylor (1950)
separates A. utowanae from A. cymbops (a
schiedii group Anolis). Stuart (1955) stated
that A. utowanae should be viewed “with
suspicion” because it was known from a
single specimen from a well-studied region.
This statement was perhaps the first inkling
of the questionable status of A. utowanae.
Etheridge (1959) X-rayed A. utowanae and
placed it in his chrysolepis series on the basis
of its possession of caudal transverse pro-
cesses, a V-shaped parietal crest, 24 presacral
vertebrae, and an inscriptional rib formula of
2:2. This series inference is not unreasonable
considering the geography, as the traits of A.
utowanae are also found in some Caribbean
Beta anoles (Etheridge, 1959). Also, there is
more variation within series than Etheridge
was aware of in 1959, and known 4A.
conspersus were not included in his study.

No. 538

Hardy and McDiarmid (1969) apparently
examined the holotype of A. utowanae and
noted its distinctness relative to other
Sinaloan anoles. Lieb (2001:53) stated that
A. utowanae was a junior synonym but did
not specify of what.

The taxonomic assignment of the A.
utowanae type specimen adds to the growing
refinement of Mexican anole taxonomy
(Lieb, 2001; Kohler, 2012; Nieto-Montes de
Oca et al., 2013). Although much work
remains to be done, progress is underway
and several additional issues are likely to be
cleared up in the near future.

Two questions remain unanswered regard-
ing A. utowanae. First, how did the proper
identification of A. utowanae elude the
careful workers on anoles post-Barbour? In
particular, how did ultimate anole expert
Ernest Williams not recognize A. utowanae
as A. conspersus when he had coauthored
with Underwood (Underwood and Williams
1959) the seminal contribution on Jamaican
anoles (the lineage from which A. conspersus
evolved), was undoubtedly familiar with
Barbour’s travels given their shared history
as curators of herpetology at the MCZ, and
had the pertinent type specimen and com-
parative material readily available to him in
the MCZ herpetology collection for decades?
The answer may be that Mexico was the one
area of Anolis in which Williams lacked
confidence; he relied heavily on Carl Lieb for
Mexican anole information, frequently send-
ing him material for identification (pers.
comm. from Williams 1992). Lieb conversely
worked almost exclusively in Mexico so there
was no reason for him to be familiar with the
distantly related West Indian forms.

Second, what really happened with Bar-
bour’s “Indian” story? The detail in the tale
encourages acceptance of its veracity. This
degree of detail likely contributed to willing-
ness to entertain the possibility of the validity
of this species in the face of multiple failed

2014

collecting attempts at the type locality (by,
e.g., my group in 2011) and gradual appre-
ciation of the aberrance of the specimen in
Sinaloa (Hardy and McDiarmid, 1969). The
A. utowanae specimen apparently is part
of the series of A. conspersus collected by
Barbour (MCZ 31500-31504), so how did
one of these specimens get attributed to a
lizard-killing cat owner in Mazatlan?

If Barbour were the only storyteller, it
might be tempting to consider the acquisition
story a complete fabrication. Thomas Bar-
bour’s contributions to herpetology and
MCZ collections are great, but his taxonom-
ic work has escaped celebration for meticu-
lous accuracy. Some of the colleagues with
whom I discussed the case of utowanae
seemed to attribute the confusion simply to
Barbour being Barbour. Note, for example,
that Barbour and his wife disagreed on the
identity of the person who lanced some
parasites out of another passenger during
the 1931 trip, with Barbour claiming it was
he and Mrs. Barbour vehemently arguing
otherwise (Barbour 1945:124). A disagree-
ment between married persons on the minu-
tiae of a temporally distant event is not
unusual, so I’m told. But here we have an
argument about what would seem to be a
memorable aspect of a major happening. The
analogy is obvious and unavoidable: if
Barbour was capable of confusing himself
with someone who performed a minor
surgery during an emergent situation at sea,
it does not seem so unrealistic that he could
misremember when or how he obtained a
lizard.

But there is no reason to doubt Mary
Barbour’s (1932) account. She seems to have
been recording events more or less as they
happened, and her dates of trip milestones
approximately correspond to those given by
Barbour (1945; although again, these ac-
counts cannot be considered independent).
The lizard collections mentioned by Mary

STATUS OF ANOLIS UTOWANAE g

Barbour in her diary correspond to speci-
mens, localities, and collection dates listed in
the MCZ catalog. Mary Barbour noted
collection of lizards on 11 February (Grand
Cayman; “four lizards’, 1.e., the A. consper-
sus), 31 March (La Paz, Lower California:
“some rare lizards’’), 1 April (Espiritu Santo
Island; “a lot of lizards’), 2 April (San
Francisco Island; “‘some fine lizards’’), 3 April
(Agua Verde; “2 kinds of lizards’’), 4 April
(Agua Grande; “‘caught lizards by stunning
them with stones’), 6 April (San Marcos
Island; “*5 lizards”), 11 April (Mazatlan;
“Lizards for my cat’, 1.e., the purported
utowanae acquisition). The MCZ catalog lists
dates of collection for Barbour’s 1931 lizard
material as 8 February (Grand Cayman; five
A. conspersus), 1 April (Isla Gallina; 11
Phyllodactylus, four Urosaurus), 2 April (sla
San Francisco; one Phyllodactylus, four Saur-
omalus, three Uta), 3 April (Agua Grande;
two Cnemidophorus, two Uta. Puerto Escon-
dido; one Uta. Agua Verde; two Urosaurus), 6
April (Isla San Marcos; two Cnemidophorus,
four Urosaurus), 10 April (Mazatlan; one
Cnemidophorus, one Ctenosaura, A. utowa-
nae), 23 April (Chiriqui, Panama; one Crteno-
saura). Minor differences in details aside (e.g.,
number of specimens collected, specific dates
of collection for contiguous mainland locali-
ties), Mary Barbour’s diary appears to be an
accurate chronicle of the collections of
Thomas Barbour during the 1931 trip on the
ship Utowana.

Given the apparent reliability of Mary
Barbour’s account, it seems likely that the
meeting with the cattle farmer did actually
occur around 11 April 1931 near Mazatlan
and that lizards were procured at that time.
Notably, Mary Barbour (1932:100) men-
tioned observing just one instance of lizard
collection by the cattle farmer: “Suddenly we
heard his whiplash crack and saw him pick
up something on a rock near us.” Barbour
(1932:11), possibly embellishing, implied

io 4)

multiple whippings that resulted in dead
lizards: “‘from time to time he snapped it
viciously and in so doing killed the lizards
on rocks or fence posts.” The specimens
obtained from the cattle farmer may have
been a whiptail (MCZ 31588) and a cteno-
saur (MCZ 31589). No other Barbour
collections around 10-11 April are listed in
the MCZ catalog or by Mary Barbour
(1932), and these are the only specimens of
the trip purportedly from near Mazatlan
(besides the A. utowanae holotype). One of
these specimens, the ctenosaur, seems too
large to have been killed in the manner
described (the ctenosaur specimen is man-
gled; the whiptail specimen is _ pristine;
personal communication from Jonathan
Losos). Perhaps the whiptail specimen is
the lizard killed in Mary Barbour’s (1932)
account.

If the Mazatlan lizard vignette occurred as
chronicled in Mary Barbour’s diary, and A.
utowanae (= conspersus) was collected on the
same trip but much earlier on Grand
Cayman, it seems likely that Thomas Bar-
bour simply misplaced his specimens such
that a Grand Cayman lizard ended up
among lizards from Western Mexico. Mary
Barbour noted four lizards collected on
Grand Cayman, the MCZ catalog (..e.,
Thomas Barbour) records five, and actually
there were six. Mary Barbour’s underesti-
mate of the number of Grand Cayman
lizards collected may have influenced Bar-
bour’s interpretation of the six anoles caught
on this trip, but we can only speculate how
this influence could have prompted the
erroneous assignment of the utowanae spec-
imen to Mazatlan.

I would prefer an ending with more closure,
but this conclusion of an unexplained speci-
men mixup is as far as current evidence can
take us. How Barbour could cull one of a
series of collected A. conspersus—a series for
which he apparently accurately chronicled

BREVIORA

No. 538

identity, locality, and (approximately) date—
and imagine a role for this specimen in a
colorful, true collection story that took place
on the other side of the continent 2 months
later remains a mystery.

ACKNOWLEDGMENTS

Thanks to Jonathan Losos, Mason Ryan,
Levi Gray, Heath Weaver, and anonymous
reviewers for comments on the manuscript.
Joe Martinez, Jonathan Woodward, and
Tsuyoshi Takahashi were helpful and hospi-
table during work done at the Museum of
Comparative Zoology. Jose Rosado and
Jonathan Losos kindly checked MCZ spec-
imens for this paper. This research was
funded by the National Science Foundation
DEB-0844624.

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