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TEXAS JOURNAL
SCIENCE

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THE TEXAS JOURNAL OF SCIENCE

Volume 6 1 , No. 1 FEBRUARY, 2009

CONTENTS

The Effects of All-Terrain Vehicle Use on the Herpetofauna
of an East Texas Floodplain.

By T imothy R. Hunkapiller, Neil B. Ford and Kevin Herriman ............................ 3

Selection of Desert Bighorn Sheep {Ovis canadensis) Transplant Sites
in Sierra Maderas del Carmen and Sierra San Marcos y del Pino,

Coahuila, Mexico.

By Alejandro Espinosa-T, Armando J. Contreras-Balderas,

Andrew V. Sandoval and Mario A. Garcia~A ..................................................... 15

Geographic Distribution Records for Select Fishes of
Central and Southern Arkansas.

By Chris T, McAllister, Renn Tumlison and Henry W. Robison.......................... 31

Comparison of Total Lipid and Fatty Acid Compositions of Whole-Body
and Body Segments of Lertha extensa Adults (Neuroptera: Nemopteridae).

By Ozlem Cakmak, Mehmet Bashan and Ali Satar . . . . . ........45

General Notes

Myxidium serotinum (Protista: Myxozoa) from a Jefferson Salamander
{Ambystoma jeffersonianum), in Illinois.

By Chris T. McAllister, John A. Crawford and Andrew R. Kuhns ................ 61

Records of the Porcupine (Erethizon dorsatum) from the
Eastern Margins of the Edwards Plateau of Texas.

By Amy B. Baird, Gregory B. Pauly, David W. Hal and

Travis J. Laduc .............................................................................................. 65

First Record of Cymbovula acicularis
(Gastropoda: Prosobranchia: Ovulidae)
from the Coast of Tamaulipas, Mexico.

By Alfonso Correa-Sandoval and Ned E. Strenth. .................................. ....... 67

Author Instructions ..............................................................................................73

Membership Application

THE TEXAS JOURNAL OF SCIENCE
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Manuscript Editor:

Frederick B. Stangl, Jr., Midwestern State University
Associate Editor for Botany:

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Associate Editor for Chemistry:

John R. Villarreal, The University of Texas-Pan American
Associate Editor for Computer Science:

Nelson Passos, Midwestern State University
Associate Editor for Environmental Science:

Thomas LaPoint, University of North Texas
Associate Editor for Geology:

Ernest L. Lundelius, University of Texas at Austin
Associate Editor for Mathematics and Statistics:

E. Donice McCune, Stephen F. Austin State University
Associate Editor for Physics:

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TJS Manuscript Editor
Department of Biology
Midwestern State University
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Scholarly papers reporting original research results in any field of
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The Texas Journal of Science. Instructions to authors are published one or
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AFFILIATED ORGANIZATIONS
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Texas Section, Mathematical Association of America
Texas Section, National Association of Geology Teachers
Texas Society of Mammalogists

TEXAS I OF SCI61(1):344

FEBRUARY, 2009

THE EFFECTS OF ALL^TER^IN VEHICLE USE ON THE
HERPETOFAUNA OF AN EAST TEXAS FLOODPLAIN

Timothy R. Hunkapiller, Neil B* Ford and Kevin Herriman

Department of Ecology and Evolutionary Biology
University of Tennessee, Knoxville, Tennessee 37996
Department of Biology, University of Texas at Tyler,

Tyler, Texas 75799 and
Texas Parks and Wildlife Department
Lindale, Texas 75771

Abstract “Recreational all-terrain vehicle (ATV) use is known to cause extensive
environmental damage in xeric habitats, however their impact has been poorly
studied in floodplain ecosystems. Floodplains are complex and important wetland
habitats due to their high primary productivity and biodiversity. In this study,
amphibians and reptiles are used as indicators of environmental stress from ATV use
in a floodplain at the Old Sabine Bottom Wildlife Management Area (OSBWMA),
Smith County, Texas. The herpetofauna were surveyed using cover board arrays
(wood and tin) placed at nine sites: three within a forest, three along trails open to
ATV traffic, and three along trails closed to ATV traffic. No significant differences
were observed in abundance, richness, evenness, or diversity of amphibians and
reptiles found among these three treatments (P>0.05). The apparent lack of effect
could be due to restricted use and the recent opening of the OSBWMA to ATVs,
and/or the natural resilience of floodplain ecosystems to disturbance. Additionally,
the impact of ATV use may have been masked by the relatively short time period of
the study (14 months) and low rainfall during the investigation.

Off-road vehicle use has increased greatly since the 1970s, from
approximately 5 to 35 million annual users (Wisdom et al. 2004;
USDA 2004). Minimal research has been conducted on the impact
of off-road vehicle trails on ecosystems (Phillips & Alldredge 2000;
Gaines et al. 2003). Additionally, only xeric regions have been
widely examined in studies of the effects of off-road vehicles on
ecosystems (Iverson et al 1981; Adams et ah 1982; Webb &
Wilshire 1983).

Recreational all-terrain vehicle (ATV) use can affect wildlife by
direct injury or death, by noise pollution, through fragmentation of
habitat, and by creating migration barriers (e.g., Webb & Wilshire
1983; Reijnen et al. 1996; Forman & Alexander 1998; Bonnet et ah
1999). Such effects are known to cause changes both in the

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

abundance and diversity of organisms in biological communities
(Pearson et aL 1999; Findlay & Bourdages 2000; Trombulak &
Frissell 2000). Habitat fragmentation and associated edge effects
are considered to have detrimental effects on community
composition by specifically impacting interior species (Wilcox &
Murphy 1985; Robinson & Quinn 1992; Spellerberg 1998). Even
less conspicuous effects of ATV use may harm community
structure. Palis & Fischer (1997) found that off-road vehicle use
disrupts pond floor microtopography by breaking the hardpan that
lies below the pond and causing a shorter hydroperiod, thus
decreasing the availability of ephemeral ponds for breeding
amphibians. Furthermore, a variety of chemicals derived from
gasoline additives can be introduced into a habitat leading to direct
toxic effects and indirect disruption of food webs (Trombulak &
Frissell 2000; Forman et al. 2003).

Floodplains are a complex and important wetland ecosystem
because their high productivity leads to increased biodiversity
(Reice 1994; Bayley 1995). The flood-pulse concept, characterized
by the predictable advance and retreat of water on a floodplain, is
postulated to maintain diversity in a system (Bayley 1995; Ostfeld
& Keesing 2000). The primary productivity is augmented by an
influx of nutrients that are washed into temporary pools and the
forest floor resulting in an increase in plant and animal abundance
due to extra energy availability (Bayley 1995; Sparks 1995).
Additionally, temporary pools that form during the recession phase
(Junk 1973; Sparks et al. 1998) hold fish and ftinction as breeding
sites for many amphibians. Both forest floor and temporary pools
provide readily available food sources for many carnivores (Modes
et al. 1998; Shiel et al. 1998). Despite their biological importance,
floodplain ecosystems are among the most threatened natural areas
in the eastern United States and, of all wetland types, floodplains
have suffered the greatest losses (Bayley 1991; Graham et al.
1997).

HUNKAPILLER, FORD & HERRIMAN

Amphibians and reptiles are often locally abundant, easy to
sample, and respond to an assortment of subtle environmental
changes, making them good indicators of wetland health (Heyer et
ah 1994; Welsh & Ollivier 1998; Christy & Dickman 2002).
Alterations of such environmental factors as temperature
(Seebacher et al. 2003), precipitation (Heyer et al. 1994), pH
(Wyman & Hawksly-Lescault 1987), salinity (Christy & Dickman,
2002), and ultraviolet radiation (Blaustein et al. 1998) have been
shown to cause changes in amphibian and reptile abundance and
diversity in an area. Because of their importance in ecosystem
structure and function, herpetofauna have also been used as
indicators of ecosystem integrity (Duellman & Trueb 1994;
Petranka 1998; Lips 1999). The amphibian and reptile community
in an east Texas floodplain that is partially open to ATV travel was
evaluated to assess the impacts of ATV use on floodplain
ecosystems.

Methods

Study site-T\iQ Old Sabine Bottom Wildlife Management Area
(OSBWMA) is located in northern Smith County, Texas and is
bordered along its northern edge by the Sabine River. With 2087
ha of hardwood forest, the OSBWMA is one of the largest
contiguous bottomland hardwood forests remaining in Texas. The
primary vegetation type is a diverse Water Oak-Elm- Sugarberry
Forest. Dominant vegetative overstory is comprised of water oak
(Quercus nigra), willow oak (Quercus phellos), overcup oak
(Quercus lyrata), pecan and hickory {Cary a sp.), cedar elm (JJlmus
crassifolia), water elm (Planera aquatica), ash {Fraxinus sp.),
American hornbeam {Carpinus caroliniana), and black willow
{Salix nigra). Periodic flooding occurs at the OSBWMA due to
heavy rainfall, opening of the floodgates at the Lake Fork Dam, and
overflow from the Iron Bridge Spillway on Lake Tawakoni,
however, no flooding occurred during this study. Flooding usually
occurs in the winter and spring months and results in the filling of
both natural ephemeral pools within the forest and the man-made

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

soil compacted ephemeral pools formed from tire ruts in the trails
(Hampton & Ford 2007).

Several trails through the OSBWMA were created by the use of
motorized vehicles prior to Texas Parks and Wildlife Department
(TPWD) ownership of the property. Current trail use includes
hiking, horseback riding, and beginning in 2004 the restricted use
of all-terrain vehicles for public hunting access on two designated
trails from October 1 to May 3 1 each year. No recreational riding is
allowed. The previous use of motorized vehicles and AT Vs caused
deep tire ruts and compaction of the soil on the trails. Over the
course of this study, the trails open to ATV use received an average
of one to two users per day during the October to December time
period, with less use through the end of May. The on-site staff of
the OSBWMA traversed all trails on the area conducting routine
trail maintenance and management activities by ATV or medium
sized tractor on a year round basis. The personnel at the OSBWMA
attempted to restrict their use of the trails to drier periods to reduce
impacts (Shaun Crook, pers. comm.).

Methods -^\nQ areas were chosen to monitor reptile and
amphibian populations: three within the forest (at least 50 m from
any edge), three along trails that are open for ATV traffic, and three
along trails that are closed to ATV traffic (very limited use since
1996). Each area was divided into five stations, each 50 m apart.
Each station consisted of two wooden cover boards (ca. 175 by 80
cm) and two pieces of corrugated tin (ca. 200 by 80 cm). One of
each cover type (wood and tin) was placed on each side of the trail
at each station. Because there were no trails within the undisturbed
forest plots, each station within the forest consisted of one piece of
both tin and wood placed approximately two meters from another
cover array to emulate the same sampling system as on the trails.
The cover items were placed in November 2004 and checked at
least once a week from November 2004 to December 2005. In this
wetland system cover boards are an effective survey method in a
very short time frame (Hampton & Ford 2007). Individual

HUNKAPILLER, FORD & HERRIMAN

amphibian and reptiles captured under boards were identified, sexed
and marked when possible, and subsequently released. Individual
identification marking consisted of scale clipping, toe clipping, or
PIT tagging depending on the size and species of the captured
individual.

Statistical methods and analyses.-^owQXdX community indices
were computed including Simpson’s index of diversity (Krebs
1999), Smith and Wilson’s index of evenness (Smith & Wilson
1996; Krebs 1999), and Morisita’s index of similarity (Morisita
1959; Krebs 1999). These indices of diversity and evenness as well
as abundance and richness of herpetofauna species were compared
among sites using an analysis of variance (Systat Software Inc.
2004) to determine if there was a significant difference among
treatments.

Results

A total of 403 amphibians and reptiles representing 16 species
were recorded under cover boards. Totals included two
salamander, one anuran, three lizard, and 10 snake species. Of the
total animals captured, 31 (7%) were salamanders, 23 (6%) were
anurans, 236 (59%) were lizards, and 113 (28%) were snakes
(Table 1). Ten species were captured in all of the treatments.
Forty-five marked individuals were recaptured during the sampling
period. Thirty-one recaptures were lizards.

The ATV trails had the highest species richness. The highest
abundance was observed in the forested control areas. However,
abundance, richness, evenness, and diversity of amphibians and
reptiles were not significantly different among ATV, non-ATV
active trails, or control treatments (f=0.04, 1.05, 0.26, 1.61, P>0.05
respectively; Table 2). Based on Morisita’s index of similarity,
community structures were extremely similar (Ca.=1.00 for all
comparisons).

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

Table 1. Numbers of amphibian and reptile species captured on ATV trails, Non-ATV
trails, and within forest control plots at the Old Sabine Bottom Wildlife Management
Area, Smith County, Texas.

Taxon

ATV

Non-ATV

Control

Total

Class Amphibia

Order Caudata

Ambystoma opacum

Ambystoma texanum

Order Anura

Rana utricuJaria

Class Reptilia

Order Squamata

Eumeces fasciatus

Eumeces laticeps

Scincella lateralis

116

Agkistrodon piscivorus

Agkistrodon contortrix

Pantherophis obsoleta

Farancia abacura

Lampropeltis getula

Nerodia erythrogaster

Nerodia fasciata

Nerodia rhombifer

Storeria dekayi

Thamnophis proximus

Total Individuals

138

124

141

403

Total Species

Discussion

Habitats in the three areas where cover boards were placed were
visibly different. Damage in the area where AT Vs were in current
use included destruction of vegetation within and next to trails,
packing of soil in some parts of the trail, and loosening of mud in
deeper, wet areas creating suspended silt. The ATV restricted trails
showed some packing and older ruts from previous vehicle use, but
much less vegetation damage. The forested area had natural
depressions and damage from tree falls but little or no human
impact. The methods used to place boards in each area resulted in
some stations being near pools or rutted areas and some stations

HUNKAPILLER, FORD & HERRIMAN

Table 2. Abundance, richness, diversity, and evenness for amphibians and reptiles
captured on ATV trails, non~ATV trails, and forest control plots at the Old Sabine
Bottom Wildlife Management Area, Smith County, Texas.

ATV

Non ATV

Control

Abundance

138

124

141

0.04

0.962

Richness

1.05

0.406

Diversity

0.846

0.839

0.816

0.26

0.780

Evenness

0.409

0.460

0.427

1.61

0.855

being near dry parts of the trail A total of 54 captures of only three
species of amphibians were recorded. This reflects a much lower
abundance than in previous work at the OSBWMA (Hampton 2004;
Hampton & Ford 2007). These studies utilized boards in close
proximity to ephemeral pools or deep ruts on trails. The lower
number of amphibians obtained during the current study may relate
to the random placement of cover objects in relationship to pools of
water. Additionally, an unusually low amount of precipitation
occurred in the second half of 2005. The average rainfall in nearby
Dallas, Texas during November and December is 6.12 cm. In 2004
the average precipitation for November and December was 8.50 cm
but for 2005 was only 0.60 cm (NOAA 2006) reflecting the very
dry second half of the study period. During this study 349 reptiles
belonging to 13 species were recorded. These numbers are more
typical for the site and reflect that lizard and snake numbers were
less influenced by low rainfall.

Species richness, abundance, diversity, and evenness of
herpetofauna did not vary among trails open to ATV use, trails
closed to ATVs, and the internal forest areas at the OSBWMA
(Table 1). This suggests that the changes to the trails by the ATV
use did not affect the biodiversity of amphibians and reptiles in this
floodplain ecosystem during the study period. This lack of effect
contrasts with previous studies of off-road disturbance (e.g., Iverson
et al. 1981; Adams et al 1982; Webb & Wilshire 1983; Luckenbach
& Bury 1983; Palis & Fischer 1997; Wisdom et al. 2004). Several

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

hypotheses are proposed concerning the lack of significant impact
of ATV use at OSBWMA.

First, the current study was limited to a 14-month duration. This
time period included the first year the OSBWMA was open to ATV
traffic since establishment of the wildlife management area in 1996.
Although physical and vegetation damage from AT Vs was evident,
the number of vehicles on each trail was very low (an average of 1-
2 users per day; Shaun Crook pers. comm.). Direct mortality from
AT Vs would have been unlikely and impacts like siltation in ruts
where amphibian eggs or larva occurred also may have been
limited. Previous researchers have also suggested that time lags of
greater than a year can exist between disturbance and ecological
responses (Magnuson 1990), and most studies in upland or desert
areas have been at least two years in length (Luckenbach & Bury
1983). Additionally, these aforementioned areas were usually open
to and extensively used by off-road vehicles for many years prior to
data collection, (Luckenbach & Bury 1983; Wisdom et al. 2004).
The abundances and diversity of amphibians and reptiles at the
OSBWMA may still be affected over time by ATV use.

A second possibility is that floodplain communities are resilient
to disturbances that include ATV activity. Previous studies of off¬
road vehicle disturbance have been conducted in upland forest
(Palis & Fischer 1997; Wisdom et al. 2004) or xeric desert habitats
(Iverson et al. 1981; Adams et al. 1982; Webb & Wilshire 1983;
Luckenbach & Bury 1983). One of the main effects of ATV
disturbance is habitat fragmentation, which causes migration
barriers and leads to decreased mobility and limited prey access
(Nour et al. 1998; Cushman 2006). This affect may be less
apparent in floodplain ecosystems because the major source of
nutrient influx is from flooding. The temporary pools formed by
the recession of floods become stocked with fish and create
amphibian breeding grounds. As the pools dry the fish and larva
are available food sources for many carnivores. Ruts created by
AT Vs simulate these pools and may also contain fish and

HUNKAPILLER, FORD & HERRIMAN

amphibian larva. The survival of organisms in those ruts, as
opposed to natural pools, is not known, but some species associated
with floodplains may be adapted to deal with perturbations.
However, observations made during this study revealed that
siltation in deep ruts appeared to kill some anuran eggs. A
secondary aspect of floods would be that they allow for animal
dispersal. Though the impact of the trails during flooding is not
known, the trails may acts as corridors as the water often is
channeled through them.

Frequent disturbances are known to cause complex effects on
ecosystems that can either amplify or mask anthropogenic affects
(Swetnam & Betancourt 1998; Hobbs & Morton 1999; Sherman
2001). It is possible that the lack of effect from ATV disturbance
was confounded by the low amount of rainfall activity during the
study, whereas this habitat is normally subjected to multiple flood
events each winter. It seems intuitive that AT Vs driving through
pools of amphibian egg masses or larva would have a negative
impact on those species. Indeed, it is almost certain that the
drought lowered amphibian activity during this study. Although
these conclusions are that the limited ATV use in this floodplain
appeared to have no significant effect on the herpetofauna, the
authors caution that fiarther study is needed. It is suggested that
these data represent a good baseline for additional studies,
particularly with the amphibians of the OSBWMA,

Acknowledgements

We thank Larry LeBeau and Shaun Crook of Texas Parks and
Wildlife for access to the study site and information regarding ATV
use at the OSBWMA. We thank Dr. Ron Gutberlet, Dr. Darrell
Pogue, and Andree Clark for comments on earlier versions of this
manuscript. We thank Stephan Lorenz, Jessica Coleman, Paul
Hampton, Casey Wieczorek, Robert Hunkapiller, Jr., and Rachel
Buerger for field assistance.

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

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TRH at: thunkapi@utk.edu

TEXAS I OF SCI. 61(1):15™30

FEBRUARY, 2009

SELECTION OF DESERT BIGHORN SHEEP {OVIS CANADENSIS)
TRANSPLANT SITES IN SIERRA MADERAS DEL CARMEN AND
SIERRA SAN MARCOS Y DEL PINO, COAHUILA, MEXICO

Alejandro Espmosa-T*^ Armando J. Contreras-Balderas**,

Andrew V* Sandoval, and Mario A. Garcia-A

*Desert Bighorn Sheep Restoration Program, CEMEX, Sustainability Vice-presidency,
Av. Independencia 901 Ote. Colonia Cementos, C.P. 64520, Monterrey, N.L Mexico and
**Laboratorio de Ornitologia, U.AN.L . Apartado Postal 425

San Nicolas de los Garza, N.L, Mexico 66450

Abstract*-Between January 2004 and November 2006, a Geographic
Information System (GIS) based habitat evaluation was conducted on the Sierra
Maderas del Carmen (MDC) and Sierra San Marcos y del Pino (SMP). The objective
of this study was to identify the most suitable sites for the restoration of desert
bighorn sheep {Ovis canadensis) in Coahuila, Mexico. Priority transplant sites were
selected based on potential contact with domestic sheep and goats and free ranging
aoudads {Ammotragus lervia), amount and juxtaposition of escape terrain, and water
availability. Priority transplant sites contain >15 km^ of escape terrain, water, and are
spatially segregated from exotic ungulates. A total of 1,159 km^ of MDC was
evaluated; 23% (271 km^) was suitable habitat for desert bighorn sheep. Two priority
transplant sites consisting of 25 and 34 km^, respectively, were delineated in MDC.
In the SMP, a total of 871 km^ was evaluated, and 20% (175 km^) was classified as
suitable habitat. One area consisting of 18 km^ was selected in SMP as a priority
transplant site.

Resumen*- De enero de 2004 a noviembre del 2006, desarrollamos un Sistema
de Informacion Geografica (SIG) para la evaluacion del habitat en las Sierras de
Maderas del Carmen (MDC) y San Marcos y del Pino (SMP) en Coahuila, Mexico.
El objetivo del estudio file el de identificar los sitios mas adecuados para la
restauracion de borrego cimarron (Ovfr canadensis). La seleccion de los sitios
prioritarios para el transplante de cimarrones frie basado en el potencial de contacto
con borregos domesticos y cabras, y exoticos ferales como los aoudads {Ammotragus
lervia), y la cantidad de la conjuncion de terreno de escape y disponibilidad de agua.
Los sitios prioritarios para un transplante tienen un area de >15 km^ de terreno de
escape, con disponibilidad de agua y estan espacialmente segregados de ungulados
exoticos. Se evaluaron 1,159 km^ en MDC; 23% (271 km^) es habitat adecuado para
borrego cimarron. Dos sitios prioritarios para transplante de 25 y 34 km^,
respectivamente, fiieron identificados en MDC. En SMP, se evaluaron 871 km^, de
los cuales el 20% (175 km^) fiie clasificado como habitat adecuado para la especie.
Un sitio de 18 km^ fiie seleccionado en SMP como area prioritaria de transplante.

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

Desert bighorn sheep (Ovis canadensis) in Mexico are classified
as “sensitive”, and a priority species for recovery and restoration
(SEMARNAP 2000). Historically, desert bighorn occurred in a
wide geographic area in the states of Chihuahua, Coahuila, Nuevo
Leon, Sonora, Baja California, and Baja California Sur (Baker
1956; Leopold 1959; Cossio 1974; Tinker 1978; Sandoval 1985).
Viable populations are still found in Sonora, Baja California, and
Baja California Sur, but the species has apparently been extirpated
from Chihuahua, Coahuila, and Nuevo Leon (Krausman et ah 1999;
Espinosa et ah 2006).

Translocations into former habitat have been widely used to
restore extirpated populations of bighorn sheep (Krausman 2000),
and transplants account for >50% of all present-day populations of
bighorn sheep (Bailey 1990). However, bighorn sheep restoration
programs can be time-consuming, costly, and bureaucratically
challenging (Zeigenfuss et al. 2000). The scarcity of sufficient seed
stock and the difficulty of rasing desert bighorn in captivity makes
it imperative that potential transplant sites be adequately assessed,
and limiting factors mitigated. A number of qualitative habitat
rating procedures have been developed to evaluate bighorn habitat
(Hansen 1980; Holl 1982; Armentrout; Brigham 1988). More
recently. Geographic Information System (GIS) and a landscape
approach have been used to increase the success of restoration
programs (Dunn 1996; Singer et al. 2000; Zeigenfuss et al. 2000;
Johnson & Swift 2000; McKinney et al. 2003; Locke et al. 2005;
Espinosa et al. 2007).

The application of GIS at the landscape perspective is an
effective and efficient means of evaluating large areas of bighorn
habitat because: (1) it provides consistent measurements across all
study areas, thereby reducing bias towards any one study area, (2) a
measurement of the amount and patchiness of bighorn habitat and a
final score and ranking can be obtained directly from the values of
the habitat components, (3) it provides for the evaluation of a large
geographic area with much less effort that if it was evaluated

ESPINOSA^T, ET AL.

directly in the field, and (4) the amount and distribution of bighorn
habitat can be displayed graphically (Espinosa et al 2007).

In Coahuila, the initial restoration of desert bighorn sheep took
place in 2000 on Sierra Maderas del Carmen (MDC) (Sandoval &
Espinosa 2001; McKinney & Delgadillo 2005). Based on habitat
suitability indices Espinosa et al. (2007), ranked MDC and Sierra
San Marcos y del Pino (SMP) in the top six potential restoration
sites in Coahuila. In addition to the availability of suitable habitat,
MDC is under federal protection (SEMARNAP 1997), as is a
portion of SMP (SEMARNAP 1999). Both areas also have active
wildlife conservation and management programs. The objective of
this study was to identify sites of minimum of 15 km^ of escape
terrain for release areas for desert bighorn sheep within MDC and
SMP.

Study Site

Sierra Maderas del Carmen (MDC) are located in extreme
northern Coahuila, at a latitude between 28° 42* 18.2849" and 29°
21’ 29.4179" N, and a longitude between 102° 22* 04,5783" and
102° 55* 04.03.6100" W (Figure 1). The Rio Grande (Rio Bravo)
separates Maderas del Carmen from Big Bend National Park,
Texas, MDC are a northeast-southwest trending mountain range,
with surrounding desert basins on the west and east sides, and is
part of the Chihuahuan Desert as defined by Brown (1982). It is a
tilted sedimentary fault block with a northeast-trending dip-slope
along with Tertiary volcanic rocks of both intrusion and extrusive
origins (Wood et al 1999). Elevations range from approximately
500 m along the Rio Grande, to 2,720 m on the highest peaks. The
terrain varies from desert flats to rugged canyons with numerous
vertical cliffs. Most precipitation occurs during summer and early
fall, and varies from approximately 100-200 mm in the foothills to
200-300 mm in the higher elevations (SEMARNAP 1997). The
annual mean temperature is approximately 22°C, with winter
temperatures dropping below 0 °C.

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Fig. 1. Location of Sierra Maderas del Carmen (MDC) and San Marcos y del Pino
(SMP) in Coahuila Mexico.

Ungulates inhabiting MDC include mule deer {Odocoileus
hemionus), white-tailed deer {Odocoileus virgianus carminis), a
reintroduced population of elk {Cervus elaphus) (Gibert et al.
2005), javelina {Pecari tajacu), and a reintroduced population of
desert bighorn sheep (McKinney & Delgadillo 2005). Major
predators include mountain lion {Puma concolor), black bear
{Ursus americanus), bobcat {Lynx rufus), and coyote {Cams
latrans).

ESPINOSA-T, ET AL.

The vegetation of MDC is highly diverse, and varies from desert
shrublands to high montane forests. The highest elevations (>2,330
m) are characterized by mixed conifer forests, dominated by
Coahuila fir {Abies coahuilensis), Douglas fir {Pseudotsuga
menziesii), and Arizona pine {Pinus arizonica). Numerous species
of oaks {Quercus spp.) are associated with the mixed conifer
forests. At mid-elevations (1,400 to 2,500 m), the forests are
replaced by woodlands dominated by oak and chaparral species
such as pointleaf manzanita (Arctostaphylos pungens), mountain
mahogany {Cercocarpus montanus), and desert ash {Fraximus
greggi). Vegetation in the lower elevations is typical of the
Chihuahuan Desert, consisting of succulent-scrub, stem succulents,
and semi-desert grasslands (Wood et al. 1999).

Sierra San Marcos y del Pino (SMP), are located in central
Coahuila, at a latitude between 26° 17' 18.5616" and 26° 53'
44.183" N, and a longitude between 101° 19' 12.25" and 102° 09'
14.1859" W (Figure 1). Desert bighorn sheep persisted in this area
until ca. 1950's (Espinosa et al. 2006). SMP is situated adjacent to
the Cuatro Cienegas Federal Wildlife and Plant Protected Area
(SEMARNAP 1999). Land ownership is a mixture of private and
ejidos (communal settlements). Of significance is Rancho Pozas
Azules from Pronatura (a non-government conservation
organization), with active ecological research and conservation
programs. Extensive livestock grazing occurs along the foothills,
and at higher elevations in the southern portion of the mountain
range.

The climate of SMP is arid, characterized by hot summers and
cool winters. Most of the precipitation occurs during winter and
varies from 100 to 440 mm, depending on elevation. Summer
temperatures may exceed 30°C, and during winter may drop below
12°C (SEMARNAP 1999). Permanent springs are found in the
northern and southern part of the mountain range. Rainfall
dependent tinajas (potholes) occur throughout the area. A
significant feature of SMP is an intermittent stream located in

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Rocillo Canyon, which bisects the central part of the mountain
range.

Large mammals include populations of white-tailed deer,
javelina, black bear, mountain lion, bobcat and coyotes.

Vegetative communities found on SMP vary greatly with
increasing elevation and resultant precipitation. Lower elevations
are characterized by desert shrubs, succulent-scrub, and stem
succulents. Succulent-scrub and stem succulent communities are
important to desert bighorn sheep, and are dominated by leaf and
stem succulents such as lechuguilla, candelilla, guapilla (Hechtia
ramillosa), ocotillo, and sangre de drago (Jatropha dioica). The
understory consists of side-oats grama and threeawns {Aristida
spp.) (Vela Coiffier 2000). Desert shrublands are replaced by
mountain shrubs, oak woodlands, and conifer forests at higher
elevations.

Methods and Materjals

The evaluation of habitat and selection of transplant sites for
desert bighorn sheep in MDC and SMP was accomplished between
January 2004 and November 2006. This analysis used combined
remote sensing imagery in the context of a Geographic Information
System (GIS), in order to identify and measure habitat components
essential to desert bighorn. Also used was the 2004 Institute
Nacional de Estadistica Geografia e Informatica (INEGI) digital
elevation models (DEM Raster Tiff Format), comprised of 30 by 30
m cells. Portions of DEM’s encompassed by both study areas were
extracted and converted to Universal Transverse Mercator (UTM)
projection, using Arcview 3.2. Slope was measured with
neighborhood analysis (Webster 1988), to obtain a measure of
escape terrain (slopes > 60%). Suitable habitat was defined as cells
between 20 and 59 % slope gradients situated < 150 m from escape
terrain (Espinosa et al. 2007). Satellite imagery (LANDSAT
ETM), with a resolution of 30 by 30 m/pixel was utilized to develop

ESPINOSA-T, ET AL.

a coverage of vegetation types preferred by bighorn sheep using a
supervised image classification procedure. Images Path030,
Row040 (March 2003) were used for MDC, and images Path029,
Row042 and Path0295 Row041 (18 September and 4 October 2000)
were used for SMP.

Satellite imagery was utilized to identify vegetation types using
the program ERJ3AS IMAGINE. Previous vegetation surveys in
MDC and SMP were used as spectral signatures in the classification
of the satellite images in developing the coverage of vegetation. In
order to delineate open vegetation preferred by desert bighorn
sheep, this study used a filtering process to identify only desert
succulent-scrub (DSS) and semi-desert grassland (SDG)
community types. The filtered vegetation file was intersected with
a coverage of slopes > 60% to create a coverage of escape terrain
with open vegetation.

Suitable habitat was defined as cells between 20 and 59% slope
gradients situated < 150 m from escape terrain, intersected with the
coverage of DSS and SDG. A coverage of impacts was digitized
from INEGI topographic maps (1:50,000) and included a 3.5 km
buffer around residential areas; 2 km buffer around paved roads;
and a 500 m buffer around secondary roads. Impacted areas were
eliminated from total suitable habitat (Espinosa et al. 2007).

Location of water sources were digitized from INEGI
topographic maps and field notes. Water sources located during
field work were marked with a Global Positioning System (GPS).
Water availability was then defined as the amount of suitable
habitat situated <3,5 km from permanent water sources (springs)
and tinajas, (temporary water sources available during the summer)
situated < 200 m of escape terrain. A coverage of water sources
was intersected with a coverage of escape terrain that had been
buffered to 200 m to create a coverage of water sources near escape
terrain. A 3.5 km buffer was then created around these water

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

sources, and this coverage was intersected with the coverage of
suitable habitat to measure water availability.

The selection of initial release sites for desert bighorn sheep was
based on the amount of escape terrain, water availability, and the
absence of exotic ungulates, the data for the absence ore presence
of exotic ungulates was obtained trough direct field verification.
Release sites should contain a minimum of 15 km^ of escape terrain
to support a viable population (McKinney et al. 2003). Several
studies have substantiated the affinity to water by desert bighorn
sheep (Dolan 2006). Given the above, only those areas containing a
minimum of 15 km^ of escape terrain and water sources were
considered as initial release sites.

A direct sampling to obtain verification points and to verify GIS
results was conduced through ground and aerial verification
surveys. The intent was to visually locate steep, broken topography
characteristic of escape terrain (slopes > 60), with open vegetation
(DSS and SDG). Secondly, it was important to refiite or
substantiate the presence of free-ranging exotic ungulates, ground
verification surveys were accomplished by vehicle or on foot. A
Cessna 182 and Robinson Raven helicopter were used for aerial
reconnaissance of the habitat, covering the entire perimeter of the
study areas at a mean distance of 100 m from the mountain side.
GPS was used to mark verification and photo points; these data was
entered in a field notebook. Finally, a review of the reports on
habitat use, movements and distribution of the transplanted free-
ranging desert bighorn population in Maderas del Carmen was
conducted.

Results

Sierra Maderas del Carwe/i. -Between January 2004 and
November 2006, GIS was implemented and verified in MDC. It
was possible to complete a habitat verification reconnaissance
overflight using fixed wing aircraft in April 2004, covering the
entire perimeter of the mountain, and obtaining 15 habitat

ESPINOSA-T, ET AL.

verification points (places where the habitat characteristics were
verified). In September 2006, a helicopter survey was conducted
covering the western, northern, and a small portion of the east side
of the mountain. During 30 days of field surveys, three water
sources were located and 26 verification points were registered.

Of the 1,159 kin^ evaluated in MDC, 270 km^ (27%) was found
to be potential habitat for desert bighorn sheep (Figure 2 in white).
Two initial release sites were delineated containing a total of 59
km^ of habitat (Figure 2 in black). The Site 1 abuts the Rio Grande,
and consists of 25 km^. Site 2 covers 34 km^ and contains Tinaja
(pothole) los Chivos, intermittent water flow and manmade water
sources in San Isidro Canyon. Site 2 actually represents the release
area for the free-ranging bighorn population in MDC, during 2004
and 2005 (McKinney & Delgadillo 2005).

Habitat use data on the free-ranging population indicate a core
use area within a radius of approximately 4 km of the release site;
identified as site 2 during this study (Figure 2). The exception was
3 adult males that dispersed >15 km to the north from the release
site within a few months following their release (B. R. McKinney
pers, comm.).

Survery did not locate exotic ungulates in the areas selected as
initial release sites (Fig 2, no. 1 and 2). However, domestic goats
attended by herders were observed in the northwestern and southern
part of MDC, approximately 5 km from the proposed bighorn sheep
release sites. According to Carlos Sifiientes (pers. comm.) the
Director of the Protected Area of MDC, free-ranging aoudad occur
in the northern part of MDC, also their presence exist across the Rio
Grande in Big Bend National Park, Texas (Skiles pers. comm.), and
in the Black Gap Wildlife Management Area, Texas (Foster 2002).

Sierra San Marcos y del Pmo.-Between January 2004 and May
2006, GIS was implemented and verified in SMP. In April 2004, a
habitat verification reconnaissance was conducted encompassing

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

Fig. 2. Potential bighorn sheep habitat (white) and priority transplant sites 1 and 2
(black) in Sierra Maderas del Carmen.

the entire periphery of the mountain using fixed-wing aircraft, and
obtained 24 verification points. During 26 days of ground surveys
30 habitat verification points and six water sources were located.

A total of 871 km^ was evaluated in SMP; 20% (174 km^) was
classified as potential bighorn sheep habitat (Figure 3 in white).
Within the potential habitat, 1 initial release site consisting of 18
km ^ was delineated (Figure 3 in black). This area contains
seasonal tinajas and one permanent spring in Quintero Canyon.

No exotics were observed in the initial release site. However,
herds of free-ranging domestic goats were observed in the southern
portion of SMP, approximately 60 km from the proposed bighorn
sheep release site, and a mixed herd of penned goats and domestic
sheep in the village of Antiguos Mineros del Norte situated <10 km
from the proposed release site. Free-ranging aoudad have not been

ESPINOSA-T, ETAL.

Fig.3. Potential bighorn sheep habitat (white) and priority transplant site (black) for
Sierra San Marcos y del Pino.

reported in SMP, however, records do exist for Sierra la Fragua
located <2 km northwest of SMP (Espinosa et al. 2006).

Discussion

Desert bighorn sheep habitat evaluation through the use of GIS
in the Chihuahuan Desert has been done in New Mexico (Dunn
1996), Texas (Locke et al. 2005), Chihuahua, Coahuila, and Nuevo
Leon (Colchero et al. 2003), and Coahuila (Espinosa et al. 2007).
The work by Espinosa et al. (2007) is most pertinent to this current
study, because they included both MDC and SMP. Espinosa et al.
(2007) conducted an evaluation and ranking of potential bighorn
sheep habitat in the known historical range of the bighorn sheep in
Coahuila. Digital elevation models with a resolution of 30 by 30
m/pixel were used to analysis topography. INEGI land use and
vegetation maps with a scale of 1:250,000 were used for vegetation

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

analysis, and INEGI topographic maps (1:50,000) to determine water
availability, that was defined as the amount of suitable habitat situated
<3.5 km from permanent water sources and tinajas, situated < 200 m
of escape terrain. The above study identified 299 km^ of potential
bighorn habitat and only 8 km^ of water availability in MDC, and
351km^ of potential habitat and 5 km^ of water availability in SMP
(Espinosa et al. 2007). The scale of data sources used in GIS habitat
evaluation studies influence the final results (Johnson & Swift 2000).
This is attested to by the results of the present study, which identified
277 and 174 km^ of potential habitat, respectively, in MDC and SMP.
This is less than the amount delineated by Espinosa et al. (2007) for
MDC and SMP, using a smaller scale vegetation data source
(1:250,000). More precise results can be obtained through the use of
higher resolution data sources, in the case of SMP the combination of
the complexity of its topography with 30 canyons, and the use of
smaller scales for the vegetation analysis (1:250,000), overestimated
the results of the potential bighorn habitat in the study of Espinosa et
al. (2007).

All permanent and temporary water sources should be identified in
the evaluation of potential habitat for desert bighorn sheep (Locke et
al. 2005). The use of more precise water sources data in this present
study resulted in the identification of a greater amount of water
availability, 59 and 18 km^, respectively, for MDC and SMP. Initial
release sites can thus be identified and quantified with more precision.

The wide-spread occurrence of goats and domestic sheep may be
the major obstacle for restoration of desert bighorn sheep in Coahuila
(Espinosa et al. 2007). These exotics have proven detrimental to
bighorn sheep due to the transmission of lethal diseases (Sandoval
1988; Gross et al. 2000; Rudolph et al. 2003; Rominger 2006). This
study documented the presence of goats and domestic sheep ranging
<15 km from sites selected for the initial release of desert bighorn
sheep. Bighorn and domestic sheep must be spatially separated a
distance of > 13.5 km to minimize the possibility of contact between
the 2 species (DBC Technical Staff 1990), this can be to close to
avoid contact between the bighorn and the domestics relatives.

ESPINOSA-T, ET AL.

maintain separation is imperative. Habitat management guidelines for
bighorn sheep and domestic goats are lacking, nonetheless, every
effort must be made to avoid contact between the two species to
minimize the potential for disease transmission.

To date no records exist of disease transmission between aoudad
and bighorn sheep. However, their habitat requirements are very
similar and social and resource competitions are possible risks on
sympatric range. Texas and New Mexico euthanize aoudads when
encountered in desert bighorn sheep habitat, the same procedures need
to be applied for Coahuila.

Development of water sources should be considered as a
management component to enhance desert bighorn sheep habitat
(Dolan 2006). Water sources for desert bighorn sheep should be
located <8 km apart and in close proximity to escape terrain (Douglas
& Leslie 1999). MDC and SMP contain a large amount of suitable
habitat for desert bighorn sheep; a major limiting factor is the lack of
available water. Dense thickets of carrizo {Arundo donax) and salt
cedar (Tamarix ramosisima), along the northern part of MDC form an
effective barrier to the Rio Grande thus making it unavailable as a
water source for desert bighorn. In SMP a paved highway presents a
potential formidable barrier to bighorn movements and high tourist
traffic at Poza de la Becerra (Cuatro Cienegas Protected Area),
effectively denies this large spring as a potential source of water.
Given the above, the authors believe that development of water
sources in MDC and SMP should be considered as part of the bighorn
restoration program in Coahuila.

Although this study did not document exotic ungulates in the
proposed bighorn sheep release sites identified in MDC and SMP,
they do occur <15 km from these sites. Notwithstanding the 13.5 km
buffer recommended by the DBC Tech Staff (1990), it is the belief of
the authors that this distance is inadequate to prevent intermingling
between bighorn sheep and exotics due to the extent and contiguous
nature of the habitat in MDC and SMP. Finally, free-ranging goat
herds should be prohibited; instead requiring daily herding and

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

penned up situations at night. The eradication of aoudad is an on¬
going activity at Black Gap Wildlife Management Area, Texas
(Pittman pers. comm.), and is under consideration in Big Bend
National Park (Skiles pers. comm.). Given the close proximity of
these two areas to MDC, the same effort needs to be applied for
MDC.

Acknowledgements

The present work was made possible thanks to financing by the
Desert Bighorn Sheep Restoration Program of CEMEX. The support
of M. Valdez of Unidos Para La Conservacion AC is appreciated, and
S. Lagham of Environmental Flights, in the aerial verifications. For
their support in San Marco y del Pino to Pronatura Noreste (Reserva
Poza Azules). The valuable support in the field of J. A. Delgadillo-
Villalobos, R. Martinez, B. R. McKinney, and B. P. McKinney
(CEMEX), is appreciated; as well as the following persons-O.
Gonzalez De Leon, M. Gonzalez F, F. Villaneuva. The present work
represents partial fulfillment of the requirements for a doctorate
degree in Biological Sciences, with a speciality in Wildlife
Management and Sustainable Development by the senior author at the
Universidad Autonoma of Nuevo Leon.

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Bailey, J. A. 1990. Management of Rocky Mountain bighorn sheep herds in Colorado.

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potenciales para la reintroduccion y el manejo del borrego cimarron {Ovis
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sheep in North America. Bonne and Crockett Club. New York, USA, 302 pp.

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sheep in vicinity of desert bighorn habitat. Desert Bighorn Council Transactions.
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desert bighorn sheep (Ovis canadensis mexicana) in Coahuila, Mexico. The
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of Historical Deset Bighorn Sheep Habitat in Coahuila, Mexico. Desert Bighorn
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poblacional de una poblacion de wapiti (Cervus elaphus) reintroducida en Maderas
del Carmen, Coahuila. Memorias del XVIII Congreso Nacional de Zoologia,
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AE at: alejandro.espinosa@cemex.com

TEXAS J. OF SCI. 61(1):31=44

FEBRUARY, 2009

GEOGRAPHIC DISTRIBUTION RECORDS FOR
SELECT FISHES OF CENTRAL AND SOUTHERN ARKANSAS

Chris T. McAllister, Renn Tumlisoii and Henry W, Robison

RapidWrite, 102 Brown Street
Hot Springs National Park, Arkansas 71913
Department of Biology, Henderson State University
Arkadeiphia, Arkansas 71999 and
Department of Biology, Southern Arkansas University
Magnolia, Arkansas 71754

Abstract.-New geographic records are documented for 16 taxa of fishes within nine
families (Anguillidae, Atherinopsidae, Catostomidae, Centrarchidae, Cyprinidae,
Fundulidae, Hiodontidae, Icturalidae, Percidae) from 20 counties of the southern portion
of Arkansas. Of these, 26 new county records for 13 (81%) of the species are presented.
Other findings include new records and range extensions for rarely collected species such
as the mooneye (Hiodon tergisus), bluehead shiner (Pteronotropis hubbsi), mud darter
{Etheostoma asprigene), swamp darter {Etheostoma fusiforme), and the undescribed
Ouachita darter (Percina sp,). Most importantly, the brown madtom, Notorus phaeus is
reported from Arkansas for only the second time since its original discovery in Columbia
County in 1972.

Robison & Buchanan (1988) provided a summation on the
geographic distribution of the fishes of Arkansas. Over the last half
decade, new geographic records for fishes of the state have been
reported (McAllister et ah 2004; 2006; 2007; 2008) to help
supplement previously published historical data. The purpose of this
report is to update the status of additional fishes of the central and
southern portions of the state.

Materials and Methods

Specimens were collected between August 1981 and April 2005
from streams throughout various localities in central and southern
Arkansas, including watersheds in 20 counties (Ashley, Bradley,
Clark, Columbia, Dallas, Drew, Garland, Hempstead, Hot Spring,
Howard, Lafayette, Little River, Miller, Montgomery, Nevada,
Ouachita, Phillips, Pike, Saline, and Union). Collections were made
with standard nylon seines (1.8 by 0.5 m and 2.7 by 0.5 m of 3.2 mm
mesh) or dipnets. Specimens were preserved in 10% formalin and
later transferred to 45% isopropanoL Specimens were field identified,

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

verified in the laboratory, and vouchers deposited in the collections at
Southern Arkansas University, Magnolia, Arkansas (SAU),
Henderson State University, Arkadelphia, Arkansas (HSU), and the
University of Louisiana-Monroe Museum of Natural History, Monroe,
Louisiana (NLU). Detailed data provided on the new sites are as
follows: (county, specific locality [township, section, and range when
available], date, museum accession number [if known], number of
specimens in parentheses, comments). In addition, several
unpublished records deposited in the NLU collection but not reported
by Robison & Buchanan (1988) are included below.

List of Species

Material examined -ThQ following is a listing of collection
localities for fishes collected in southern Arkansas.

ANGUILLIDAE

Anguilla rostrata (Lesueur) {n = 10). CLAUK CO.: SE
Arkadelphia at River Park, Ouachita River (Sec. 17, T7S, R19W). 30
April 1987. HSU 1351 (3); County Club pond, Arkadelphia. 30 March
1991. HSU 1353 (1); Ouachita River, 3.2 km S of Arkadelphia (Sec.
28, T7S, R19W). 21 March 2003. HSU 2893 (1). HOT SPRING
CO.: Ouachita River (Sec. 31, T3S, R17W). 15 April 1997. HSU
1979, 2099 (2); Remmel Dam at Jones Mill (Sec. 36, T3S, R18W). 13
May 1997. HSU 2674 (1). LITTLE RIVER CO.: Little River, river
run E of Dam (Sec. 26, T12S, R28W). 21 April 1991. HSU 1081 (1).
OUACHITA CO.: Little Missouri River, 1.6 km N of Reader on
unnamed gravel road (Sec. 18, T1 IS, R18W). 19 February 1983. NLU
(1). These specimens supplement previous records from the state.
American eel populations have experienced a drastic decline in
Arkansas due to construction of dams which block migration in large
rivers.

HIODONTIDAE

Hiodon tergisus Lesueur (n = 3). OUACHITA CO.: 24.1 km NE
of Chidester at confluence of Little Missouri and Ouachita rivers at
Tates Bluff Recreation Area (Sec. 1, TllS, R18W). 19 February
1983. NLU (1). UNION CO.: Ouachita River at U.S. 167 (Sec. 10,

MCALLISTER, TUMLISON & ROBISON

T16S, R14W). 26 May 1997. SAU (1); Ouachita River at U.S. 82
(Sec. 14, T18S, RIOWS). 17 July 1999. SAU (1). This large river
species is rarely encountered in Arkansas. These two new county
records document significant collections for the lower Ouachita River.

CYPRINIDAE

Notropis maculatus (Hay) {n = 48). BRADLEY CO.: Snake
Creek at Broad (Sec. 30, T16S, R9W). 17 June 2002. SAU (3).
COLUMBIA CO.: Dorcheat Bayou at co. rd., 4.8 km SW of
Philadelphia (Sec. 16, T18S, R22W). 4 September 1993. SAU (1);
Dorcheat Bayou at St. Hwy. 160, 6.4 km E of Taylor (Sec. 9, T19S,
R22W). 19 May 2004. SAU (2). DREW CO.: Cut-Off Creek at St.
Hwy. 35, 1.1 km E of Collins (Sec. 31, TBS, R4W). 13 April 1993.
SAU (2). LAFAYETTE CO.: Bodcau Creek at co. rd„ 1.6 km N of
Lewisville (Sec. 7, T15S, R23W). 5 July 1992. SAU (1); Bayou
Bodcau at U.S. 82 (Sec. 7, T16S, R23W). 11 October 1995. SAU
(2). LITTLE RIVER CO.: Cypress Creek at St. Hwy. 234 in
Winthrop (Sec. 7, TllS, R31W). 6 June 1989. SAU (1); Little River
backwater at U.S. 71, 3,2 km N of Wilton (Sec. 24, TllS, R29W). 5
October 2001. SAU (2). NEVADA CO.: Middle Creek, 14.5 km N
of Prescott on St. Hwy. 19 (Sec. 27, T9S, R23W). 19 February 1983.
NLU (1); Terre Rouge Creek, 11.3 km SE of Prescott on St. Hwy. 24
(Sec. 3, TBS, R22W). 19 February 1983. NLU (1); Caney Creek, 4.8
km N of Bluff City on St. Hwy. 24 (Sec. 22, TllS, R20W). 19
February 1983. NLU (26). UNION CO.: Smackover Creek at co. rd.
68, 3.2 km N of Norphlet (Sec. 3, TBS, R15W). 20 September 1992.
SAU (1); Grand Marais Lake at Felsenthal (Sec. 16, TBS, RlOW).
18 September 1996. SAU (5). Collections of this lowland cyprinid
from Lafayette and Nevada counties represent new county records.
This shiner was generally collected from quiet, backwater areas
devoid of current but often with vegetation.

Notropis texanus (Girard) (n = 6). COLUMBIA CO.: Dorcheat
Bayou at St, Hwy. 160, 6.4 km E of Taylor (Sec. 9, TBS, R22W). 19
May 2004. SAU (1). LAFAYETTE CO.: Bayou Bodcau at U.S. 82
(Sec. 7, TBS, R23W). 11 October 1995. SAU (1). UNION CO.:
Smackover Creek at Co. Rd. 68, 3.2 km N of Norphlet (Sec. 3, TBS,

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

R15W). 20 September 1992. SAU (1); Big Comie Creek at St. Hwy.
15, 25.7 km SW of El Dorado (Sec. 35, T19S, R18W). 5 October
1995. SAU (3). Specimens from Lafayette County document a new
county record and represent the southwestemmost locality of N.
texanus in the state.

Pteronotropis hubbsi (Bailey & Robison) {n = 24). ASHLEY
CO.: Thompson Creek, 11.3 km NW Crossett (Sec. 11, T17S, R9W).
5 July 1996. HSU 1340 (13); slough off Saline River at Stillions, 14.5
km NW Crossett (Sec. 4, T17S, R9W). 5 July 1996. HSU 1319 (1);
Ouachita River, 9.7 km W Crossett at U.S. 82 (Sec. 14, T18S, RlOW).
11 April 1997. HSU 2008 (3). NEVADA CO.: Middle Creek, 14.5
km N of Prescott on St. Hwy. 19 (Sec. 27, T9S, R23W). 19 February
1983. NLU (1); Caney Creek, 4.8 km N of Bluff City on St. Hwy. 24
(Sec. 22, TllS, R20W). 19 February 1983. NLU (6). This species
was formerly included in the genus Notropis; the subgenus
Pteronotropis was elevated to genus rank by May den (1989). The
bluehead shiner has a spotty distribution in lowlands of the Red and
Ouachita river systems of southern Arkansas and was previously
known from only 10 localities in the state, including two disjunct sites
in the Little River system (Bailey & Robison 1978; Robison &
Buchanan 1988). This shiner was also listed as a species of special
concern in Arkansas (Robison & Buchanan 1988; Anonymous 2004)
and the Nature Conservancy considers P. hubbsi vulnerable (S3) in
the state (NatureServe 2008). This study documents two new county
records in tributaries of the Little Missouri and Saline rivers.

Semotilus atromaculatus (Mitchill) {n = 134). ASHLEY CO.:
Ouachita River, 10.5 km W Crossett off St. Hwy. 82 (Sec. 14, T18S,
RlOW). 11 April 1999. HSU 2009 (1). CLARK CO.: unnamed
tributary to Little Deceiper Creek, 8.0 km W Arkadelphia (Sec. 24,
T7S, R20W). 8 February 1997. HSU 1681 (3). DALLAS CO.:
unnamed tributary to L’Eau Frais Creek (Sec. 8, T7S, R17W). 5 April
1997. HSU 2118 (4). GARLAND CO.: tributary to Pleasant Run
Creek, 2.4 km S Lonsdale (Sec. 26, T2S, R17W). 29 March 1991.
HSU 286 (2); Cooper Creek, 1.6 km S of jet. St. Hwys. 171 & 290
(Sec. 6, T4S, R18W). 20 August 1993 & 29 July 1995. HSU 99, 1 1 14

MCALLISTER, TUMLISON & ROBISON

(7, 1); Cooper Creek, 2.4 km S jet. St. Hwy. 171 & 290 (Sec. 8, T4S,
R18W). 23 August 1993. HSU 189 (2). HEMPSTEAD CO.: Bois
d’Arc Creek, 3.2 km SW jet. St. Hwys. 73 & 195 (Sec. 13, T12S,
R26W). 6 April 1991. HSU 1231 (1). HOT SPRING CO.; Prairie
Bayou, 3.2 km NW of New DeRoche (Sec. 31, T4S, R19W). 20
August 1993. HSU 103 (1); 9.7 km W of Malvern at Blakely Creek
(Sec. 15, T4S, R18W). 20 August 1993. HSU 1312 (1); 8.0 km NW
Bismarck at Valley Creek (Sec. 28, T4S, R21W). 27 August 1993.
HSU 165 (3); 4.8 km N Bismarck at Big Hill Creek (Sec. 30, T4S,
R20W). 27 August 1993. HSU 170 (15); Mt. Carmel Creek, 7.2 km N
jet. St. Hwy. 84 & 128 (Sec. 17, T4S, R19W). HSU 181 (6); tributary
of Big Hill Creek, 3.2 km N of Lambert (Sec. 1, T5S, R21W). 5
February 1994. HSU 980 (3); uimamed creek, 8.0 km NW of Lambert
(Sec. 28, T4S, R21W). 5 February 1994. HSU 984 (2); unnamed
creek, 4.8 km NE of jet. St. Hwy. 84 & 128 (Sec. 25, T4S, R19W). 20
March 1994. HSU 1001 (4); Mt. Carmel Creek at St. Hwy. 128 bridge
(Sec. 17, T4S, R19W). 21 July 1995. HSU 1193 (1); Curl Creek, 5.2
km W of St. Hwy. 128 on Land Camp Rd (Sec. 36, T4S, R20W). 22
July 1995. HSU 1102 (1); tributary to Prairie Bayou, 4.8 km W St.
Hwy. 128 (Sec. 25, T4S, R20W). 22 July 1995. HSU 1127, 1135 (1,
4); tributary to Valley Creek (Sec. 35, T4S, R21W). 28 July 1995.
HSU 1155 (1); tributary to Valley Creek (Sec. 26, T4S, R21W). 28
July 1995. HSU 1166 (1); tributary to Blakely Creek, 0.8 km S
O’Neal Trail (Sec. 17, T4S, R18W). 29 July 1995. HSU 1 147 (1); Big
Hill Creek at Tower Road (Sec. 30, T4S, R20W). 4 August 1995.
HSU 1163 (1); Curl Creek (Sec. 27, T4S, R20W). 4 August 1995.
HSU 1169, 1172 (4, 4); tributary to Ouachita River (Sec. 7, T4S,
R17W). 16 February 1997. HSU 1469 (4); Stone Quarry Creek at U.S.
270 (Sec. 32, T3S, R17W). 1 April 1997. HSU 1593 (1); Dyer Creek
at end of Jenney Lane (Sec. 36, T3S, R17W). 7 March 1999. HSU
2531 (4); Curl Creek, 6.4 km NE Bismarck (Sec. 27, T4S, R20W). 15
January 1999. HSU 2533 (2); Tigger Creek, 1.6 km from county line
off U.S. 270E (Sec. 31, T4S, R20W). 10 April 1999. HSU 2672 (1).
HOWARD CO.: 6.4 km W Dierks at Saline River near Bluff Creek
(Sec. 23, T7S, R29W). 2 March 1991. HSU 1230 (1). LITTLE
RIVER CO.: Little River at wall of dam (Sec. 26, T12S, R28W). 23
February 1991. HSU 1233 (1). MONTGOMERY CO.: Dry Mazam

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

Creek (Sec. 14, T3S, R24W). 4 May 1991. HSU 42 (8); Little
Missouri River at Little Missouri Falls (Sec. 6, T4S, R27W). 1 May
1993. HSU 159 (2); Collier Creek at St. Hwy. 8, 2.4 km N Caddo Gap
(Sec. 12, T4S, R25W). HSU 261 (3); Lick Creek, tributary to Caddo
River (Sec. 24, T3S, R26W). 2 April 1994. HSU 370 (5); tributary of
Little Missouri River at dirt rd. (Sec. 27, T4S, R27W). 19 April 1991.
HSU 1607 (1); Greasy Branch (Sec. 35, T4S, R27W). 2 May 1997.
HSU 2144 (3). NEVADA CO.: Middle Creek, 14.5 km N of Prescott
on St. Hwy. 19 (Sec. 27, T9S, R23W). 19 February 1983. NLU (1).
PIKE CO.: spring drainage ditch, 7.2 km NE of Murfreesboro at
Wayside Park off Co. Rd. 379 (Sec. 35, T7S, R25W). 18 June 1983.
NLU (1); Little Missouri River (Sec. 5, T5S, R27W). 16 March 1997.
HSU 2023 (1); 1.6 km N of Langley on St. Hwy. 269 at first bridge
(Sec. 12, T5S, R27W). 16 March 1997. HSU 1574 (2); Bear Creek
(Sec. 34, T5S, R25W). 20 April 1997. HSU 1928 (2); Rock Creek, 1.6
km SE of Salem off St. Hwy. 70 (Sec. 18S, T5S, R24W). 20 April
1997. HSU 1628 (3); Wolf Creek at St. Hwy. 29 bridge, 1.6 km SW
Antoine (Sec. 27, T8S, R23W). 17 April 1999. HSU 2628 (1).
SALINE CO.: unnamed creek at 1106 West Place in Benton (Sec. 1,
T2S, R15W). 13 March 1994. HSU 482 (6); tributary to McNeil
Creek, 0.4 km N of 1-30 on Congo Rd. (Sec. 35, TIS, R15W). 13
March 1994. HSU 509 (2); Brushy Creek, 6.4 km NW Exit 106 off I-
30 at Fairplay Rd. (Sec. 34, T2S, R16W). 5 April 1994. HSU 498 (2);
Brushy Creek, 4.8 km NW Exit 106 off 1-30 at Fairplay Rd. (Sec. 35,
T2S, R16W). 29 April 1994. HSU 811 (1); unnamed creek behind
Watson Place in Benton (Sec. 30, TIS, R15W). 29 April 1994. HSU
1053 (1). New county records for this headwater cyprinid are
reported for Ashley, Hempstead, Little River, and Nevada counties,
and the records from Howard and Pike counties represent the first
collections since 1960 (see Robison & Buchanan 1988). The creek
chub inhabits the smallest, clearer headwater streams.

CATOSTOMIDAE

Moxostoma poecilurum (Jordan) {n = 13). CLARK CO.: L’Eau
Frais Creek at St. Hwy. 7, 8.9 km S Jet. St. Hwy 51 and 7 (Sec. 1,
T8S, R18W). 25 March 1994. HSU 543 (1); L’Eau Frais Creek at St.
Hwy 128, 0.8 km of Joan (Sec. 22, T7S, R18W). 2 May 1994 and 1

MCALLISTER, TUMLISON & ROBISON

March 2006, HSU 575, 3164 (1, 3); Tupelo Creek at St Hwy, 7
bridge (Sec. 35, T7S, R19W). 11 March 2005. HSU 2927 (1).
COLUMBIA CO.: Smackover Creek at U.S. Hwy 79, 5.6 km NE of
McNeil (Sec. 26, T15S, R20W). 25 September 1991. SAU (1); Big
Creek at St Hwy. 98, 6.4 km S of Village (Sec. 3, T18S, R19W). 5
November 1993. SAU (1); Sloan Creek at St. Hwy. 57 (Sec. 11,
T16S, R19W). 5 November 1993. SAU (1). NEVADA CO.:

Grassy Lake, 9.7 km NE of Prescott off St. Hwy. 67 (Sec. 1, TIOS,
R22W). 2 August 1981. NLU (1); Middle Creek, 14.5 km N of
Prescott on St. Hwy. 19 (Sec. 27, T9S, R23W). 19 February 1983.
NLU (1); Caney Creek, 4.8 km N of Bluff City on St. Hwy. 24 (Sec.
22, TllS, R20W). 19 February 1983. NLU (1). UNION CO.: Big
Comie Creek at St. Hwy. 15, 25.7 km SW of El Dorado (Sec. 35,
T19S, R18W). 5 October 1995. SAU (1). The blacktail redhorse is
generally confined to the southern Coastal Plain streams of the lower
Ouachita River system of southcentral Arkansas (Robison &
Buchanan 1988). This is the first time M poecilurum has been
reported from Union County.

ICTALURJDAE

Noturus phaeus Taylor {n = 1). COLUMBIA CO.: Horsehead
Creek at U.S. 19, 12.9 km SW of Magnolia (Sec. 32, T18S, R21W).
21 November 2001. SAU (1). To the authors' knowledge, this
madtom had not been collected in the state since 1972 when Robison
(1974) reported three specimens from a spring-fed tributary of
Horsehead Creek in Columbia County, 3.5 km S Macedonia (Sec, 29,
T18S, R21W). Subsequently, it was previously listed as endangered
and considered extremely rare by Robison & Buchanan (1988).
However, N. phaeus is not currently listed in any category by the
Arkansas Game and Fish Commission. Interestingly, the Nature
Conservancy lists the species as SH (possibly extirpated) in Arkansas
(NatureServe 2008).

FUNDULIDAE

Fundulus dispar (Agassiz) {n = 40), ASHLEY CO.: Thompson
Creek, 11.3 km NW Crossett (Sec. 11, T18S, R9W). 5 July 1996.
HSU 1347 (6); Ouachita River, 10.5 km W Crossett at St. Hwy 82

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(Sec. 14, T18S, RlOW). 11 April 1997. HSU 2004 (3). BRADLEY
CO.: Moro Creek at Moro Bay State Park (Sec. 21, T16S, R12W).
17 October 1998. SAU (1); Snake Creek at Broad (Sec. 30, T16S,
R9W). 17 June 2002. SAU (1); L’Aigle Creek at co. rd., 14.5 km
South of Hermitage (Sec. 18, T16S, RlOW). 10 June 2005. SAU (2).
CLARK CO.: Tupelo Creek (T7S, R19W). 25 February 1997. HSU
2337 (1). HOT SPRING CO.: tributary to Ouachita River under
bridge (Sec. 22, T6S, R18W). 2 February 1997. HSU 2237 (1).
OUACHITA CO.: Bragg Lake, 2.0 km SE Bragg City at St. Hwy. 24
(Sec. 33, T12S, R18W). 12 April 1997. HSU 2051 (4); Freeo Creek at
St. Hwy. 9, 3.2 km S Dallas County line (Sec. 36, TllS, R16W). 7
June 1997. HSU 2178 (4). NEVADA CO.: Caney Creek, 4.8 km N
of Bluff City on St. Hwy. 24 (Sec. 22, TllS, R20W). 19 February
1983. NLU (8). UNION CO.: Lapile Creek at North Road, 8.0 km N
of Huttig (Sec. 25, T18S, RllW). 16 June 1992. SAU (1); Lapile
Creek at Lapile Road (Sec. 31, T18S, RllW). 16 June 1992. SAU
(1). Grand Marais Lake at Felsenthal, (Sec. 16, T19S, RlOW). 18
September 1996. SAU (2); Shallow Lake, lower Ouachita River
backwater (Sec. 37, T19S, RlOW). 20 January 1999. HSU 2256 (5).
This study documents a new county record for this lowland
topminnow in Nevada County.

ATHERINOPSIDAE

Menidia audens Hay {n = 47). ASHLEY CO.: Thompson Creek,
11.3 km NW Crossett (Sec. 11, T17S, R9W). 5 June 1996. HSU 894
(1). BRADLEY CO.: Moro Creek at Moro Bay State Park (Sec. 21,
T16S, R12W). 17 October 1998. SAU (3). CLARK CO.: Caddo
River at St. Hwy 67 bridge, Caddo Valley (Sec. 31, T6S, R20W). 10
& 26 April 1994. HSU 473, 520 (1, 2); DeRoche Creek at St. Hwy 28
bridge (Sec. 7, T6S, R19W). 20 March 1994. HSU 839 (4); Saline
Bayou bridge at St. Hwy 7 (Sec. 16, T7S, R19W). 31 August 2005.
HSU 3129 (1). DALLAS CO.: unnamed creek at St. Hwy 128, 6.4
km N of Sparkman (Sec. 35, T9S, R17W). 20 January 1997. HSU
1344 (2); Tulip Creek at St. Hwy 8 (Sec. 27, T8S, R16W). 18
February 1999. HSU 2371 (3). HOT SPRING CO.: Lake DeGray at
Lenox Marcus crossing. June 1993. HSU 137 (20). LITTLE RIVER
CO.: Little River, run E of dam (Sec. 26, T12S, R28W). 16 February

MCALLISTER, TLMLISON & ROBISON

1991 & 14 April 199L HSU 289, 27 (1, 6); Little River at Millwood
Dam (Sec. 26 T12S, R28W). March 1991. HSU 313 (1). PHILLIPS
CO.: Old Town Lake, flood gate of St. Hwy. 44 (Sec. 30, T3S, R3E).
23 March 2005. HSU 3002 (2). Suttkus & Thompson (2002)
discussed the rediscovery of this silverside in the Pearl River of
Louisiana and Mississippi and provided compelling evidence that M
audens is a valid species. New county records are reported for
Ashley, Bradley, Clark, and Hot Spring counties.

CENTRARCHIDAE

Centrarchus macropterus (Lacepede) {n = 18). ASHLEY CO.:
slough off Saline River at Stillions, 14.5 km NW Crossett (Sec. 4,
T17S, R9W). 5 July 1996. HSU 1317 (1); tributary to Hanks Creek,
12.1 km E Crossett (Sec. 21, T18S, R7W). 5 July 1996. HSU 1368
(4). CLARK CO.: McNeeley Creek, 6.4 km S Beime off St. Hwy.
51 (Sec. 31, TIOS, R20W). 20 April 1997. HSU 2156 (1); Clear Lake
in Joan off St Hwy. 51 (Sec. 23, T7S, R18W). 8 May 1999. HSU
2649 (1); L'Eau Frais Creek at Ouachita River (Sec. 1, T8S, R19W).
11 March 2005. HSU 3038 (1). HEMPSTEAD CO.: Yellow Creek,
0.4 km from railroad on St Hwy. 32 (Sec. 17, T12S, R27W). 23
March 1991. HSU 1235 (1). HOT SPRING CO.: L’Eau Frais Creek
at St Hwy. 222 (Sec. 15, T6S, R17W). 9 March 1997. HSU 1884 (1).
OUACHITA CO.: Freeo Creek at St. Hwy. 9, 8.9 km S of Dallas
County line (Sec, 36, T22S, R16W). 7 June 1997. HSU 2175 (1).
NEVADA CO.: Caney Creek, 4.8 km N of Bluff City on St. Hwy, 24
(Sec. 22, TllS, R20W). 19 February 1983. NLU (4); Middle Creek,
14,5 km N of Prescott on St Hwy. 19 (Sec. 27, T9S, R23W). 19
February 1983, NLU (1), UNION CO.: Lapoile Creek, 5.2 km NE
Strong (Sec. 18, T18S, R1 1 W). 22 March 1997. HSU 1688 (2), These
new records supplement those of McAllister et at (2004); however,
Hempstead is a new county record.

Lepomis marginatus (Holbrook) {n = 117). ASHLEY CO.:
Thompson Creek, 1 1,3 km NW Crossett (Sec. 11, T17S, R9W). 5 July
1996, HSU 1346 (4); tributary to Hanks Creek, 12.1 km E Crossett
(Sec. 21, T18S, R7W), 5 July 1996. HSU 1367 (9); slough off Saline
River, 14.5 km NW Crossett at Stillions (Sec. 1, T17S, R9W). 6 July
1996. HSU 1364 (1). BRADLEY CO.: Snake Creek at Broad (Sec.

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

30, T16S, R9W). 17 June 2002. SAU (4). CLARK CO.: Tupelo
Creek, 6.4 km S Arkadelphia at St. Hwy. 7 bridge (Sec. 35, T7S,
R19W). 22 March 1997 & 4 May 1999. HSU 1791, 2349 (2, 2);
Beech Creek, 3.2 km SE Gurdon (Sec. 11, TIOS, R20W). 27 March
1997. HSU 1644 (2); McNeeley Creek, 6.4 km S Beime off St. Hwy.
51 (Sec. 31, TIOS, R20W). 20 April 1997. HSU 2162 (3).
COLUMBIA CO.: Horsehead Creek at U.S. 19, 12.9 km SW of
Magnolia (Sec. 32, T18S, R21W). 21 November 2001. SAU (6).
LAFAYETTE CO.: Bayou Bodcau at U.S. 82 (Sec. 7, T16S,
R23W). 11 October 1995. SAU (1). MILLER CO.: Millwood
Lake at Paraloma Landing at end of St. Hwy. 234 (Sec. 29, TllS,
R28W). 18 June 2002. SAU (3). NEVADA CO.: Caney Creek, 4,8
km N of Bluff City on St. Hwy. 24 (Sec. 22, TllS, R20W). 19
February 1983. NLU (54). PIKE CO.: Saline River, 8.0 km W of
Delight on St. Hwy. 26 (Sec. 8, T8S, R24W). 19 June 1982. NLU
(1). UNION CO.: Big Comie Creek at St. Hwy. 15, 25.7 km SW of
El Dorado (Sec. 35, T19S, R18W). 5 October 1995. SAU (1); Grand
Marais Lake at Felsenthal (Sec. 16, T19S, RlOW). 18 September
1996. SAU(17);CalionLakeatCalion(Sec. 22, T16S, R14W). 17
May 2003. SAU (7). The dollar sunfish is a widespread species of
the Coastal Plain province in Arkansas. The specimen from Pike
County represents a new county record.

Lepomis miniatus Jordan {n = 42). CLARK CO.: Little Deceiper
Creek on St. Hwy. 51, 1.6 km W of Arkadelphia (Sec. 26, T7S,
R20W). June 1992. HSU 128 (1); Caddo River at St. Hwy. 7 bridge
in Caddo Valley (Sec. 31, T6S, R20W). 20 March & 17 April 1994, 1
April 1997. HSU 828, 740, 1937 (1, 1, 1); Caddo River, 3.2 km W of
1-30 bridge in Caddo Valley (Sec. 36, T6S, R20W). 4 May 1994. HSU
775 (1); Tupelo Creek, 6.4 km SE of Arkadelphia off St. Hwy. 7 (Sec.
35, T7S, R19W). 24 February 1995, 25 February 1999, & 11 March
2005. HSU 1240, 2345, 2350, 2979 (1, 2, 1, 1); Gentry Creek (boat
ditch), 6.4 km N Gurdon off U.S. 67 (Sec. 3, T9S, R20W). 13 January
1996. HSU 1276 (1); Saline Bayou, 1.6 km E Arkadelphia on St.
Hwy. 7 (Sec. 16, T7S, R19W). 2 March 1996. HSU 1283 (1); lower
dam of Caddo River at Caddo Valley (Sec. 36, T6S, R20W). 25
February & 1 March 2005. HSU 2899, 2909 (1, 1). COLUMBIA
CO.: Bayou Dorcheat Creek at St. Hwy. 160, 6.4 km E of Taylor

MCALLISTER, TLMLISON & ROBISON

(Sec. 5, T19S, R22W). 6 October 2003. SAU (2); Horsehead Creek
at U.S, 19, 12.9 km SW of Magnolia (Sec. 32, T18S, R21W). 21
November 2001. SAU (1). GARLAND CO.: mouth of Hot Springs
Creek, Lake Hamilton (T3S, R18W). 30 April 1999. HSU 2279 (1).
HOT SPRING CO.: U Eau Frais Creek, 12,1 km E Donaldson (Sec,
1, T6S, R17W). 22 March 1997. HSU 1493 (1); Saline Bayou, 4.0 km
S Friendship (Sec. 23, T6S, R19W). HSU 2719 (1). LAFAYETTE
CO.: Lake Erling at St Hwy. 160 (Sec. 35, T19S, R23W). 14 June
2002. SAU (5). NEVADA CO.: Middle Creek, 14.5 km N of
Prescott on St. Hwy. 19 (Sec. 27, T9S, R23W). 19 February 1983.
NLU (2); Caney Creek, 4.8 km N of Bluff City on St. Hwy. 24 (Sec.
22, Ills, R20W). 19 February 1983. NLU (3). OUACHITA CO.:
Bragg Lake, 2 km SE Bragg City off St. Hwy. 24 (Sec. 33, T12S,
R18W). 12 April 1997. HSU 2053 (1); Freeo Creek at St Hwy. 9, 8.9
km S Dallas County line (Sec. 36, TllS, R16W). 7 June 1997. HSU
2717 (3). UNION CO.: Grand Marais Lake at Felsenthal (Sec. 16,
T19S, RlOW). 18 September 1996. SAU (3); Big Comie Creek at St.
Hwy. 15, 9.7 km SW of El Dorado (Sec. 35, T19S, R18W). 5

October 1995. SAU (4); Calion Lake at Calion (Sec. 22, T16S,
R14W). 17 May 2003. SAU (1). This fish was formerly regarded as
a subspecies of Lepomis punctatus. However, Warren (1992)
examined morphological variation and considered L. punctatus
miniatus a separate species. New county records for the red-spotted
sunfish are documented for Garland and Nevada counties.

PERCIDAE

Etheostoma asprigene (Forbes) (n = 9). BRADLEY CO.: More
Creek at Moro Bay State Park (Sec. 21, T16S, R12W). 17 October
1998. SAU (1). HEMPSTEAD CO.: Ozan Creek, 1.1 km W of
McCaskill on St. Hwy. 24 (Sec. 28, T9S, R26W). 26 Sept. 1983.
NLU (2). HOWARD CO.: Muddy Fork Creek, 12.9 km E of Dierks
on FAS south of Muddy Fork (Sec. 33, T7S, R27W). 21 Nov. 1981.
NLU (1). UNION CO.: Grand Marais Lake at Felsenthal (Sec. 16,
T19S, RlOW). 18 September 1996. SAU (5). Hempstead and
Howard counties represent significant southwestward range
extensions as well as the westernmost distribution for E. asprigene in
the state.

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

Etheostoma fusiforme (Girard) {n = 15). COLUMBIA CO.:
Bayou Dorcheat at co. rd., 6.4 km NE of Bussey (Sec. 16, T18S,
R22W). 6 October 2003. SAU (2). NEVADA CO.: Caney Creek,
4.8 km N of Bluff City on St. Hwy. 24 (Sec. 22, TllS, R20W). 19
February 1983. NLU (1). PIKE CO.: spring drainage ditch, 7.2 km
NE of Murfreesboro at Wayside Park off co. rd. 379 (Sec. 35, T7S,
R25W). 18 June 1983. NLU (6). UNION CO.: Grand Marais Lake at
Felsenthal (Sec. 16, T19S, RlOW). 18 September 1996. SAU (2);
Norris Creek at NE comer of Strong (Sec. 32, T18S, R12W). 22
March 1997. HSU 2132 (1); Calion Lake at Calion (Sec. 22, T16S,
R14W). 17 May 2003. SAU (3). Robison & Buchanan (1988)
considered the swamp darter to be a threatened species in the state and
the Nature Conservancy (NatureServe 2008) lists it as imperiled (S2).
Few previous records exist for E. fusiforme in Arkansas, and new
county records for Columbia, Nevada, and Pike counties significantly
add to the knowledge of its current distribution in the state.

Percina sp. {n = 3). CLARK CO.: Caddo River, 3.2 km W of 1-30
bridge in Caddo Valley (Sec. 36, T6S, R20W). J. Hardage. 4 May
1994. HSU 767 (1). HOT SPRING CO.: Ouachita River, river road
at Friendship (Sec. 12, T6S, R19W). 27 April 2005. HSU 3102 (2).
This undescribed species, an endemic of the Ouachita River, is one of
special concern (Robison & Buchanan 1988; Robison 1992;
Anonymous 2004). The Clark County record suggests that the former
lower Caddo River population may not be extirpated by the tailwater
effects of DeGray Dam, a suggestion offered by Buchanan (1984) and
reiterated by Robison & Buchanan (1988). More recently, Gagen et
al. (2002) provided information on habitat and abundance of the
Ouachita darter. This percid species is currently being described by
HWR and R. C. Cashner.

Results and Summary

This study documents the collection of 530 specimens,
representing 16 taxa within nine families. The following noteworthy
species were collected: American eel {A. rostrata), mooneye (//.
tergisus), bluehead shiner (P. hubbsi), taillight shiner (N. maculatus),
weed shiner {N. texanus), creek chub {S. atromaculatus), blacktail

MCALLISTER, TUMLISON & ROBISON

redhorse (M poecilurum), brown madtom {N. phaeus), flier (C
macropterus), dollar sunfish (L. marginatus), redspotted sunfish (Z.
miniatus), northern starhead topminnow (F, dispar), mississippi
silverside (M audens), mud darter (Z. asprigene), swamp darter {E.
fusiforme), and the undescribed Ouachita darter (Percina sp.).

In summary, this study documents 26 new county records for 13
(81%) of the species collected. Most importantly, this study includes
new records and extensions of the known geographic ranges for rarely
collected species, including H. tergisus, P. hubbsi, E. asprigene, E.
fusiforme, and Percina sp. nov. nr, nasuta, while N. phaeus is reported
from Arkansas for only the second time since its original discovery
more than 35 years ago.

Acknowledgments

Special thanks to Dr. N. H. Douglas, University of Louisiana-
Monroe for use of the collection of fishes housed in the ULM
Museum of Natural History, including those collected by R. A. Loe.
Also, thanks to previous SAU Vertebrate Natural History classes, and
former SAU students J. Rader, C. Brummett, N. Covington, and K.
Ball, and HSU ichthyology classes, and former HSU students J.
Abernathy, K. Bailey, B. Baker, B. Chancellor, M. Clark, J. & D.
Collins, S. Davis, W. Daggett, J. Daniel, A. DeLaughter, R. Dorer, D.
Dyer, D. Fendley, R. Fisher, B. Fluker, J. Hardage, S. Henson, B,
Hesington, R. Hicks, J. Hooks, C. Horton, T. James, S. Jordan, J.
Jumper, R. Long, J. Nix, J. Patterson, C. Petty, J. Pinkerton, A.
Rainwater, C. Pope, J. Rigsby, J. Russell, S. Ryders, L. Self, D. & J.
Thompson, D. Turner, K. Watt, T. West, and A. Weston for assistance
in collecting. We also thank the Arkansas Game and Fish
Commission for providing scientific collecting permits to HWR and
RT.

Literature Cited

Anonymous. 2004. Arkansas endangered, threatened, and species of special concern.

Arkansas Game & Fish Comm. Rep., January 9, 2004. 6 pp.

Bailey, R. M. & H. W. Robison. 1978. Notropis hubbsi, a new cyprinid from the
Mississippi River basin. Occas. Pap. Mus. Zook, Univ. Michigan, 683:1-21.

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

Buchanan, T. M. 1984. Status of the Longnose Darter, Percina nasuta (Bailey), in
Arkansas. Arkansas Nat. Hist. Comm. Rep., June 1984, 29 pp.

Gagen, C. J., K. R. Moles, L. J. Hlass & R. W. Standage. 2002. Habitat and
abundance of the Ouachita Darter {Percina sp. nov.). J. Arkansas Acad. Sci.,
56:230-234.

Mayden, R. L. 1989. Phylogenetic studies of North American minnows, with emphasis
on the genus Cyprinella (Teleostei: Cypriniformes). Univ. Kansas Mus. Nat. Hist.
Misc. Publ., 80:1-189.

McAllister, C. T., S. E. Barclay & H. W. Robison. 2004. Geographic distribution
records for the Flier, Centrarchiis macropterus (Perciformes: Centrarchidae), from
southwestern Arkansas. J. Arkansas Acad. Sci., 58:131-132.

McAllister, C. T., H. W. Robison & T. M. Buchanan. 2006. Noteworthy geographic
distribution records for the Golden Topminnow, Fundulus chrysotus
(Cyprinodontiformes: Fundulidae), from Arkansas. J. Arkansas Acad. Sci. 60:185-
188.

McAllister, C. T., H. W. Robison & R. Tumlison. 2007. Additional geographic records
for the Goldstripe Darter, Etheostoma parvipinne (Perciformes: Percidae), from
Arkansas. J. Arkansas Acad. Sci., 61: 125-127.

McAllister, C. T., R. Tumlison & H. W. Robison. 2008. Distribution of the Bantam
Sunfish, Lepomis symmetricus (Perciformes: Centrarchidae), in Arkansas. Texas J.
Sci., 60(l):23-32.

NatureServe, 2008. NatureServe Explorer: An online encyclopedia of life [web
application]. Version 7.0. NatureServe, Arlington, Virginia. Available
http://www.natureserve.org/explorer. (Accessed: June 29, 2008).

Robison, H. W. 1974. First record of the ictalurid catfish, Noturus phaeus, from
Arkansas. Southwest. Nat., 18:475.

Robison, H. W. 1992. Distribution and status of the Ouachita River form of the
Longnose Darter in the Ouachita National Forest, Arkansas. Final report submitted
to U.S.D.A. Forest Service, Ouachita National Forest, Hot Springs, Arkansas, 58 pp.

Robison, H. W. & T. M. Buchanan. 1988. Fishes of Arkansas. Univ. Arkansas Press,
Fayetteville, 536 pp.

Suttkus, R. D. & B. A. Thompson. 2002. The rediscovery of the Mississippi Silverside,
Menidia audens, in the Pearl River drainage in Mississippi and Louisiana. SE Fishes
Coun. Proc., 44:6-10.

Warren, M. L., Jr. 1992. Variation of the spotted sunfish, Lepomis punctatus complex
(Centrarchidae): Meristics, morphometries, pigmentation and species limits. Bull.
Alabama Mus. Nat. Hist., 12:1-47.

CTM at: drctmcallister@aol.com

TEXAS J. OF SCI. 61(1):45»60

FEBRUARY, 2009

COMPARISON OF TOTAL LIPID AND
FATTY ACID COMPOSITIONS OF WHOLE-BODY AND
BODY SEGMENTS OF LERTHA EXTENSA ADULTS
(NEUROPTERA: NEMOPTERIDAE)

Ozlem Cakmak, Mehmet Bashan and Ali Satar

Department of Biology, Dicle University
21280 Diyarbakir, Turkey

Abstract. — Total lipid and fatty acid compositions of phospholipids and
triacylglycerols fractions, prepared from whole body and selected body segments
(head, thorax and abdomen) of adults males and females of Lertha extensa Oliver
(Neuroptera: Nemopteridae), were analyzed by gas chromatography and gas
chromatography-mass spectrometry. The female abdomen has the highest level of
total lipid in body segments. Predominant fatty acid components of phospholipid
and triacylglycerol fractions in the whole body and three body segments were C16:0,
C18:ln-9 and C18:2n-6 acids, comprising more than 80 % of the fatty acid
components. Fatty acid profiles of whole body extracts differed by sex only for
phospholipid and triacylglycerol fractions. Fatty acid profiles of phospholipids from
the head and abdomen were similar to those of the whole body, but considerably
different from that of thorax. Several minor fatty acids, comprising < 2% of the
total, were noted. Detailed profiles are reported, and are compared to those of
other insects.

Fatty acids (FA) assume broad biological significance, with
important roles in three general areas. First, lipids serve as energy
reserves, generally stored as saturated and monounsaturated fatty
acids, are deployed in development, hibernation and locomotion.
Second, fatty acids form essential parts of the cellular structure as
components of cellular phospholipids (PL), particularly in biomem¬
branes. Third, certain polyunsaturated fatty acids (PUFAs) -
specifically C20:3n-6 (eicosatrienoic), C20:4n-6 (arachidonic
acid) and C20:5n-3 (eicosapentaenoic acid) - act in regulatory
roles, as substrates for the biosynthesis of prostaglandins and other
eicosanoids. With respect to insects, studies of the biology and
biochemistry of fatty acids have offered new insights, such as the
de novo biosynthesis of Cl 8 and C20 PUFAs (Stanley-Samuelson
etal. 1988).

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

Insects utilize lipids efficiently for development, reproduction
and flight. The amount and composition of lipids in an insect vary
considerably between developmental stages, selected tissues,
nutrition, starvation, sex and hormones (Beenakkers et al. 1985).
Physiological studies in adults address reproduction and
mobilization of lipids during flight, a period of high metabolic
demand (Goldsworthy & Mordue 1989). Lipid and FA
compositions of insects have been studied for whole-body
preparations (Nikolova et al. 2000) and, to a lesser extent,
individual segments of the body.

FA profiles have been documented for a taxonomic array of
insects, but changes in FA composition of individual body
segments have been addressed for comparatively few species
such as Periplaneta americana (Linnaeus) (Blattaria: Blattidae)
(Jurenka et al. 1987); Magicicada septendecium (Linnaeus)
(Hemiptera: Cicadidae) (Hoback et al. 1999); and Photinus pyralis
(Coleoptera: Lampyridae) (Nor Aliza et al. 2000).

The order Neuroptera has received recent interest, due to their
promise as biological control agents and a tolerance for several
modem insecticides (Canard 1998). However, few studies of the
total FA composition of the Neuroptera have been performed
(Fast 1970; Zinkler 1975; Lemesle et al. 1997; Nelson et al.
2003). Larvae of the family Nemopteridae are carnivorous,
although adults are unique for their long, filamentous or ribbon-like
hind wings, and feed exclusively on pollen and nectar with long
specialized mouth parts, whilst all larvae are carnivores (Mansell
1992). Lertha extensa is endemic to Turkey (Satar & Ozbay 2004).

There are no previous descriptions of FA composition of PL and
triacylglycerol (TG) fractions of any neuropteran except Lertha
sheppardi Kirby (Neuroptera: Nemopteridae) (Cakmak et al.
2007). This study provides data concerning fatty acid composition
of PL and TG fractions prepared from whole-body and each of the

CAKMAK, BASHAN & SATAR

threebody segments (head, thorax and abdomen) of adult specimens

of Lertha extensa.

Materials And Methods

Biological specimens -Lertha extensa adults were collected with
nets and light traps from Diyarbakir, Turkey (37° 54’N, 40° 14’E; at
an altitude of about 850 m) throughout June and July 2006.
Samples were individually isolated into plastic boxes of 10 by 5 by
5 cm and supplied with drops of water, commercially available
pollen, and fixed flowers (family Apiaceae = Umbelliferae).

Analysis of insects ' fatty acids -\nsQCts were anesthetized by
chilling on ice, and processed for lipid extraction and analysis
following the methods of Bling & Dyer (1959). For whole-body
insect analyses, three groups of 20 adult males, and 20 adults
females were used per sample in 4 mL of chloroform/methanol
(2:1, v/v). For analyses of isolated body segments, 96 anesthetized
males and females were dissected, and three sets of 32 samples
were collected, then processed for extraction. Each group of
samples was transfered into 3 mL of chloroform/methanol (2:1,
v/v). Autoxidation of unsaturated components was minimized by
adding 50 pi of 2% butylated hydroxytoluene in chloroform to each
sample during the extraction process. Total lipid measures were
conducted using the method of Christie (1982).

Total lipid extracts were dried under a stream of N2, then PL and
TG fractions were isolated by thin-layer chromatography (TLC),
using Silica Gel G TLC plates (20 X 20 cm, 0.25 mm thick). After
applying the total lipid extracts, the TLC plates were developed in
petroleum ether:diethyl ether:acetic acid (80:20:1, v/v). Lipid
fractions were made visible by spraying the TLC plates with 2\T-
dichlorofluorescein (Supelco, Supelco Park, PA, USA), and PL and
TG fractions were identified by corresponding standards.

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

PL and TG fractions were scraped into reaction vials, and the
associated fatty acids were transmethylated by refluxing the
fractions in acidified methanol for 90 min at 85^C. The fatty acid
methyl esters (FAMEs) were extracted from the reaction vials three
times with hexane, and concentrated. Results were expressed as the
percentage of each fatty acid with respect to the total fatty acids.
Internal standard was not used.

Gas chromatography -ThQ FAMEs were analyzed by gas
chromatography using a Ati Unicam 610 gas chromatograph
equipped with a SP-2330 capillary column (30 m by 0.25 mm i.d.,
0.2 pm film thickness, Supelco), a flame ionization detector and an
Unicam 4815 recording integrator. A split injection of 0.5 pi was
used. The temperature condition detector was 250°C. The oven
temperature was set at 180°C for 5 min then reached to 200°C with
a ramp rate of 2 /min, and then held for 15 min. FAMEs were
identified by comparisons of retention times with authentic
standards (Sigma Chemical Co., St. Louis, MO, USA). Individual
FAMEs were identified by comparisons with the chromatographic
behaviors of authentic standards.

Gas chromatography-mass spectrometry -C\\Qm\cdX structures
of fatty acid methyl esters were confirmed by capillary gas
chromatography-mass spectrometry (GC-MS) (HP 5890-E series
GC-System, Hewlett-Packard, Palo Alto, CA, USA) with mass-
selective detection. An Innowax column (30m by 0.25 mm i.d.,
0.25 pm film thickness) was used, and the temperature was
programmed from 150 to 230 ®C at a 2 °C/min increase with an
initial hold of 6 min. The carrier gas was helium (1 mL/min) and
the split ratio was 1:50. The injection port and the detector
temperatures were 250°C and 300°C respectively. The mass
spectrometer was operated in the electron impact ionization mode
(70 eV). Chemical structures of the FAMEs were determined by
comparison of the spectra with the Wiley 275 and Nist 98 databank,
and by comparing obtained spectra with that of authentic standards.

CAKMAK, BASHAN & SATAR

Fig. 1. Total lipid of body segments of Lertha externa as a percentage of total wet
weight.

Statistical analys is. -Stsitistical analysis included ANOVA to
determine variability, F-tests to identify treatment effects, and least
significant difference and student's Mest (Snedecor & Cochran
1967) to identify significant differences between groups (P<0.05).

Results

The percentage of total lipid of body segments of L. extensa
are shown in Fig. 1. The abdomen contain highest level of total
lipid percentage in all body segments. The female abdomen
contain higher total lipid than male {F =656.417; df= 3,8;
P<0.001). No significant difference was observed in total lipid
of head {F =12A1\ P>0.05) and thorax (F =11.247; P>0.05)
between male and female. The fatty acid compositions of PL
and TG prepared from whole-body of adult males and females
L. extensa are displayed in Table 1. Predominant fatty acid
components of whole-body PL and TG were Cl 6:0 (palmitic),
C18:0 (stearic acid), C18:ln-9 (oleic) and C18:2n-6 (linoleic) acids,
occuping over 85% of the fatty acids, in which C18:ln-9 was most
abundant, attaining to ca. 40%. The fatty acid profiles of PL and

THE TEXAS JOURNAE OF SCIENCE, VOL. 61, NO. 1, 2009

Table 1. Proportion of fatty acids, as percentage of total fatty acids, associated with
phospholipids and triacylglycerols prepared from whole-body of Lertha extensa.

Fatty acid

Phospholipids

Triacylglycerols

Female

Male

Female

Male

(means*±5'.Z).)#

(means *±5". D. )#

(means *±5. D. )#

(means*^:^./).)#

C14:0

1.58±0.04a

0.89±0.03b

2.28±0.15a

2.78±0.15a

C15:0

0.51±0.02a

0.32±0.02b

1.71±0.07a

0.24±0.02b

C16:0

11.50±0.70a

lL63±2.01a

26.07±1.23a

28.61±1.52a

C17:0

0.13±0.02a

0.18±0.02a

1.16±0.09a

0.65±0.02b

C18:0

12.70±1.15a

10.87±0.54a

5.32±0.16a

5.77±0.41a

ZSFA

26.42±1.71a

23.86±2.50a

36.54±2.13a

38.05±1.49a

C16:ln-7

6.65±0.90a

5.62±0.53a

7.04±0.07a

6.26±0.08a

C18:ln-9

38.14±0.89a

36.57±1.17a

44.01±2.31a

41.63±1.38b

C20:ln-9

0.20±0.09a

0.26±0.09a

0.10±0.09a

0.14±0.08a

IMUFA

44.99±1.47a

42.45±1.65a

51.16±3.06a

47.89±2.39a

C18:2n-6

22.25±1.03a

23.68±0.19b

11.08±1.07a

8.33±1.09b

C18:3n-3

4.30±0.02a

9.03±0.20b

1.21±0.02a

5.63±0.59b

C20;4n-6

0.65±0.02a

0.51±0.02a

C20:5n-3

0.80±0.02a

0.43±0.02b

ZPUFA

28.00±2.69a

33.65±0.21b

12.29±1.32a

13.96±1.08a

* Averages of three replicates using 20 adult males, and 20 adult females per repli¬
cates.

# Means with the same letter do not significantly different between sexes, P>0.05.
Each section is separately evaluated.

SEA: Saturated Fatty Acids; MUFA: Monounsaturated Fatty Acid; PUFA: Polyun¬
saturated Fatty Acid.

TG prepared from whole-body were essentilally similar
between female and males, although sexual differences in the
proportion were found in some components such as 18:3n-3
(linolenic), the content being higher in males than females, in
both lipid fractions. Among minor components occuping less
than 2% of fatty acids. Cl 5:0 (pentadecanoic acid) and Cl 7:0
(heptadecanoic acid) existed both in PL and TG fractions, but
C20:4n-6 and C20:5n-3 were hardly detected in TG. Odd-
chain fatty acid and eicosanoid precursor C20 PUFAs were
detected by flame ionization gas chromatography only
sporadically; they were in low titres. More sensitive analysis by
gas chromatography-mass spectrometry confirmed that these
components are present in both fractions. Compared with PLs,

CAKMAK, BASHAN & SATAR

the FA profiles of TGs prepared from L. extensa had higher
proportions of C16:0, C18:ln~9 and lower proportions of two
PUFAs, C18:2m6 and C18:3m3.

The fatty acid compositions of PL prepared from the head^
thorax, abdomen were presented in Table 2. The major
components are C16:0, C18:ln-9 and C18:2n»6. The profiles
of these components varied among the body segments which
were analysed. The major component present in lowest
proportion, Cl 6:0, comprised about 25% of phospholipid
fatty acids in head and abdomen, but made up about 16% in
female and 14% in male of thorax. Similarly, C18:2n-6
accounted for about 20% of the phospholipid fatty acids in
the male abdomen, and 41% of the female thorax
phospholipid fatty acids. Although there are sexual
differences in the proportions of several fatty acids of PL
prepared from head, they were more prominent in those
prepared from thorax and abdomen. Fatty acid profile in PL
prepared from head and abdomen was largely similar to that of
whole-body (Table 1), but considerably different from that of
thorax. In head and abdomen, C18:ln~9 was predominant,
attaining to ca. 35% of the fatty acids, and C16:0 and C18:2n“6
respectively occupied 25%, whereas, in thorax, C18:2n-6 was the
most abundant component, occuping 40%, followed by C18:ln-9
(25%) and Cl 6:0 (15%). Some differences among body
segments were detected in the minor components: Cl 7:0 and
C22:ln-9 (docosenoic acid) were specific to abdomen, C20:2n"6
(eicosadienoic acid) not detected in head, while C20:ln-9
(eicosenoic acid), C20:4n”6 and C20:5n-3 were present in all
segments and whole body.

The fatty acid profiles of TG fractions prepared from three
body segments of adult are presented in Table 3. Fatty acid
profiles in TG prepared from head and abdomen were largely
similar to that of whole-body (Table 1), but considerably

Table 2. Proportion of fatty acids, as percentage of total fatty acids, associated with phospholipids prepared from body segments of
Lertha externa.

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

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CAKMAK, BASHAN & SATAR

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* Averages of three replicates using 32 adult males, and 32 adult females per replicates.

# Means with the same letter do not significantly different between sexes, P>0.05. Each section is separately evaluated.

SFA: Saturated Fatty Acids; MUFA: Monounsaturated Fatty Acid; PUFA: Polyunsaturated Fatty Acid

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

different from that of head and abdomen, as in the case of PL
(Table 2). In head and abdomen, C18:ln-9 was predominant,
attaining to ca. 40% of the fatty acids, followed by Cl 6:0 (ca.
30%) and C18:2n-6 (less than 16%), whereas, in thorax, C16:0
was the most abundant component, occuping over 30%, followed
by C18:ln-9 (25%) and C18:2n-6 (23%).

For minor components, some differences were observed
among body segments: Cl 5:0 was present only in abdomen;
Cl 7:0 was not in head; C20:4n-6 and C20:5n-3 was in head
and thorax of males, but not of females, whereas C20:ln-9
present in all segments with both sexes.

Discussion

Changes in lipid stores occur due to flight, mating activities,
and oviposition (Gilby 1965) in the adult stage. The amount of
total lipid was observed to vary among body segments and sexes
of L. extensa. Total lipid amount is significantly highest in
female abdomen. This difference may depend on accumulation
of lipids in the eggs. Abdomen contains the reproduction system,
digestive system, and the most of the fat body. Triglycerides,
which con-stitute 50 to 70% of the fat of insects throughout their
development, are located mostly in their fat bodies and serve as a
source of energy for various metabolic purposes. During ovarial
develop-ment, lipids synthesized in the fat body are transported
to the developing ovary, and stored for use in embryogenesis.

The major fatty acids in the lipids of L. extensa, are Cl 6:0,
Cl 8:0, C18:ln-9 and C18:2n-6, as reported for some other
Neuroptera and most other insect orders (Stanley-Samuelson et
al. 1988; Cakmak 2006; Cakmak et al. 2007). Aside from these
major components, many fatty acids were detected at low
proportions of the PLs and TGs. The additional components
include shorter chain fatty acids, such as C14:0, odd-chain fatty

CAKMAK, BASHAN & SATAR

acids, including C15:0, C17:0 and longer chain saturated and
unsaturated fatty acids, C20:ln-9, C20:2n”6, C20:4n-6, C20:5n“3,
and C22:ln~9,

Fatty acid profiles of PL and TG prepared from whole-
body were essantially similar between female and males,
although sexual differences in the proportion were found
about some components such as C18:3n-3, the content being
higher in males than females, in both lipid fractions. High
levels of C18:3n--3 could be based on a number of
physiological factors. One of the major functions of C18:3n-
3 and PUFAs in general, is that they serve as a structural
component of membranes to maintain proper fluidity and
permeability. C18:3n-3 is also a precurser to the eicosanoids,
including prostaglandins, leuko-trienes, and thromboxanes
(Stanley 2006).

Fatty acid compositions are not fixed in insects and can
change seasonally to perform special functions that may be
critical for survival. Many factors affect the shape of fatty acid
profiles, in particular, development (Cakmak et al. 2007), diet
(Bozkus 2003; Stanley-Samuelson & Dadd 1981; Stanley-
Samuelson et al. 1985), diapause status (Shimizu 1992; Hodkova
et al. 1999; Bashan et al 2002; Bashan & Cakmak 2005), and
body segments (Nor Aliza et al. 2000) exert strong influences on
the shape of fatty acid profiles.

Fatty acid profiles of PL and TG prepared from head and
abdomen was largely similar to that of whole-body, but
considerably different from that of thorax. The principal storage
site for insect lipids is the fat body, and the lipid composition of
the whole insect probably reflects the lipid composition of the fat
body. This organ consist of aggregates of cells forming lobes or
sheets of tissue, which is spread throughout the body and invests
the internal organs. Its spatial arrengement in the abdomen,

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

where large fat deposits are found in close association with the
gut, facilitates the uptake of dietary nutritions. Therefore, it is
not surprising that the fatty acid composition of abdomen and
whole-body were similar but some minor differences were
detected in both fractions, such as Cl 6:0 and C18:3n-3. In
both fractions, proportions of C18:ln-9 were lower in adult in
females than males, which seem most likely to be associated with
the processes of egg development and oviposition. When
neuropterans pupate, larvae dig a small cavity in the soil and spin
a loose silken cocoon around themselves. Many holometabolous
insects exhibit similar behavior, but neuropterans are unusual
because their silk is produced by Malpighian tubules (excretory
organs) and spun from the anus. In contrast, most other
endopterygote insects produce silk in modified salivary or labial
glands and spin it with their mouthparts. Only one other order,
the Coleoptera, makes silk in the same manner as Neuroptera.
Lipids also occur in the silk (Meyer 2005), and it is possible the
fatty acid differences in the abdomen stem from consumption of
some fatty acids while making the cocoon.

PL fraction of thorax of P. americana contained the highest
linoleic acid (38%) while oleic acids (49%) was found primarily
in abdomen (Jurenka et al. 1987). Similarly, high level of
linoleic acid was detected in PL extracted fom thorax of M
septendecim (45%) (Hoback et al. 1999) and Tenebrio molitor
Linnaeus (Coleoptera: Tenebrionidae) (Howard & Stanley-
Samuelson 1990). In both sexes of L. extensa, it was detected
that the ratio of PUFAs in thorax had the highest level due to
linoleic acid in body segments. This result was valid in both
fractions of the insect. In order to fly, insects require flight
muscles that constitute at least 12 to 16% of their total mass, and
flight performance increases as this percentage increases.
However, flight muscles are energetically and materially
expensive to build and maintain, and investment in flight muscles
constrains other aspects of function, particularly female fecundity

CAKMAK, BASHAN & SATAR

(Marden 2000). Fatty acid accumulation in the thorax, where
flight muscles are located, makes it a convenient, ready-made
energy source. Localization of the fatty acid-containing vesicles
in the thorax also supports the possibility that lipid energy is
required for rapid muscle contractions (Georgia & Mohammed
2002). Since insect flight muscles are among the most active in
nature, they have extremely high rates of fiiel supply and flights
of insects are lergely fiieled through fatty acid oxidation. The
lipid substrate is transported as diacylglycerol in the blood,
employing a unique and efficient lipoprotein shuttle system
(Horst et ah 1993). These findings indicate that the accumulation
and utilization of these FAs differ in body segment of both sexes.

Arachidonic acid typically occurs in very small proportions in
PLs of terrestrial insects, ranging from no more than traces to less
than 1% of PL fatty acids, while C20:5n"3 is often missing
entirely from insect lipids. In this study, the proportions of
C20:4n-6 and C20:5n-3 were detected body segment and whole-
body of both fractions (less than 1.8%), especially in PL fraction.
Contrarily, C20:4n-6 made up approximately 21% of the PL fatty
acids prepared from whole-body of males and females of P.
pyralis, and from heads and thoraces prepared from males (Nor
Aliza et al. 2000). Ogg & Stanley-Samuelson (1992) detected
C20:4n“6 approximately 12% of the PL fraction of head of
Manduca sexta (Linnaeus) (Lepidoptera: Sphingidae). These
insects are peculiar among terrestrial insects with respect to
maintaining high proportions of PL C20:4n-6, which might
support an unusually high capacity to generate prostaglandins
and other eicosanoids in these insects.

Acknowledgment

This study was supported by Dicle University Research Fund.
Project number is DUAPK 03 FF- 02.

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 1, 2009

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OC at: ocakmak@dicle.edu.tr

TEXAS J. SCI 61(1), FEBRUARY, 2009

GENERAL NOTES

mXIDIUMSEROTINUM(mOTlSTA: MYXOZOA)
FROM A JEFFERSON SALAMANDER
{AMBYSTOMA JEFFERSONIANUM), IN ILLINOIS

Chris T. McAllister^ John A. Crawford and Andrew R. Kuhns

RapidWrite, 102 Brown Street
Hot Springs National Park, Arkansas 71913
Indiana School of Medicine-Terre Haute, 135 Holmstedt Hall
Terre Haute, Indiana 47809-9989 and
Illinois Natural History Survey, 1816 S. Oak Street
Champaign, Illinois 61820

The Jefferson salamander, Ambystoma jeffersonianum (Green,
1827) ranges from southeastern New York through Pennsylvania
and eastern and southern Ohio to southern Indiana, and
southward to southcentral Kentucky and northern Virginia
(Conant & Collins 1998). However, because of extensive
hybridization with the blue-spotted salamander {Ambystoma
laterale), the precise range of novel populations is uncertain
(Bogart & Klemens 1997), In Illinois, the species is listed as
imperiled (S2) by the Nature Conservancy (NatureServe 2008)
and has been reported from only two counties of the state (Clark
and Edgar) (Brodman 2005). As such, A. jeffersonianum is listed
as a threatened species in Illinois (Illinois Endangered Species
Protection Board 2006).

Little is known about the endoparasites of A. jeffersonianum.
Rankin (1945) reported the trematode, Brachycoelium
salamandrae in A. jeffersonianum from Massachusetts, Fischthal
(1955) noted unidentified immature trematodes and
metacercariae from the species in New York, and Anderson
(1960) documented the nematode, Cosmocercoides dukae in this
salamander from Ontario, Canada. Herein is provided new host
and geographic distribution records for an endoparasite of A.
jeffersonianum from Illinois,

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

An adult female (snout-vent length = 67 mm) A.

jeffersonianum was collected on 2 April 2008 by minnow trap
from Zone 16 in Clark County, Illinois (39.1948°N, 87.4724° W).
It was returned to the laboratory and euthanized with a dilute
chloretone solution. The gall bladder was removed and
processed for myxozoans according to methods of McAllister &
Trauth (1995). The voucher specimen is deposited in the
Arkansas State University Herpetological Museum (ASUMZ),
State University, Arkansas as ASUMZ 3 1 144. A parasite
voucher was deposited in the United States National Parasite
Collection (USNPC), Beltsville, Maryland as USNPC 100988.

This salamander was found to be infected with a myxozoan
fitting the description of Myxidium serotinum by Kudo &
Sprague (1940). Numerous free spores and spherical to ovoidal
trophozoites were found in bile contents. Ovoidal bivalved
spores containing two polar capsules were observed as well as
finer details of the spore shell that are typical of M serotinum (cf.
McAllister et al. 1995).

This parasite has been previously reported from a variety of
amphibians, including at least 16 species of frogs and toads
within four families and four species of salamanders within two
families (McAllister & Trauth 1995; McAllister et al. 2008).
Salamander hosts include the spotted salamander (Ambystoma
maculatum) from Arkansas and Texas (McAllister et al. 2008),
marbled salamander {Ambystoma opacum) from Arkansas
(McAllister & Trauth 1995), small-mouthed salamander
{Ambystoma texanum) from Texas (McAllister & Upton 1987),
and two-lined salamander {Eurycea bislineata) from West
Virginia (Clark & Shoemaker 1973). A similar species,
Myxidium melleni Jirku, Bolek, Whipps, Janovy, Kent & Modry,
2006 was described from western chorus frogs {Pseudacris
triseriata) and Blanchard’s cricket frogs {Acris blanchardi) from
Nebraska (Jirku et al. 2006). More recently, M melleni was

TEXAS J, SCI. 61(1), FEBRUARY, 2009

reported in the Cajun chorus frog, Pseudacris fouquettei
{^Pseudacris feriarum) from Texas by McAllister et ah (2008).

In summary, this study provides the first report of M
serotinum in a threatened salamander species (A. jeffersonianum)
from Illinois. For comparative purposes, it is suggested that
additional salamanders, including related A. laterale from Illinois
be examined for endoparasites. Like A. jeffersonianum, A.
laterale has already been reported to harbor some of the same
parasites including unidentified metacercariae (Muzzall &
Schinderle 1992) and B. salamandrae (Muzzall & Schinderle
1992) in Michigan, and C dukae (Coggins & Sajdak 1982) in
Wisconsin, and may share others, including M serotinum.

Acknowledgments

We thank the Illinois Department of Natural Resources for
providing funding from contributions to the Illinois Wildlife
Preservation Fund. The specimen was collected under an Illinois
Threatened and Endangered Species Permit (#05-118) issued to
ARK, We also thank P. A. Pilitt (USNPC) and S. E. Trauth
(ASUMZ) for curatorial assistance and C. R. Bursey (Penn.
State- Shenango) for information on salamander parasites.

Literature Cited

Anderson, R. C. 1960. On the development and transmission of Cosmocercoides
dukae of terrestrial molluscs in Ontario. Can. J. ZooL, 38:801-825.

Bogart, J. P. & M. W. Klemens. 1997. Hybrids and genetic interactions of mole
salamanders (Ambysioma jeffersonianum and A. laterale) (Amphibia:
Caudata) in New York and New England. Amer. Mus. Nov. No. 3218, Amer.
Mus. Nat Hist., New York. 78 pp.

Brodman, R, 2005. Ambystoma jeffersonianum (Green, 1827) Jefferson
salamander. Pp. 611-613, in Amphibian declines: The conservation status of
United States species (M. Lannoo, ed.), Univ. California Press, xxi + 1-1094.
Clark, J. G. & J. P. Shoemaker. 1973. Eurycea bislineata (Green), the two-lined
salamander, a new host of Myxidium serotinum Kudo & Sprague, 1940
(Myxosporida, Myxidiidae). J. ProtozooL, 20:365-366.

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

Coggins, J. R. & R. A. Sajdak. 1982. A survey of helminth parasites in the
salamanders and certain anurans from Wisconsin. Proc. HelminthoL Soc,
Washington, 49:99-102.

Conant, R. & J. T. Collins. 1998. A Field Guide to Reptiles and Amphibians of
Eastern and Central North America, ed. (expanded). Houghton Mifflin,
Boston. 616 pp.

Fischthal, J. H. 1955. Ecology of worm parasites in south-central New York.
Amer. Midi. Nat, 53:176-183.

Illinois Endangered Species Protection Board, 2006. Endangered and Threatened
Species List. One Natural Resources Way. Springfield, Illinois.
http://dnr.state.iLus/espb/datelist.htm

Jirku, M., M. G. Bolek, C. M. Whipps, J. Janovy, Jr., M. L. Kent & D. Modry.
2006. A new species of Myxidium (Myxosporea: Myxidiidae), from the
western chorus frog, Pseudacris triseriata triseriata, and Blanchard’s cricket
frog, Acris crepitans bl anchor di (Hylidae), from eastern Nebraska:
Morphology, phylogeny, and critical comments on amphibian Myxidium
taxonomy. J. ParasitoL, 92:611-619.

Kudo, R. & V. Sprague. 1940. On Myxidium immersum (Lutz) and M. serotinum
n. sp., two myxosporidian parasites of Salientia of South and North America.
Rev. Med. Trop. ParasitoL Bacteriol. Clin. Lab., 6:65-73.

McAllister, C. T. & S. E. Trauth. 1995. New host records for Myxidium
serotinum (Protozoa: Myxosporea) from North American amphibians. J.
ParasitoL, 81:485-488.

McAllister, C. T. & S. J. Upton. 1987. Endoparasites of the smallmouth
salamander, Amby stoma texanum (Caudata: Ambystomatidae) from Dallas
County, Texas. Proc. HelminthoL Soc. Washington, 54:258-261.

McAllister, C. T., C. R. Bursey & S. E. Trauth. 2008. New host and geographic
distribution records for some endoparasites (Myxosporea, Trematoda,
Cestoidea, Nematoda) of amphibians and reptiles from Arkansas and Texas,
U.S.A. Comp. ParasitoL, 75:241-254.

McAllister, C. T., S. E. Trauth & B. L. J. Delvinquier. 1995. Ultrastmctural
observations on Myxidium serotinum (Protozoa: Myxosporea) from Bufo
speciosus (Anura: Bufonidae), in Texas. J. HelminthoL Soc. Washington,
62:229-232.

Muzzall, P. M. & D. B. Schinderle. 1992. Helminths of the salamanders
Ambystoma t. tigrinum and Ambystoma laterale (Caudata: Ambystomatidae)
from southern Michigan. J. HelminthoL Soc, Washington, 59:201-205.

Rankin, J. S., Jr. 1945. An ecological study of the helminth parasites of
amphibians and reptiles of western Massachusetts and vicinity. J. ParasitoL,
31:142-150.

CTM at: drctmcallister@aoLcom

TEXAS T SCI. 61(1), FEBRUARY, 2009

RECORDS OF THE PORCUPINE {ERETHIZON DORSATUM) FROM
THE EASTERN MARGINS OF THE EDWARDS PLATEAU OF TEXAS

Amy B. Baird*^ Gregory B* Pauly^ David W* Hall
and Travis J. LaDuc

Section of Integrative Biology and Texas Natural Science Center
The University of Texas, Austin, Texas 78712
^Current address:

National Museum of Natural History, P.O. Box 9517,

2300RA Leiden, The Netherlands

During the first half of the 1900s, the porcupine {Erethizon
dorsatum) in Texas was restricted to the northernmost Panhandle
and parts of the Trans-Pecos (Bailey 1905; Taylor & Davis 1947;
Hall & Kelson 1959), By the mid- 1900s, observations suggested
that the range was expanding into the southern Panhandle, and onto
the western Edwards Plateau (Milstead & Tinkle 1958).
Subsequent accumulations of voucher specimens over the past
several decades have served to both substantiate these earlier
observations and to document the expansion of the porcupine
eastward across the southern Rolling Plains (e.g. Tyler & Joles
1997 and Caire et ah 1989 for Oklahoma; Dalquest & Homer 1984
for north-central Texas) and elsewhere across Texas (e.g., Davis
1974; Davis & Schmidly 1994; Use & Hellgren 2001). The species
has presently come to occupy most of the western two-thirds of the
state (Stangl et al. 1991; Schmidly 2004),

Goetze (1998) documented the occurrence of the porcupine over
the western two-thirds of the Edwards Plateau, and the two sight
records reported by Milstead & Tinkle (1958) from Mason and Ken-
counties remain the southeastemmost records from both the region
and the state. This study reports on two specimens that extend the
known range of the species to the eastern margin of the Edwards
Plateau, a distance of approximately 160 km from the nearest
voucher records of Kimble and Concho counties reported by Goetze
(1998). Both were salvaged as road-killed animals, and the skulls
were retained as vouchers for deposit with the Natural Science

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

Research Laboratories of Texas Tech University (TTU). The first
specimen (TTU 108417) was taken on 4 January 2003 from Travis
County (0.4 mi NE of Hwy. 71 on Southwest Parkway; 30° 16.777’
N, 97° 54.498’ W), and the second (TTU 108416) was taken on 20
January 2007 from Burnet County (1.5 mi E of Llano-Bumet
County line on Hwy 71; 30° 30.903’ N, 98° 19.431’ W). Two
sightings from as recently as July 2008 on the Brackenridge Field
Laboratory, University of Texas campus in Austin, have also been
related to the authors by lab personnel as the first evidence of the
species locally since the lab’s inception in 1967.

The reported vouchers and recent sight records provide evidence
that the porcupine is a viable resident component of the Edwards
Plateau. Given the demonstrated ability of E. dorsatum to expand
its range in recent decades across the relatively open terrain
dominating much of the northwestern parts of Texas, there seems
little to impede its eastward progress beyond the Balconian
Escarpment, and across the blackland prairies and postoak belt.

We thank David M. Hillis of the University of Texas for
assistance in the collection of the Burnet County specimen, and
Larry Gilbert, Director of UT Brackenridge Field Laboratory, for
providing information and documentation on the porcupine
observed there. Specimens were collected under Texas Parks and
Wildlife Scientific Permit (SPR-02010-133).

Literature Cited

Bailey, V. 1905. Biological survey of Texas. North American Fauna 25:1-222.
Caire, W., J. D. Tyler, B. P. Glass, & M. A. Mares. 1989. Mammals of Oklahoma.

University of Oklahoma Press, Norman, xiii + 567 pp.

Dalquest, W. W., & N. V. Homer. 1984. Mammals of North-Central Texas.

Midwestern State University Press, Wichita Falls, Texas, 254 pp.

Davis, W. B. 1974. The Mammals of Texas. Texas Parks & Wildlife Press, Austin,
294 pp.

Davis, W. B., & D. J. Schmidly. 1994. The Mammals of Texas. Texas Parks &
Wildlife Press, Austin, 338 pp.

Goetze, J. R. 1998. The mammals of the Edwards Plateau, Texas. Special
Publications, Museum of Texas Tech University, 41:1-263.

TEXAS J. SCI. 61(1), FEBRUARY, 2009

Hall, E. R., & K. R. Kelson. 1959. The Mammals of North America. Ronald Press,
New York, 2:viii + 547-1083,

Use, L. M, and E. C. Hellgren. 2001. Demographic and Behavioral Characteristics
of North American Porcupines (Erethizon dorsatum) in Pinyon-Juniper
Woodlands of Texas. American Midland Naturalist, 146(l):329-338.

Milstead, W. W. and D. W. Tinkle. 1958. Notes on the porcupine {Erethizon
dorsatum) in Texas. Southwestern Naturalist, 3(l-4):236-237.

Schmidly, D. J. 2004. The Mammals of Texas, revised edition. University of Texas
Press, Austin, 501 pp.

Stangl, Jr., F. B., R. D. Owen, and D. E. Morris-Fuller. 1991. Cranial variation and
asymmetry in southern populations of the porcupine, Erethizon dorsatum. Texas
Journal of Science, 43 (3): 237-259.

Taylor, W. P., & W. B. Davis. 1947. The mammals of Texas. Texas Game, Fish,
and Oyster Commission, Austin, Bull. 27:1-79

Tyler, J. D. & S. Joles. 1997. The Porcupine in Oklahoma. Proceedings, Oklahoma
Academy of Science, 77(1): 107-1 10.

ABB at: baird(§nnm.nl

FIRST RECORD OF CYMBOVULA ACICULARIS
(GASTROPODA: PROSOBRANCHIA: OVULIDAE) FROM
THE COAST OF TAMAULIPAS, MEXICO

Alfonso Correa-Sandoval and Ned E, Strenth

Laboratorio de Zoologia, Instituto Tecnologico de Cd. Victoria,

A.P. 175, C.P. 87010, Cd. Victoria, Tamaulipas, Mexico (ACS) and
Department of Biology, Angelo State University,

San Angelo, Texas, 76909

Members of the family Ovulidae are found in warm marine
waters as ectoparasites on sea whips and sea fans upon which they
feed (Cate 1973; Rosenberg 1989; Redfem 2001). They are largely
restricted to the microhabitat provided by their association with the
various host species (Abbott & Morris 1995). The shell may be
oblong, ftisiform or elongate. The surface is smooth with the
aperture extending the length of the shell and it lacks an operculum
(Abbott 1974; 1986). Notable among living specimens is the
expanded fleshy mantle which covers the shell and exhibits a

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

variety of vivid coloration patterns (Keen 1971; Rehder 1981).
Species of Ovulidae are known from the American Atlantic coast
and exhibit distributions within the Virginian subprovince and
Carolinian and Caribbean provinces (Abbott 1974; 1986; Andrews
1992). Five species, Cyphoma gibbosum (Linnaeus), C macgintyi
Pilsbry, C allenae (Cate), C intermedium (Sowerby) and
Simnialena uniplicata (Sowerby), are currently known from the
state of Tamaulipas in Mexico and have been reported from the
northern and southern coastal zones (Leal 1978; Fregoso 1986;
Perez 1993).

Cymbovula acicularis (Lamarck), has been reported from North
Carolina to Brazil, Bermuda and the West Indies (Morris 1975,
Warmke & Abbott 1975; Abbott & Morris 1995; Abbott & Dance
1998). Fossil specimens are known from the Pleistocene of South
Carolina (Richards 1962). This species is currently known from the
western Gulf of Mexico from Texas (Tunnell & Chaney 1970; Ode
1973) south to Veracruz, Yucatan (Vokes & Vokes 1983; Garcia-
Cubas & Reguero 2004) and Quintana Roo (Oliva-Rivera &
Navarrete 2007). This species, however, has not been previously
recorded by prior surveys from the state of Tamaulipas (Fregoso
1986; Perez 1993; Perez-Rodriguez 1997; Rodriguez-Castro et al
2005). This study reports the collection of Cymbovula acicularis
from near the village of La Pesca on Barra Soto la Marina
(23°42’06'’N and 97°45'00”W) which is located along the central
Tamaulipas coast 51 km east of Soto la Marina. This area is
approximately 260 km south of the Texas coast, less than 50 km
north of the Tropic of Cancer and exhibits a warm-subhumid
climate (INEGI 1981). The area exhibits an isotherm with an
average annual temperature of 25° C and average salinity of 33.9
ppt (Rodriguez-Castro 2002).

Two specimens of C acicularis were collected during a field
trip to the La Pesca area on 9 May 2007. The gastropods were
found on specimens of yellow sea whips {Leptogorgia sp.:
Gorgoniidae) upon which they are known to both live and feed

TEXAS J. SCI. 61(1), FEBRUARY, 2009

Figure 1. Cymbovula aciciilaris from La Pesca, Tamaulipas, Mexico. Scale = 1 mm.

(Rehder 1981; Britton & Morton 1989). The specimens are
deposited in the Malacological Collection of the Institute
Tecnologico de Ciudad Victoria (ITCVZ 5021). A second trip to
the same site on 6 November, 2007 resulted in the collection of two
additional specimens which are also deposited with the Institute
Tecnologico de Cd. Victoria collections (ITCVZ 5118).

The four specimens from La Pesca are yellow, elongated, thin
and measure 8-17 mm in length (Figure 1). The aperture extends
along the total length of the shell; the lip is thin and sharp.
Columella with an opaque anterior thickening and with
punctuations. Columellar area flattned. Surface brilliant, with
spirals growth lines, more or less equidistant, slightly irregular
towards the posterior area of the shell. The morphology of these
specimens agrees with the descriptions of Redfem (2001) and
Garcia-Cubas & Reguero (2004).

This record of C. acicularis from La Pesca brings the total
number of marine gastropods reported from the state of Tamaulipas
to 165. This is second only to Yucatan in the total number of
reported species among those states in Mexico which border the
Gulf of Mexico.

70 THE TEXAS JOURNAL OF SCIENCE-VOL. 6 1 , NO. 1 , 2009

Acknowledgments

We wish to thank Fred G. Thompson for both his response and
assistance to our request during this study for an examination of
the Collection of Mollusks of the Florida Museum of Natural
History, University of Florida, in Gainesville, Florida. Special
appreciation is extended to Dr. Martha Reguero of the
Universidad Nacional Autonoma de Mexico and Dr. Fabio
Moretzsohn of the Harte Research Institute at TAMU-Corpus
Christi for suggestions that greatly improved this manuscript.
Also to Anabel Gutierrez for her assistance in the preparation of
the manuscript and to Gonzalo Guevara for the preparation of
figure 1.

Resumen.-El gastropodo marino Cymbovula acicularis es
registrado por primera vez para la malacofauna de Tamaulipas,
Mexico.

Literature Cited

Abbott, R. T. 1974. American Seashells. Van Nostrand Reinhold. New York,
663 pp.

Abbott, R. T. 1986. Seashells of North America. A Guide to Field
Identification. St. Martin’s Press. New York, 280 pp.

Abbott, R. T. & P. A. Morris. 1995. A Field Guide to Shells of the Atlantic and
Gulf Coast and the West Indies. Houghton Mifflin. New York, 350 pp.

Abbott, R. T. & S. P. Dance. 1998. Compendium of Seashells. Odyssey
Publishing. China, 41 1 pp.

Andrews,!. 1992. A Field Guide to Shells of the Texas Coast. Gulf Publishing
Company. Houston, Texas, 176 pp.

Britton, J. C. & B. Morton. 1989. Shore Ecology of the Gulf of Mexico.
University of Texas Press. Austin, Texas, 387 pp.

Cate, C. N. 1973. A Systematic Revision of the Recent Cypraeid Family
Ovulidae (Mollusca, Gastropoda). The Veliger 15 (Supplement): 1-1 16.

Fregoso, J. A. 1986. Contribucion al conocimiento de la fauna malacoldgica de
la costa del Estado de Tamaulipas, Mexico. Tesis Licenciatura. Escuela de
Ciencias Bioldgicas. Universidad del Noreste, Tampico, Tamaulipas, 133 pp.

Garcia-Cubas, A. & M. Reguero. 2004. Catalogo ilustrado de moluscos
gasteropodos del Golfo de Mexico y Mar Caribe. Instituto de Ciencias del

TEXAS J. SCL 61(1), FEBRUARY, 2009

Mar y Limnologia. Universidad Nacional Autonoma de Mexico. Mexico, D.
F., 171 pp.

INEGI. 1981. Instituto Nacional de Estadistica, Geografia e Informatica. Carta
de Climas. Esc. 1:1000,000. Secretaria de Programacion y Presupuesto.
Mexico, D.F.

Keen, A. M. 1971. Sea Shells of Tropical West America. Stanford University
Press. Stanford, California, 1064 pp.

Leal, L. 1978. Estudio taxonomico de los moluscos marines (Gastropoda-
Pelecypoda) representtativos de la region de Soto la Marina, Tamaulipas,
Mexico. Tesis Licenciatura. Facultad de Ciencias Biologicas. Universidad
Autonoma de Nuevo Leon. San Nicolas de Los Garza, Nuevo Leon, 107 pp,

Morris, P. A. 1975. A Field Guide to Shells of the Atlantic and Gulf Coast and
the West Indies. Houghton Mifflin. Boston, Massachusetts, 330 pp.

Oliva-Rivera, J. J. & A. de Navarrete. 2007. Larvas de moluscos gasteropodos
del Sur de Quintana Roo, Mexico. Hidrobiologica 17(2):151-158.

Ode, H. 1973. A survey of the molluscan fauna of the northwest Gulf of Mexico
- preliminary report (continued). Texas Conchologist 9: 60-72.

Perez, F. A. 1993. Contribucidn al conocimiento de los gasteropodos (Mollusca:
Gastropoda) de la costa de Tamaulipas, Mexico. Tesis Licenciatura. Escuela
de Biologia. Universidad del Noreste. Tampico, Tamaulipas, 144 pp.

Perez-Rodriguez, R. 1997. Moluscos de la plataforma continental del Atlantico
Mexicano. Universidad Autonoma Metropolitana-Xochimilco. Mexico, D.
F., 260 pp.

Redfem, C. 2001. Bahamian Seashells. A Thousand Species from Abaco,
Bahamas. Bahamianseashells.com. Boca Raton, Florida, 280 pp.

Rehder, H. A. 1981. National Audubon Society Field Guide to North American
Seashells. Alfred A. Knopf New York, 894 pp.

Richards, G. H. 1962. Studies on the Marine Pleistocene: The Marine
Pleistocene of the Americas and Europe; The Marine Pleistocene Mollusks of
Eastern North America. Trans. Am. Phil. Soc., New Series 52(3): 1-141.

Rodriguez-Castro, J. H. 2002. Sistematica y zoogeografia de los gastropodos y
bivalvos marinos de la costa del Estado de Tamaulipas, Mexico. Tesis
Maestria en Ciencias. Instituto Tecnologico de Cd. Victoria. Cd. Victoria,
Tamaulipas, 248 pp.

Rodriguez-Castro, J. H., A. Correa-Sandoval & N. Strenth. 2005. Gastropodos
marinos de Tamaulipas: Pp. 88-96, in: Biodiversidad Tamaulipeca Vol. 1. L.
Barrientos, A. Correa-Sandoval, J. V. Horta and J. Garcia (Editors). Instituto
Tecnologico de Cd. Victoria. Cd. Victoria, Tamaulipas, 272 pp.

Rosenberg, G. 1989. Aposematism evolves by individual selection: evidence
from marine gastropods with pelagic larvae. Evolution, 43(8): 1811-1813.

Tunnell, J. W., Jr. & A. H. Chaney. 1970. A checklist of mollusks of Seven and
One-Half Fathom Reef, northwest Gulf of Mexico. Contributions in Marine
Science, 15: 193-203.

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 1, 2009

Yokes, H. E. & E. H. Yokes. 1983. Distribution of Shallow-water Marine
Mollusca, Yucatan Peninsula, Mexico. Middle American Research Institute,
Publication 54. Tulane University, New Orleans, 183 pp.

Warmke, G. L. & R. T. Abbott. 1962. Caribbean Seashells. Dover Publications.
New York, 348 pp.

AC-S at: agutierr@uat.edu.mx

THE TEXAS JOURNAL OF SCIENCE— VOL. 61, NO. 1, 2009

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Jones, T. L. 1971. Vegetational patterns in the Guadalupe Mountains,
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Smith, J. D. 1973. Geographic variation in the Seminole bat, Lasiurus
seminolus. J. Mammal., 54(l):25-38.

Smith, J. D. & G. L. Davis. 1985. Bats of the Yucatan Peninsula. Occas.
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THE TEXAS JOURNAL OF SCIENCE— VOL. 61, NO. 1, 2009

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Jones, T. L. 1975. An introduction to the study of plants. John Wiley &
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265, in Native grasses of North America (R. R. Dunn, ed.), Univ. Texas
Studies, 205 :xx+ 1-630.

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Davis, G. L. 1975. The mammals of the Mexican state of Yucatan.

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THE TEXAS JOURNAL OF SCIENCE

Volume 61, No. 2 May, 2009

CONTENTS

Vegetation of South Padre Island, Texas: Freshwater and Brackish Wetlands.

By Frank W. Judd and Robert 1. Lonard. . . . . . . . 83

Seasonal Trophic Ecology of the White- Ankled Mouse, Peromyscus pectoralis
(Rodentia: Muridae) in Central Texas.

By John T. Baccus, John M. Hardwick, David G. Huffman

and Mark A. Kainer . . . . . 97

Changes In Vegetation Patterns and their Effect on Texas Kangaroo Rats
(Dipodomys el at or).

By Allan D. Nelson, Jim R. Goetze, Elizabeth Watson

and Mark Nelson . . 119

Breeding Biology of the Bam Swallow (Hirundo rustic a) in Northeast Texas
with Temporal and Geographic Comparisons to other North American Studies.

By K. T. Turner and J. G. Kopachena . . . . 131

General Notes

Reproductive Cycle of the Central American Mabuya,

Mabiiya unimarginata (Squamata: Scincidae) from Costa Rica.

By Stephen R. Goldberg . . . . . 147

New Geographic Distribution Records for Parajulid Millipeds
(Diplopoda: Julida), in Arkansas and Texas.

By Chris T. McAllister and Henry W. Robison . . . . 151

Noteworthy Records of Dragonflies (Odonata: Anisoptera) From
Jones and Taylor counties of Central Texas.

By Thomas E. Lee, Jr., Amisha J. Patel, Benjamin W. Johnson
and Roy C. Vogtsberger

157

THE TEXAS JOURNAL OF SCIENCE
EDITORIAL STAFF

Managing Editor:

Ned E. Strenth, Angelo State University
Manuscript Editor:

Frederick B. Stangl, Jr., Midwestern State University
Associate Editors:

Allan D. Nelson, Tarleton State University
Jim R. Goetze, Laredo Community College
Associate Editor for Botany:

Janis K. Bush, The University of Texas at San Antonio
Associate Editor for Chemistry:

John R. Villarreal, The University of Texas-Pan American
Associate Editor for Computer Science:

Nelson Passos, Midwestern State University
Associate Editor for Geology:

Ernest L. Lundelius, University of Texas at Austin
Associate Editor for Mathematics and Statistics:

E. Donice McCune, Stephen F. Austin State University

Manuscripts intended for publication in the Journal should be submitted in
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Stephenville, Texas 76402
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AFFILIATED ORGANIZATIONS
American Association for the Advancement of Science,

TEXAS J. OF SCI. 61(2):83-96

MAY, 2009

VEGETATION OF SOUTH PADRE ISLAND, TEXAS:
FRESHWATER AND BRACKISH WETLANDS

Frank W. Judd and Robert 1. Lonard

Department of Biology, University of Texas- Pan American
Edinburg, Texas 78541

Abstract -Species composition and importance, species richness, species diver¬
sity, and evenness were compared among 14 freshwater and brackish wetlands in the
secondary dunes and vegetated flats topographic zone of South Padre Island, Texas.
Twenty-four different species were found among the sites, but the greatest species
richness at a site was only 14. Mean species diversity was 0.855 and mean evenness
was 0.797. Both parameters showed little variability among sites. South Padre
Island wetlands had significantly lower species richness and species diversity than
freshwater and brackish marshes of the adjacent Texas mainland. Conversely,
evenness was similar among island and mainland wetlands. Most of the island
wetlands shared 62% to 67% of their species and similarity in species composition
was significantly greater for island wetlands than mainland wetlands. Bulrush
(Schoenoplectus pungens) was the dominant species at nine wetlands. Umbrella
grass (Fiiirena simplex) and marshhay cordgrass {Spartina patens) each were
dominant in two wetlands. Bushy bluestem {Andropogon glomeratus) was dominant
in one wetland. Bie meager data currently available suggests that species richness of
wetland communities is similar on northern and southern portions of Padre Island, but
similarity in species composition is modest. Species importance in wetland
communities varies markedly between northern and southern areas of Padre Island.

The most conspicuous physiographic feature along the Gulf of
Mexico coastline of Texas is a series of five barrier islands that
enclose several shallow bays. The flora and vegetation of South
Padre Island are relatively well known compared to other areas of
the barrier island chain (Dahl et al. 1974; Judd et al. 1977; Lonard
et al. 1978; Lonard & Judd 1980; Lonard & Judd 1981; Judd &
Sides 1983; Judd & Lonard 1985; Judd & Lonard 1987; Judd et al.
1989; Lonard & Judd 1989; Judd et al. 1991; Lonard et al. 1991;
Everitt et al. 1991; Everitt et al. 1992; Lonard & Judd 1993; Judd et
al. 1994; Lonard & Judd 1997; Judd et al. 1998; Lonard & Judd
1999; Everitt et al. 1999; Lonard et al. 1999; Judd et al. 2007) but
little is known of the composition of freshwater and brackish
wetland communities on South Padre Island or any of the other
Texas barrier islands.

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

Judd et al. (1977) reported that dense meadows of fresh water
marsh communities occurred within the secondary dunes and
vegetated flats topographic zone of South Padre Island and
identified five common species. Similarly, Lonard et al. (1999)
commented that depressions that often support development of
marsh communities occur frequently in the secondary dunes and
vegetated flats topographic zone. These wetland plant communities
were said to be dominated by a combination of sedges and grasses
including Scirpus pungens (= Schoenoplectus pungens), Spartina
patens^ Fimbristylis castanea, Fuirena simplex and Rhynchospora
color ata by Lonard et al. (1999). Thus, the only information
available is a partial list of species that occur in the wetlands of
South Padre Island. A complete list of species for even one wetland
is lacking and there is no information on the abundance of the
species comprising a wetland community or the fidelity of species
among wetland sites. Consequently, a quantitative comparison of
wetland communities at 14 sites on South Padre Island is provided
herein to rectify these insufficiencies in the knowledge of the
vegetation of South Padre Island.

Materials and Methods

The locations of wetland communities studied are given in Table
1. Four of the 14 sites contained standing water at the time of
sampling. Salinity was recorded at each of the four sites using a
temperature compensated hand-held refractometer (Table 1).
Marshes were considered freshwater if salinity was 0.0 to 0.5 ppt
and brackish if salinity was 0.5 to 17.0 ppt .

The line intercept method (Canfield 1941) was used to quantify
species abundance. Seventy meters of transects were sampled at
each site. Each transect was divided into 10 m intervals and
readings were taken along the length of each interval. First, the
total cover of dead vegetation intercepted was recorded without
regard to species. For live vegetation, each species intercepted by
the line was rated individually without separation into strata.
Species and foliage cover were recorded and from these data the

JUDD «& LONARD

Table 1. Study site locations, dates of sampling, and mean salinity of sites with standing
water.

Site Location Sample Date Mean Salinity (ppt)

26°09’43.26”

'N,

97°10’27.49”

' W

5-21-07

dry

26°10’13.16”

'N,

97°10’32.18”

5-21-07

dry

26°13’00.52”

'N,

97°10’56.16”

6-14-07

dry

26°13’04.89”

97°10’57.26”

6-14-07

dry

26°13’25.55”

97°10’57.67”

5-21-07

2.0

26°14’22.16”

97°11’ 10.28”

8-4-07

0.0

26°14’28.30”

97°11’09.16”

9-24-07

5.5

26°14’47.10”

97°11’14.44”

9-24-07

dry

26°17^03.69”

97°11 ’45.26”

11-5-07

dry

26° 17’ 5 1.20”

97°11’55.86”

11-5-07

dry

26°19’ 12.36”

97°12’1L11”

3-26-08

diy

26°19’29.37”

97°12’2L68”

3-26-08

12.0

26°20’44.42”

97°12’36.77”

3-27-08

dry

26°21’33.38”

97°12’48.32”

3-27-08

dry

frequency of occurrence, relative frequency, relative cover and an
importance value, which is the sum of relative frequency and
relative cover, were calculated. The importance value was used to
determine dominant species.

Similarity of species composition among wetland sites was
calculated using Sorensen’s Coefficient of Community (Krebs
1999). Species importance value was used as the measure of
abundance for calculating species diversity indices. Species
diversity was assessed using the Shannon diversity index (Brower
et al. 1998; Krebs 1999). Evenness was determined as the ratio of
heterogeneity (H’) to maximum heterogeneity (H’ max) (Brower et
al. 1998; Krebs 1999). Nomenclature and common names follow
Jones &Wipff (2003).

Results

Study sites were located in the southern two-thirds of South
Padre Island along a south to north axis in the secondary dune and
vegetated flats topographic zone (Judd et al. 1977). Distance
between successive study locations ranged from 120 m to 5,010 m
(mean = 1,846.6 m, SD = 1,764.9 m). Twenty-four different
species were found in the 14 sites sampled (Table 2). Twenty of

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

Table 2. Species present, their periodicity, growth form and percent occurrence among
sites.

Species

Periodicity

Growth Form

Occurrence

Andropogon glomeratiis

Perennial

Grass

78.6

Eragrostis secimdiJJora

Perennial

Grass

7.1

Panicum amariim

Perennial

Grass

57.1

Paspalum monostachyum

Perennial

Grass

64.3

Schizachyriwn littorale

Perennial

Grass

50.0

Spartina patem

Perennial

Grass

71.4

Sporobolus airoides

Perennial

Grass

14.3

Sporohohis virginicus

Perennial

Grass

71.4

Eleocharis geniculata

Annual

Sedge

42.9

Fimbristylis castanea

Perennial

Sedge

21.4

Fuirena simplex

Perennial

Sedge

85.7

Rhynchospora colorata

Perennial

Sedge

57.1

Schoenoplectus pungens

Perennial

Sedge

92.9

Typha domingensis

Perennial

Cattail

14.3

Agalinus sp.

Annual

Broad-leaved herbaceous

28.6

Bacopa monnieri

Perennial

Broad-leaved herbaceous

21.4

Blutaparon vermiciilare

Perennial

Broad-leaved herbaceous

7.1

Borrichia frutescens

Perennial

Broad-leaved suffrutescent

35.7

Conoclinium betonicifolium

Perennial

Broad-leaved herbaceous

92.9

Flaveria brownii

Annual

Broad-leaved herbaceous

7.1

Iva texensis

Annual

Broad-leaved herbaceous

100.0

Lythnim alatum

Perennial

Broad-leaved herbaceous

21.4

Samolus ebracteatus

Perennial

Broad-leaved herbaceous

71.4

Sol id ago sempervirens

Perennial

Broad-leaved herbaceous

78.6

the species were perennials and four were annuals. Eight species
were grasses, five were sedges, 10 were broad-leaved herbaceous
species and one species was a cattail (Table 2). No woody species
were found. Thirteen of the species occurred in more than half of
the sites and Iva texensis (sumpweed) was present at each site.
Conoclinium betonicifolium (mist flower) and Schoenoplectus
pungens (bulrush) each occurred in all but one of the wetlands
(92.9%).

Sampling at Site 1 produced 14 species (58.3% of the total).
Site 2 added four new species and resulted in 75% of the total for
the 14 sites. After Site 2, no more than a single species was added

JUDD & LONARD

at a site. Sampling through Site 9 produced 95.8% of the total
species and only a single new species was added thereafter (at Site
12).

Species richness at the wetland sites ranged from 7 to 14 and 1 1
of the wetlands (78.6%) had 12 or more species (Table 3). Mean
species richness was 11.9 (SD = 2.17) and the 95% confidence
interval ranged from 10.6 to 13.2. Species diversity values ranged
from 0.533 to 1.005 (Table 3). Mean species diversity was 0.855
(SD = 0.134) and the 95% confidence interval extended from 0.778
to 0.932. Evenness showed even less variation (Table 3). The
mean was 0.797 (SD = 0.084) and the 95% confidence interval
ranged from 0.750 to 0.844.

Judd & Lonard (2004) provide data on species richness, species
diversity, and evenness in freshwater, brackish, and saltwater
marshes in the Rio Grande Delta of the Texas mainland adjacent to
South Padre Island. The means for freshwater and brackish
marshes were calculated from their data and compared with the
means for these parameters in the South Padre Island wetlands. Six
mainland freshwater marshes had a mean species richness of 25.3
(SD = 6.25), mean species diversity of 1.394 (SD = 0.120), and
mean evenness of 0.788 (SD = 0.05). Nine mainland brackish
marshes had a mean species richness of 17.9 (SD = 5.578), mean
species diversity of 1.262 (SD = 0.184) and mean evenness of 0.783
(SD = 0.056). South Padre Island wetlands had significantly lower
species richness than both freshwater (t = 7.279, df, P < 0.001)
and brackish (t = 3.656, 2\ df, P < 0.01) mainland marshes.
Likewise, South Padre Island wetlands had significantly lower
species diversity than mainland freshwater marshes (t = 8.502, 18
df,P< 0.001) or mainland brackish marshes (t = 5.700, 21 df^P<
0.001). However, there was no significant difference in evenness
between South Padre Island wetlands and either mainland
freshwater marshes (/ = 0.243, df, P > 0,5) or mainland brackish
marshes (t = 0.44, 21 df,P> 0.5).

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

Table 3. Comparison of species ricliness (N), species diversity (H’), and evenness (J’)
among wetlands on South Padre Island, Texas.

Wetland

H’

J’

1.005

0.877

0.787

0.708

0.848

0.761

0.947

0.878

1.001

0.873

0.876

0.812

0.918

0.801

0.809

0.726

0.994

0.892

0.899

0.775

0.719

0.667

0.738

0.533

0.631

0.904

0.838

Similarity in species composition among marshes was high
(Table 4) with coefficients for 91 pair-wise comparisons ranging
from 0.300 to 0.857. The mean coefficient of similarity was 0.644
(SD = 0.138) and the 95% confidence interval of the mean ranged
from 0.615 to 0.673. Most of the wetlands shared 62% to 67% of
their species. The similarity is significantly greater (t = 9.499,104
df,P< 0.001) than the mean of 0.322 {SD = 0.116) for 15
coefficients of freshwater marshes and the mean of 0.258 {SD =
0.123) for 36 coefficients of brackish water mainland marshes {t =
16.176, 125 df,P< 0.001) reported for the adjacent Texas mainland
by Judd & Lonard (2004).

Cover of dead plant material ranged from 17.1% to 97.3%
among the sites. The mean was 67.4% {SD = 26.1). Cover of live
plants ranged from 27.4% to 80.5% and the mean was 56.9% {SD =
22.7). Much of the dead and live plant material overlapped, but
relatively little of the surface was bare. The top three species in
importance were abundant in both frequency (range 57.1% to
100%) and cover. The sum of the relative cover for the top three
species in importance at a site ranged from 63.5% to 97.7%. The
other species present at a site contributed little to the abundance of

JUDD & LONARD

Table 4. Comparison of Sorensen’s community similarity coefficients among freshwater
and brackish wetlands of South Padre Island, Texas.

Site

.667

.593

.692

.714

.615

.786

.593

.667

.615

.636

.476

.692

.846

.800

.741

.720

.815

.615

.692

.696

.720

.381

.400

.640

.720

.741

.720

.741

.615

.538

.696

.640

.381

.300

.640

.769

.750

.769

.720

.818

.750

.500

.421

.750

.846

.857

.815

.741

.750

.692

.455

.381

.769

.800

.640

.727

.667

.500

.526

.750

.741

.815

.750

.615

.455

.476

.692

.769

.696

.640

.476

.400

.640

.696

.640

.381

.400

.640

.727

.444

.353

.818

.500

.421

.833

.667

.600

.526

vegetation at the site. Consequently, data on species importance are
presented for only the first three species in importance at each site
(Table 5). Schoenoplectus pungens (bulrush) was among the top
three species in importance in 13 of the wetlands (92.9%) and it
was the dominant species in nine wetlands (64.3%) (Table 5).
Fuirena simplex (umbrella grass) and Spartina patens (marshhay
cordgrass) each were dominant in two wetlands and Andropogon
glomeratus (bushy bluestem) was dominant in one (Table 5). Two
of the dominants were sedges (bulrush and umbrella grass) and two
were grasses (bushy bluestem and marshhay cordgrass). None of
the broad-leaved species were dominants and only four were one of
the top three species at a particular site.

Discussion

Water in the wetlands of the secondary dunes and vegetated flats
zone of South Padre Island occurs as a result of rainfall. Thus, it is
fresh when it falls and it becomes progressively more brackish as
water evaporates and the remaining standing water accumulates salt
from wind transport. Consequently, a given wetland might contain
freshwater if sampled soon after a rain and brackish water if

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

sampled a week or more later. Thus, South Padre Island may not
have wetlands that are permanently fresh or brackish. Salinity
varies with the proximity of rain events. The wetlands are dry
much of the time; especially so in drought years. Only 28.6% of
the wetlands contained water at the time we sampled them. This
was likely due to the fact that rainfall was below normal in seven of
the 1 1 months of this study.

Each of the South Padre Island wetlands was sampled only once.
Thus, it is not known if there is significant seasonal variation in any
of the parameters studied here. However, it seems unlikely because
only four species (16.7 % of the total, Table 2) are annuals and only
two of these four species ranked in the top three species in
importance at a given site (Table 5). None of the four species was
dominant at a site.

South Padre Island freshwater and brackish wetlands have fewer
species and lower species diversity than mainland freshwater and
brackish marshes of the adjacent Rio Grande Delta (Judd & Lonard
2004), but evenness is similar in the island and mainland wetlands.
Only six (24%) of the species occurring in South Padre Island
wetlands in this study were also found in mainland wetlands by
Judd & Lonard (2004). All six species occurred in brackish
marshes on the mainland. Schoenoplectus pungens and
Conoclinhim betonicifolium occurred only in mainland brackish
marshes, Typha domingensis and Bacopa monnieri were found in
freshwater, brackish, and salt marshes of the adjacent mainland.
Sporobolus virginicus and Borrichicia frutescens occurred in
brackish and salt marshes on the mainland. Three of the species
were dominant in one or more marshes on the mainland. Typha
domingensis was dominant in one of six freshwater marshes and in
three of nine brackish marshes on the mainland. Borrichia
frutescens was dominant in one of nine brackish marshes and two
of 1 1 salt marshes on the mainland. Sporobolus virginicus was
dominant in two salt marshes on the mainland. None of the
dominant species in island wetlands were also dominant in

JUDD & LONARD

Table 5. Comparison of species importance among freshwater and brackish wetlands on
South Padre Island, Texas, Freq. = frequency, Rel. Freq. = relative frequency, Rel.
Cover = Relative Cover, Imp. Val. = importance value (sum of relative frequency
and relative cover).

Site

Species

Freq.

Rel.

Freq.

Cover

Rel.

Cover

Imp.

Val.

Schoenoplectiis pungem

100.0

14.3

14.23

35.5

49.8

Spartina patens

85.7

12.2

11.21

28.0

40.2

Iva texensis

1 1 additional species

85.7

Total

12.2

Cover

5.61

40.03

14.0

26.2

Schoenoplectus pimgens

100.0

15.7

65.64

81.6

97.3

Iva texensis

87.5

13.7

4.31

5.4

19.1

Conoclinium betonicifolinm

1 0 additional species

87.5

Total

13.7

Cover

2.16

80.5

2.7

16.4

Schoenoplectiis pimgens

100.0

17.9

13.13

47.9

65.8

Iva texensis

100.0

17.9

10.31

37.6

55.8

Spartina patens

10 additional species

85.7

Total

15.4

Cover

1.09

27.44

4.0

19.4

Schoenoplectus pungem

100.0

13.7

15.34

36.2

49.9

Solidago sempervirem

100.0

13.7

5.96

14.0

27.7

Spartina patem

9 additional species

100.0

Total

13.7

Cover

5.66

42.41

13.3

27.0

Schoenoplectus pungem

100.0

10.1

14.60

31.2

41.3

Iva texensis

100.0

10.1

11.00

23.5

33.6

Andropogon glomeratus

1 1 additional species

100.0

Total

10.1

Cover

7.60

46.70

16.3

26.4

Schoenoplectus pungem

100.0

14.0

30.70

59.8

73.8

Conoclinium betonicifolinm

100.0

14.0

5.54

10.8

24.8

Spartina patem

9 additional species

100.0

Total

14.0

Cover

2.59

51.38

5.0

19.0

Fuirena simplex

100.0

12.3

34.86

55.6

67.9

Andropogon glomeratus

100.0

12.3

16.84

26.9

39.2

Schoenoplectus pungem

1 1 additional species

100.0

Total

12.3

Cover

2.80

62.68

4.5

16.8

Fuirena simplex

100.0

14.3

62.61

79.9

94.2

Schoenoplectus pungem

100.0

14.3

6.71

8.6

22.9

Panicum amarum

10 additional species

100.0

Total

14.3

Cover

2.09

73.34

2.7

17.0

Schoenoplectus pungem

100.0

11.5

16.03

25.1

36.6

Eleocharis geniculata

71.4

8.2

16.30

25.5

33.7

Panicum amarum

100.0

11.5

10.36

16.2

27.7

10 additional species Total Cover 63.89

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

Table 5. Cont.

Site

Species

Freq.

Rel.

Freq.

Cover

Rel.

Cover

Imp.

Val.

Schoenoplectus pimgens

100.0

14.0

24.10

32.1

46.1

Andropogon glomeratiis

100.0

14.0

23.11

30.8

44.8

Fuirena simplex

100.0

14.0

10.83

14.4

28.4

1 additional species

Total

Cover

75.07

Schoenoplectus pimgens

100.0

19.4

47.94

62.7

82.1

Borrichia frntescens

100.0

19.4

21.40

28.0

47.4

Bacopa monnieri

85.7

16.7

2.90

3.8

20.5

9 additional species

Total

Cover

76.48

Spartina patens

100.0

25.0

30.36

54.0

79.0

Schoenoplectus pimgens

100.0

25.0

19.61

34.9

59.9

Borrichia frntescens

85.7

21.4

4.97

8.8

30.2

5 additional species

Total

Cover

56.18

Spartina patens

100.0

36.8

60.27

91.7

128.5

Borrichia frntescens

57.1

21.1

0.63

1.0

22.2

Iva texensis

42.9

15.8

1.17

1.8

17.6

4 additional species

Total

Cover

65.74

Andropogon glomeratiis

85.7

15.0

11.99

34.9

49.9

Schoenoplectus pnngens

100.0

17.5

7.73

22.5

40.0

Spartina patens

100.0

17.5

5.81

16.9

34.4

9 additional species

Total

Cover

34.33

mainland wetland communities. Clearly the island wetland
communities are markedly different than wetland communities of
the adjacent mainland. They have fewer species, mostly different
species, and different dominant species. It is unlikely that
additional sampling on South Padre Island would have added many
more species because sampling through the first nine sites resulted
in 23 of the 24 species found. Sampling five more sites produced
only one species. Similarity of species composition among
freshwater and brackish wetlands of South Padre Island is high.
The higher similarity among island wetlands is likely related to the
much lower total species richness (24 species) compared to the
mainland freshwater and brackish marshes (81 species each).

JUDD & LONARD

Baccus & Horton (1979) identify 13 species that occur in the
low marshy areas of the heavily vegetated barrier flat at Padre
Island National Seashore. Thus, species richness was similar to
what we found at South Padre Island (mean = 1 1.9, 95% confidence
interval = 10.6 to 13.2). However, similarity in species composi¬
tion was modest. Only five of the 13 species Baccus & Horton
(1979) list for Padre Island National Seashore were found among
the 14 sampling sites in this study. This might be six species in
common if the Eleocharis sp. they list was E. geniculata. Species
in common increases to eight if species known to occur on South
Padre Island are included (Lonard et al. 1978) but not found in this
study, and it might be increased to nine if the Eleocharis sp. was E.
geniculata^ E. montevidensis, or E. obtusa, which are known to be
present on South Padre Island (Lonard et al. 1978; this study).
Nelson et al. (2001) compare the floras of Matagorda Island,
Mustang Island, North Padre Island and South Padre Island. Each
species in the wetlands of South Padre Island is known to occur on
North Padre Island.

Baccus & Horton (1979) found that the dominant species among
the wetland species was Eleocharis interstincta. Eleocharis
interstincta was not found in this study and Lonard et al. (1978) did
not find it on South Padre Island. A different species of Eleocharis
{E. geniculata) was present at six of the 14 wetlands sampled, but it
was abundant at only one site (2”*^ in importance at Site 9).

Nelson et al. (2000) reported on the vegetation and fioristics of
four communities in the Big Ball Hill region of Padre Island
National Seashore. They identified Andropogon glomeratus and
Spartina patens as dominants in the lowland subcommunity of the
barrier flat community. Borrichia frutescens, Hydrocotyle
bonariensis, Samolus ebracteatus, and Schoenoplectus pungens
were listed as species typically present in the lowland sub¬
community. They do not suggest that this list is complete for the
lowland sites. All of the species, except H, bonariensis were
present among the sites studied at South Padre Island, but only

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

Spartina patens occurred in Baccus & Horton’s (1979) list of
species occurring in low marshy areas of the vegetated flats of
Padre Island National Seashore. Thus, there is less similarity in
species present in lowland barrier flat communities on North Padre
Island than there is between either of the studies and the species
present in the wetlands of South Padre Island.

Mean species richness in the barrier flat community (both low
and higher elevations) of Big Ball Hill region of Padre Island
National Seashore was reported to be 20.3 (Nelson et al. 2000).
This is almost double the mean species richness of 11.9 found
among the wetlands at South Padre Island. It is not surprising since
only the lowland component of the barrier flats was included.
Nelson et al. (2000) reported a mean species diversity for the
barrier flat community of 0.58 and a mean evenness of 0.61. They
do not provide a measure of variance for either mean, but both
means appear to be significantly lower than those reported here
because the values fall below the 95% confidence intervals for
species diversity and evenness in wetlands at South Padre Island.

Conclusions

Based on the meager data available, it appears that species
richness at wetland sites is similar on the northern and southern
portions of Padre Island, but there is only modest similarity in
species composition of wetland communities. There is considerable
variation in species importance among South Padre Island sites and
marked variation between southern and northern portions of Padre
Island. Additional study of Padre Island and other Texas barrier
islands is needed to ascertain if the differences in composition of
barrier island freshwater and brackish wetland communities
reported here are stochastic or related to variation in environmental
variables.

Literature Cited

Baccus, J. T. & J. K. Horton. 1979. An ecological and sedimentary study of Padre Island
National Seashore. Report to Office of Natural Resources, Southwest Region,
National Park Service, Santa Fe, New Mexico. Contract No. CX 702970059, 272 pp.

JUDD & LONARD

Brower, J. E., J. H. Zar & C. N. Von Ende. 1998. Field and Laboratory Methods for
General Ecology. WCB/McGraw-Hill, Boston, Massachusetts, U.S.A., 273 pp.

Canfield, R. H. 1941. Application of the line interception method m samplmg range
vegetation. Journal of Forestry, 39(4):388-394.

Dahl, B. E., B. A. Fall, A. Lohse & S. G. Appan. 1974. Stabilization and reconstruction
of Texas foredunes with vegetation. Gulf Universities Research Consortium, 139:1-
325.

Everitt, J. H., D. E. Escobar & F. W. Judd. 1991. Evaluation of airborne video imagery
for distinguishing black mangrove {Avicennia germinans) on the lower Texas gulf
coast. Journal of Coastal Research, 7(4): 1 169-1 173.

Everitt, J. H., D. E. Escobar, F. W. Judd & M. R. Davis. 1992. Evaluation of spot
satellite and airborne video imagery for distinguishing black mangrove {Avicennia
germinans). Pp. 169-175, in Proceedings 13*^' Biemiial Workshop on Color Aerial
Photography and Videography in the Plant Sciences, Orlando, Florida. May 6-9,
1991, (Published in 1992).

Everitt, J. H., M. A. Alaniz, D. E. Escobar, R. I. Lonard, F. W. Judd & M. R. Davis.
1999. Reflectance characteristics and film image relations among important plant
species on South Padre Island, Texas. Journal of Coastal Research, 15(3):789-795.

Jones, S. D. & J. K. Wipff. 2003. A 2003 updated checklist of the vascular plants of
Texas. Botanical Research Center, Bryan, TX. (CD-ROM). 697 pp.

Judd, F. W., R. I. Lonard & S. L. Sides. 1977. The vegetation of South Padre Island,
Texas in relation to topography. Southwestern Naturalist, 22(l):31-48.

Judd, F. W. & S. L. Sides. 1983. Effects of Hurricane Allen on the nearshore vegetation
of South Padre Island. Southwestern Naturalist, 28(3):365-369.

Judd, F. W. & R. 1. Lonard. 1985. Effects of perturbations on South Padre Island. Pp.
1855-1869, in Proceedings Fifth Symposium on Coastal and Ocean Management,
“Coastal Zone ‘85”. American Soc. of Civil Engineers. Baltimore, Maryland, 2672

pp.

Judd, F. W. & R. 1. Lonard. 1987. Disturbance and community development. Pp. 1731-
1745, in Proceedings Fifth Symposium on Coastal and Ocean Management, “Coastal
Zone’87”. American Soc. of Civil Engineers. Seattle, Washington, 5870 pp.

Judd, F. W., R. 1. Lonard, J. H. Everitt & R. Villarreal. 1989. Effects of vehicular traffic
in secondary dunes and vegetated flats of South Padre Island, Texas. Pp. 4634-4645,
in Proceedings Sixth Symposium on Coastal and Ocean Management, “Coastal Zone
‘89”. American Soc. of Civil Engineers. Charleston, South Carolina, 4978 pp.

Judd, F. W., R. 1. Lonard, J. H. Everitt & D. E. Escobar. 1991. Resilience of seacoast
bluestem bamer island communities. Pp. 3513-3524, in Proceedings Seventh
Symposium on Coastal and Ocean Management. “Coastal Zone ‘91”. American
Soc. of Civil Engineers. Long Beach, California, 3800 pp.

Judd, F. W., R. 1. Lonard, D. L. Hockaday, D. E. Escobar, J. H. Everitt & R. Davis.
1994. Remote sensing of nearshore vegetation. South Padre Island, Texas. Pp. 581-
589, in Proceedings of the Second Thematic Conference on Remote Sensing for
Marme and Coastal Enviromnents. Vol. 1. Needs, Solutions and Applications.
Environmental Research Institute of Michigan (ERIM), P, O. Box 134001, Ann
Arbor, MI 48113-4001, U.S.A, 704 pp.

Judd, F. W., R. 1. Lonard, J. H. Everitt, D. E. Escobar & M. R. Davis. 1998. Wind-tidal
flats and dune vegetation of South Padre Island, Texas. Pp. 177-184, in Vol. II,

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Proceedings Fifth International Conference on Remote Sensing for Marine and
Coastal Environments. San Diego, California. 5-7 October 1998, 584 pp.

Judd, F. W. & R. I. Lonard. 2004. Conmiunity ecology of freshwater, brackish, and salt
marshes of the Rio Grande Delta. Texas J. Sci., 56(2): 103-122.

Judd, F. W., R. I. Lonard, K. R. Summy & R. A. Mazariegos. 2007. Seasonal variation
in dune vegetation at South Padre Island, Texas. Texas J. Sci., 59(2); 1 13-126.

Krebs, C. J. 1999. Ecological Methodology. Menlo Park, California. Addison Wesley
Longman, 620 pp.

Lonard, R. L, F. W. Judd & S. L. Sides. 1978. Aimotated checklist of the flowering
plants of South Padre Island, Texas. Southwestern Naturalist, 23(3):497-510.

Lonard, R. 1. & F. W. Judd. 1980. Ph>^ogeography of South Padre Island, Texas.
Southwestern Naturalist, 25(3):3 13-322.

Lonard, R. 1. & F. W. Judd. 1981. The terrestrial flora of South Padre Island, Texas.
Texas Memorial Museum Miscellaneous Papers No. 6, 74 pp.

Lonard, R. 1. & F. W. Judd. 1989. Phenology of native angiosperms of South Padre
Island, Texas. Pp. 217-222, in Proceedings Eleventh North American Prairie
Conference. T. Bragg and J. Stubbendieck (Eds.). Univ. Nebraska Printing, Lmcoln,
Nebraska, 293 pp.

Lonard, R. L, F. W. Judd, J. H. Everitt & D. E. Escobar. 1991. Roadside associated
disturbance on coastal dunes. Pp. 2823-2836, in Proceedings Seventh Symposium on
Coastal and Ocean Management, “Coastal Zone ‘91”. American Soc. of Civil
Engineers. Long Beach, California, 3800 pp.

Lonard, R. I. & F. W. Judd. 1993. Recovery of vegetation of barrier island washover
zones. Pp. 2324-2331, in Proceeding of the Eighth Symposium on Coastal and
Ocean Management, “Coastal Zone ‘93”. American Soc. Civil Engineers. New
Orleans, Louisiana, 3512 pp.

Lonard, R. 1. & F. W. Judd. 1997. The biological flora of coastal dunes and wetlands.
Sesuviiim portulacastnim (L.) L. Journal of Coastal Research, 13(1):96-104.

Lonard, R. 1. & F. W. Judd. 1999. The biological flora of coastal dunes and wetlands.
Ipomoea imperati (Vahl) Griseb. Journal of Coastal Research, 15(3):645-652.

Lonard, R. L, F. W. Judd, J. H. Everitt, D. E. Escobar, M. A. Alaniz, 1. Cavazos III & M.
R. Davis. 1999. Vegetative change on South Padre Island, Texas, over twenty years
and evaluation of multispectral videography in determining vegetative cover and
species identity. Southwestern Naturalist, 44(3):26 1-271.

Nelson, A. D., J. R. Goetze, I. G. Negrete, V. E. French, M. P. Johnson & L. M. Macke.
2000. Vegetational analysis and floristics of four communities in the Big Ball Hill,
region of Padre Island National Seashore. Soutliwestem Naturalist, 45(4):43 1-442.

Nelson, A., J. Goetze & A. Lucksinger. 2001. A comparison of the flora of northern
Padre Island to that of Matagorda Island, Mustang Island and southern Padre Island,
Texas. Occas. Papers, Museum of Texas Tech University, Number 209:1-23.

FWJ at; Qudd@utpa.edu

TEXAS J. OF SCI. 61(2);97-118

MAY, 2009

SEASONAL TROPHIC ECOLOGY OF THE WHITE-ANKLED MOUSE,
PEROMYSCUS PECTORALIS (KODENTIA: MURID AE)

IN CENTRAL TEXAS

John T. Baccus, John M. Hardwick, David G. Huffman
and Mark A. Kainer

Wildlife Ecology Program, Department of Biology
Texas State University, San Marcos, Texas 78666

Abstract-Fruits and seeds, green foliage, and animal matter composed the
seasonal diets of Peromysciis pectoralis in central Texas. Based on changes in foods
consumed, five distinct seasons were delineated. Trophic diversity differed among
seasons, with diversity highest in summer and lowest in winter. Diets for winter and
autumn were more similar; whereas, diets for winter and late spring, winter and
summer, and late spring and autumn were more dissimilar. Peromysciis pectoralis in
central Texas are primarily frugivorous/granivorous herbivores with omnivorous
tendencies reflecting opportunistic feeding habits.

Species of Peromyscus have been characterized as granivores
with omnivorous tendencies that reflect opportunism in feeding
habits (Cogshall 1928; Baker 1971; Montgomery 1989). Most
dietary studies have compared two or more closely related species
in sympatry and generally found Peromyscus are efficient foragers
with broad diets consisting of seeds, fruits, green vegetation, and
arthropods (Cogshall 1928; Hamilton 1941; Jameson 1952;
Williams 1959; M'Closkey & Fieldwick 1975; Grant 1978; Knuth
& Barrett 1984). Before interspecific interactions between
potentially competing congeners can be fully determined, studies of
each species in the absence of potential competitors are needed to
ascertain the breadth of the trophic niche. In addition, climatic and
phenological events may cause notable changes in community
structure that affect the quantity and quality of food resources and
shape the trophic niche of a species (Waser 1978a; 1978b).
Seasonality in diet is evident in populations of most northern
species of Peromysus, Because of the paucity of seasonal data for
southern populations, a nonexistent or limited dietary cycle in these
populations has been suggested (Montgomery 1989).

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

The white-ankled mouse, Peromyscus pectoralis Osgood,
occurs in a variety of habitats over the central Mexican Plateau and
Sierra Madre Oriental in Mexico northward to southeastern New
Mexico and western and central Texas into southern Oklahoma.
The species has a propensity for rocky environments, especially
rock outcrops (Kilpatrick & Caire 1973; Schmidly 1972; 1974;
Baccus & Horton 1984; Etheredge et al. 1989). Although
substantial information exists about the habitat affinities of this
species, the trophic niche of this mouse is poorly known. Alvarez
(1963) observed the species eating fruits of nopal (prickly pear,
Opuntia lindheimeri) cactus in Tamaulipas, Mexico. In Texas,
Davis (1974) reported consumption of juniper berries, hackberry
seeds, and acorns in central Texas. Schmidly (2004) mentioned the
lack of a detailed food habits study for the species and speculated
the diet consisted of seeds, cactus fruits, lichen, fungi, and insects.
Here, the first detailed analysis and description of the seasonal
trophic niche of P. pectoralis in central Texas is presented. No
other sympatric congeners (i.e., P. attwateri) inhabited the study
site (Mullican & Baccus 1990); thus, no accounting for interspecific
competition or differential use of resources was necessary in the
analysis.

Methods

White-ankled mice were collected 5 km W. San Marcos, Hays
County Texas (29°47' N, 97°58' W) in an abandoned limestone
quarry at the eastern periphery of its distribution. The landscape
consists of large to medium size, strewn boulders and truncated
limestone outcrops. Ashe juniper trees (Juniperus ashei) dominate
the woody vegetation with trees and shrubs of green sumac {Rhus
virens), Texas persimmon (Diospyros texana), sugar hackberry
(Celtis laevigata), plateau live oak (Quercus fusiformis), agarito
(Mahonia trifoliolata), and Roosevelt weed {Baccharis neglecta)
also present. Herbaeceous vegetation consists of Johnsongrass
{Sorghum halepense). King Ranch bluestem {Bothriochola
ischaemum), little bluestem {Schizachyrium scoparium), Texas
wintergrass {Stipa leucotricha), threeawn grass {Aristida sp.).

BACCUS ET AL.

prickly pear, frostweed (Verbesina virginica), Lindheimer senna
(Cassia lindheimeriana), knotted hedgeparsley (Torilis nodosa),
prairie bluet (Heydotis nigricans), Drummond skullcap (Scutellaria
drumondii), white sweetclover (Melilotus albus), oneseed croton
(Croton monanthogynus), Indianmallow (Abutilon incanum),
common sorrel (Rumex acetosella), sensitivebriar (Schrankia
roemeriana), pepperweed (Lepidiim virginicum), and Texas
bluebonnet (Lupinus texensis). Plant taxonomy followed Hatch et
ah (1990).

Thomthwaite (1948) classified the climate of Hays County as
C1B4 (dry subhumid, mesothermal) with a mean annual potential
evapotranspiration of 99.7 to 114 cm. The mean monthly

maximum temperature is 35 C (July) with a mean monthly
minimum of 5°C (January). Mean annual precipitation is 85.7 cm.
During this study, annual precipitation was below normal (76 cm).

Monthly collections between October 1976 and September 1977
were accomplished by Museum Special snap-traps set along 100
trap-station transects (spacing 15 m) during 12 consecutive days
each month (total for study = 14,400 trap-days). Age (Schmidly
1972), standard external measurements, gender, and reproductive
condition were recorded for each mouse. White-ankled mice were
collected under Texas Scientific Permit SPR-0890-234.

Quantitative evaluations of stomach contents from 149 (67
females and 82 males) P. pectoralis followed relative-occurrence
evaluation and histomicroscopic methods (Sparks & Malechek
1968; Free et al. 1970; Hansson 1970; Westoby et al. 1976; Ellis et
al. 1977; Dawson & Ellis 1979; Cockbum 1980; Copley &

Robinson 1983; Mclnnis et al. 1983; Copson 1986; Carron et al.

1990). Stomach contents were washed through a 0.125-nim sieve,
homogenized, equally partitioned, and placed on 5 microscopic
slides. These contents were considered as a whole or 100%
irrespective of the extent of stomach fill (Hansson 1970). Twenty
randomly selected microscopic fields with food items were

100

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

examined per slide under lOOX magnification (total fields
examined = 14,900). Identified food items in each field were
recorded. Reference slides were also prepared of plant and animal
matter from the collection site after mincing in a blender to mimic
mastication.

Identification of food items and classification categories were
based on anatomical or morphological structures of plant and
animal matter. However, some matter due to extensive mastication
remained unidentified. Ingested bait was ignored. Food categories
applied to stomach contents were: (I) fruits and seeds-remains of
testa, endosperm, exocarp, or individual minute seeds; (2) green
foliage-leaf or stem tissues; (3) larval insects- nonsclerotized soft-
bodies; (4) adult insects-sclerotized parts such as antennae, elytra,
mouthparts, wings, or appendages; and (5) animal matter-primarily
muscle tissue and hair. Fruits and seeds were combined as a food
category because of a strong tendency of co-occurrence (Pitts &
Barbour 1979). Calculations of percent frequency (%f), percent
volume (%v) and relative importance (I) for each food item
followed Obrtel & Holisova (1974; 1981). Composition

percentages were obtained by dividing the total number of
fragments of a given food item by the total number of fragments of
all foods encountered. Analyses showed no difference in percent
frequency of food items in diets of males and females; therefore,
data for males and females were pooled.

Seasonal variation in food habits of small mammals has been
primarily demonstrated by two methods. A season is defined by
one method as a set number of months, usually four periods of three
months, and food items enumerated within a season (Hamilton
1941; Whitaker 1966; Houtcooper 1978; Luo & Fox 1994). The
second method lists food items consumed each month (Jameson
1952; Myers & Vaughan 1964; Bradley & Mauer 1971; Luce et al.
1980; Armgard & Batzli 1996). Both methods fail to reveal the
dynamics of rapid dietary change. It is possible that seasonal
change in animal diets may not be reflected by the four customary

BACCUS ET AL.

101

seasons; rather, based on dietary changes, one might find fewer or a
greater number of seasons.

An Index of Change based on dietary dynamics was developed
using a two-day moving average of the abundance of food items for
groups of a set number of mice. The difference between group
means for two consecutive such groups of mice was divided by the

mean collection date of the mice in
Change is represented by the formula:

both groups. The Index of

m+k

m+2k

(Z-)

- z

(Z-)

j=l

i=m

j=l

i=m+k+l

m+k

m+2k

[(I “) -

i=m

i=m+k+l

where;

k = number of mice included in the first or second half
(group) of the moving mean interval

m = accession number of the first mouse in the interval,
with m incrementing by unity after each solution
of the expression

X = percent composition of food item j in mouse i
V = total number of different food items

d = Julian day of capture for mouse i

102

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

Seasonal recognition was determined by the extent of an abrupt
change in the diet. For the purposes of this study, any change >10
was defined as a substantial dietary change and recognized as the
beginning of a season. Also, the stability of the period following an
abrupt change was considered. If the Index of Change did not vary
by a factor >5 during a period of time, this sequence of days was
considered as the term of a season. The next change in the index
>10 marked the start of the next season.

Preference for food items was determined by a modified Ivlev's
Index of Electivity (Ivlev (1961; Jacobs 1974; Krebs 1999). The
electivity index indicated whether a plant was consumed in amounts
similar to availability in the environment or active selection of the
food. A value of zero indicated randomness in resource selection, a
positive index indicated the food was selected in amounts greater
than would be expected by chance encounter, and a negative index
signified consumption of a food in quantities lower than predicted
by randomness. The z test for comparing sample proportions
(Sokal & Rholf 1969) was used to determine differences in male
and female diets.

Seasonal diversity in the diet was calculated by Brillouin's Index
of Diversity because samples were treated as collections rather than
random samples from a larger biological community (Margalef
1958; Pielou 1966; Krebs 1999). Seasonal overlap in diet was
determined by Morisita's Index of Similarity (Morisita 1959; Krebs
1999). The overlap coefficient varies from 0 when diets are
completely distinct (no food categories in common) to 1 when diets
are identical. Values >0.60 indicate significant overlap (Zaret &
Rani 1971), but this should not be construed as statistically
significance. Seasonal differences in the proportion of plant and
animal matter in the diet were tested by a Goodness-of-Fit test.

Herbaceous vegetation occurring on the study site was assessed
in January, April, August and November by randomly dropping a
0.5 m x 1 m quadrat 100 times in the study area. From these data.

BACCUS ET AL.

103

Table 1. Characteristics of the seasonal diet of Peromyscus pectoralis in central Texas.

Criterion

Winter

Early

Spring

Late

Spring

Summer

Autumn

Sample

size (n)

Number

of items (S)

Plant (%)

5 (62.5)

7 (70.0)

5 (62.5)

11(73.3)

8 (66.7)

Animal (%)

3 (37.5)

3 (30.0)

3 (37.5)

4 (26.4)

4(33.3)

Equally

common species

6.19

7.44

7.25

9.32

7.82

Trophic

diversity (H)

2.52

2.78

2.71

3.736

2.87

percent cover of each plant species was estimated (Myers &
Vaughan 1964). Seed abundance was sampled by placing a 25 cm
X 10 cm quadrat in the lower right comer of the larger quadrat at 10
sampling locations, removing the top 1-2 cm of soil, and screening
soil samples through a series of descending size sieves (2.0 mm- 180
pm). Seeds were removed, counted, and identified to species.

The arthropod fauna was sampled monthly by sweeping through
the ground vegetation along each trap-line with an insect net (38-
cm diameter) (Beiner 1955). Arthropods inhabiting the foliage of
trees were not sampled.

Results

Twenty-three food items (18 plant and 5 animal), broadly
categorized into 3 trophic groups (fruit and seed, green foliage, and
animal matter), comprised the seasonal diet of P. pectoralis in
central Texas (Table 1). No mouse exclusively consumed plant or
animal matter. There was no difference in percent frequency of
food items in diets of males and females (z = 0.0936, P > 0.34).
The percent frequency, percent volume, and importance value of
food items in the diet varied seasonally (Table 2). Prickly pear
foliage, larval and adult insects, and arachnids were the only food
items consumed in all seasons. Ten food items had a percent fre-

Table 2. Percent frequencies (%f), percent volumes (%v) and importance values (I) of items identified in the seasonal diet of Feromyscus
pectoral is in central Texas.

Winter Early Spring Late Spring Summer Autumn

104

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

'S' ^

—

VO —

rn 'if

t~- m vO

ci ri d

fS O' p

c-'‘ 'f r'-'

"sr r~; p
d m »n

\C so

»o p

rn so

'f d

p
fn d

<NJ p p

d so d

P O'.

<N p P

d d

d d d

'Sf

*r) —

fsj OO

p fS p

d vi

sd 00 d

fn —•

OO 0©

<N f'' 0©

>—

00 so

p sn p

•d d d

(NJ

so so
d d

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OO p 0©

odd

SO OO p
ri d d

a •§ M

I III

S ’I -S .2 -i

s > ^ P ft

g ^
ClP -a

i ,

; g o

^ i s s:-« ^ I

C3 2

•H *o =§

&.-« .s:

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y>:2

.ts

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OOdCsoiXCQOD

S o
■S

V c
*o w

- s2

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1-^

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Sylvilagus jloridanus

BACCUS ET AL.

105

quency of < 5%. These foods were a miscellaneous assortment of
animal tissues, green foliage, seeds, fruits, and possibly flower
inflorescence. Twenty-two percent of stomachs had plant
fragments so masticated that identification only as dicot material
was possible. No monocot, lichen, or fungal materials were
identified in any stomach. The greatest number of different food
items found in any one stomach was 6 (X= 3.26, SE = 0.73 ).
Based on percent volume and percent frequency, 2 trophic
categories (fruit and seed and animal matter) composed the bulk of
seasonal diets. Seeds and fruits were the most frequently
encountered category in the seasonal diet, occurring in 87% of
stomachs. Ashe juniper berries were the most frequently consumed
fruit. Arthropods were the primary animal matter in the seasonal
diet with 84% of stomachs containing mostly insect fragments.
Insects consumed were from the following taxa: Order
Hymenoptera, Families Formicidae and Apidae; Order Coleoptera,
Family Scarabaeidae; Order Orthoptera, Family Gryllidae; Order
Collembola; Order Lepidoptera, Family Pyralidae; and Order
Diptera.

Five seasons (winter, December-January; early spring, February-
March; late spring, April-May; summer, June-August; and autumn,
September- November) were delineated by an Index of Change
(Fig. 1). Morisita's Index of Similarity indicated no two seasons
had unity (Table 3), yet some seasons were similar because of
dietary overlap >0.60. The temporal continuity of resource use by
white-ankled mice and dynamic changes in the diet demonstrated a
general trend in which the immediate juxtaposed seasons to a given
season were most similar (i.e., winter diet was most similar to the
previous season, autumn, and the following season, late spring).
The exception to this trend was autumn. A similarity value (0.42)
between summer and autumn diets indicated substantial
dissimilarities between these juxtaposed seasons; however, autumn
and winter diets had the highest similarity. Furthermore, the
autumn diet was more similar to the early spring diet. The
communality of diet for these three seasons was based on the extent

106

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

40 r

Figure 1. Eidex of change in the diet of Peromysciis pectoralis collected in central Texas
as a function of date and season. Each of the five major peaks of the graph represent
the start of a season (M Dec = winter, E Feb = early spring, E Apr = late spring, E
Jun = sununer, E Sept = Autumn). Letters representmg months mark the early,
middle, and late portions of each month.

of fruit and seed use (primarily Ashe juniper berries) by white-
ankled mice.

The substantial and almost exclusive use of the fruit and seed
category was the distinctive feature of the winter diet (Table 2, Fig.
2). As a result, the winter diet was the most homogenous and least
diverse of any season. The high use of fruit and seed in comparison
to minimal consumption of animal matter (arthropods) and green
foliage resulted in low seasonal diversity. Only Ashe juniper and
green sumac berries were important staples, continuing the trend
first seen in autumn of a preponderance of fruit and seed in the diet.
The importance value (74.6) for Ashe juniper berries in winter was
the highest importance value of any food item for any season. The
highest seasonal percent volume (88.4%) of a trophic category
(fruit and seed) occurred during winter. Plant matter in the winter
diet was greater than animal matter {X^= 67.2; df = 1; P < 0.01).

The early spring was transitional with a decrease in the use and
importance of the fruit and seed category (32.6% of overall
seasonal percent volume) and an increase in the consumption and

BACCUS ET AL.

107

Table 3. Seasonal overlap in the diet of Peromyscus pect oralis (above diagonal) in centrs
Texas as measured by Morisita’s Index of Similarity and number of food itenr
common to all pair-wise seasonal comparisons (below diagonal).

Season

Winter

Early

Spring

Late

Spring

Summer

Autumn

Winter

0.61

0.367

0.394

0.88

Early Spring

0.688

0.622

0.603

Late Spring

0.657

0.392

Summer

0.418

Autumn

importance of green foliage. Three trophic categories (fruit and
seed, green foliage, and animal matter) comprised the diet.
Residual Ashe juniper berries and hackberries in the environment
(Table 4) and new, succulent foliage of white sweet clover, oneseed
croton, and unidentified dicot herbage were important plant items in
the diet. Larval and adult insect consumption increased (26.7% of
overall seasonal percent volume), but arthropod use ranked lower in
importance than fruit and seed and green foliage (40.7% of overall
seasonal percent volume). Early spring trophic diversity was higher
than winter. Consumption of residual winter berries and new green
foliage resulted in higher plant matter use than animal matter in the
early spring diet (A'^ = 21.67; df = 1; P < 0.01).

A shift in resource use in late spring to consumption of green
foliage and animal matter supplanted fruit and seed use in the diet.
The array of herbaceous plants in the diet expanded; however,
extreme mastication of foliage made identification difficult and
most material was classified as unidentified dicot herbage. Based
on percent frequency, percent volume, and importance value, the
most important food consumed during late spring was larval
insects. Because of high consumption of larval insects, late spring
was the only season where percent volume of animal matter
(57.4%) exceed percent volume of plant matter in the diet. The
most common larval insect in stomachs was the lepidopteran moth
family, Pyralidae. Although there was a higher consumption of
animal matter for this season, animal and plant matter use was
similar (V^= 3.24; df = 1; /> > 0.1).

108

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

□ Fruit and Seed
m Foliage
□Animal Matter

Winter Early Late Summer Autumn
Spring Spring

Season

Figure 2. Seasonal changes in the three major food categories in the diet of Peromyscus
pectoral is in central Texas.

The summer diet was the most heterogeneous of any season.
White-ankled mice continued to eat green herbaceous foliage
(23.6% of overall seasonal percent volume), but percent volume
for most herbs was small. Stem and fruit use of prickly pear was
the highest for any season. However, with maturation of grass and
woody plant seeds, the importance of seeds and fruits in the diet
increased (34.2% of overall seasonal percent volume), but percent
volume for most species was low. Adult insect consumption was
greatest in summer. Arthropods as a trophic group persisted as the
most important food category (41.0% of overall seasonal percent
volume). Adult crickets (Family Gryllidae) and beetles (Family
Scarabaeidae) were the most common insects consumed. The
highest consumption of arachnids occurred during summer.
Trophic diversity (3.74) and richness (15 different food items) were
the highest for any season. Materials attributable to 3 trophic
categories were consumed during this season, and with increased

BACCUS ET AL.

109

Table 4. Seasonal availability of the food resource of Per omy sens pect oralis. Fruits and
seeds are the total number of items counted in ten 0.25 m^ quadrats. Green foliage is
the estimated percent occurrence in one hundred 0.5 m“ quadrats.

Item

Winter

Spring

Summer

Autumn

Fruit and Seed

Jimipertis ashei

222

Rhus Virens

Celtis laevigata

Quercus fusiformis

Sorghum halepense

Cassia Lindheimeriana

Diospyros texana

Green Foliage

Sorghum halepense

Bothriochola ischaemum

Stipa leucotricha

Melilotus albus

Opuntia lindheimeri

Croton monant hog\mus

Shrankia roemeriana

Tor ills nodosa

Lepidium virginicum

Houstonia nigiicans

Lupinus texensis

Rumex acetosella

Abutilon incanum

seed use, there was a preponderance of plant matter in the summer
diet. However, there was no difference between plant and animal
matter use (X^= 2.56; df = 1; P > 0.2).

Autumn was a transitional season because of shifts and changes
in the diet. Fruit and seed consumption (86.2% of the overall
seasonal percent volume) increased and supplanted the importance
of green foliage and arthropods in the diet (Fig. 2). This season was
distinguished by an abrupt increase in the consumption of Ashe
juniper and green sumac berries and prickly pear fruit. Arthropod
use (12.0% of overall seasonal percent volume) decreased sub¬
stantially in comparison to summer. Trophic diversity decreased

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

because fewer food items were consumed. The consumption of
berries and seeds and a diminished use of arthropods resulted in a
diet dominated by plant matter (X^= 56.8; df = 1; P < 0.01).

Peromyscus pectoralis preferred certain foods in different
seasons. Texas persimmon seeds had the highest electivity value
(0.66) for any food item. No Texas persimmon trees occurred
along trap-lines, and few trees were in the immediate study area.
There were, however, an abundance of scattered raccoon {Procyon
lotor) scats with Texas persimmon seeds in the study area. These
scats concentrated Texas persimmon seeds in high density patches
that provided an immediate access to seeds that otherwise were
unavailable.

The most common fruit eaten was Ashe juniper berries. Even
though the ground under and around juniper trees was often
covered by berries, the electivity value (0.12) indicated only a slight
preference for this food item. Since the quantity of Ashe juniper
berries consumed by white-ankled mice was commensurate with
availability (Table 4), P. pectoralis opportunistically fed on these
berries. Hackberry and green sumac berries were not as common as
Ashe juniper berries and availability was lower; however, electivity
indices for these fruits (0.53 and 0.42, respectively) indicated
selection by white-ankled mice. White sweetclover was the most
abundant herbaceous plant in the spring plant availability sample
(28% ground coverage). The electivity index for white sweetclover
foliage (0.13) was similar to Ashe juniper indicating consumption
was commensurate with availability. The negative electivity
indices for seeds and/or foliage of Johnsongrass (-0.93), King
Ranch bluestem grass (-0.96), prickly pear (-0.59), oneseed croton
(-0.91), prairie bluet (-0.95), and Lindheimer senna (-0.99)
indicated avoidance. No stomachs contained live oak acorns,
agarito berries, and seeds or foliage of little bluestem grass,
sensitivebriar, knotted hedgeparsley, pepperweed, Texas
bluebonnet, common sorrel, or Indianmallow.

BACCUS ET AL.

Ill

Discussion

Species of Peromyscus are opportunistic feeders with variable
use of food resources by season and availability. Major trophic
categories in the diet are seeds, fruits, green plants, and animal
matter (Montgomery 1989). Studies of the food habits of P.
maniculatus^ P. leucopus, P. californicus, P. eremicus, P. truei, and
P, boylii (= attwateri) indicated either a moderate to common use of
seeds, rare to common use of animal matter, rare to moderate use of
green vegetation in all species except P. attwateri, and rare to
moderate use of fruits (Brown 1964; Whitaker 1966; Flake 1973;
Kritzman 1974; Vaughan 1974; Meserve 1976; Wolff et al. 1985).

Peromyscus food habits vary from season to season. Wolff et al.
(1985) found univariate differences between seasons for seven of
eight categories of food items in diets of P. maniculatus and P.
leucopus. Both species ate more fleshy fruit in summer, more
moths and butterflies in autumn, and more nuts in autumn and
winter than other seasons. Hamilton (1941) reported that 180 P.
leucopus noveboracensis collected between November and April in
central New York consumed more arthropods than nuts/seeds or
green plant matter. The diet between May and October was
primarily arthropods with lesser amounts of fruits, nuts/seeds, and
fungi. Whitaker (1963) found the primary food in the summer diet
of 142 P. leucopus from New York was nuts/seeds with lesser
amounts of arthropods and green plant matter. In addition,
Whitaker (1966) stated 1 13 P. maniculatus from Indiana consumed
principally nuts/seeds with lesser amounts of arthropods and green
plant matter. Martell and Macauley (1981) found arthropods were
the most common item and nuts/seeds, fruits, and fungi were less
common with green plant matter and achlorophyllous plant matter
present in miniscule amounts in stomach contents of 712 P,
maniculatus taken between May and September in northern
Ontario. Brown (1964) analyzed stomach contents of 20 P.
attwateri collected in March, June, September, and December in
southern Missouri and found seed use highest in June and lowest in
March, insect use highest in December and lowest in June, fruit or

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

berry use highest in March and lowest in June, consumption of
green plant matter highest in June and September and lowest in
March and December, and bulb fragments use highest in September
and lowest in June.

There was a definite seasonal variation in the diet of P.
pectoralis in contrast to the suggestion by Montgomery (1989) that
southern populations of Peromyscus indicate little or no annual
dietary cycle. Dietary trends of this species resemble the diet of
Peromyscus athvateri and other species of Peromyscus; yet, there
are differences. The seasonal diet of P. pectoralis appears
opportunistic, especially in autumn and winter use of Ashe juniper
berries, spring use of green foliage, and spring and summer
consumption of insects. In contrast, plant matter dominated the
winter diet (percent volume 91%), but the percent volume in the
summer diet was only 42.6%. No other studies of Peromyscus have
indicated this high a consumption of plant matter during winter
(Montgomery (1989). Brown (1964) found only low or trace
amounts of green plant matter in the diet of P. attwateri with the
greatest amount of plant matter consumed being seeds.

As in other species of Peromyscus, consumption of animal
matter by the white-ankled mouse had seasonal importance,
especially insect use. Although insect abundance was monitored
using sweeping of vegetation, this method did not provide adequate
samples of the availability of the insects consumed by P. pectoralis
during this study. Most adult insects eaten by P. pectoralis were
ground crickets or beetles. The most consumed larval insects were
catepillars of a pyralid moth that inhabits Ashe juniper trees. Insect
consumption in late spring and summer were the highest reported
for Peromyscus for these seasons (Flake 1973, Kritzman 1974,
M’Closkey and Fieldwick 1975, Batzli 1977). Overall a similar
consumption of animal matter by P. pectoralis and P. attwateri
occurred in spring, summer, and autumn (Brown 1964). Insect use
by P, pectoralis and P. athvateri in early spring and autumn was
comparable, but winter consumption was dissimilar. The disparity

BACCUS ET AL.

113

in insect consumption during winter by P. pectoral is in central
Texas compared to other species of Peromyscus was probaly
phenological. The mild winters of central Texas allows for an
extended period of activity by adult insects compared to those
inhabiting northern latitudes and high altitude environs where
winter temperatures are below freezing for an extended time. In
Hays County, Texas, it is not unusual to have <5 days in winter
with temperatures at or below freezing.

A major difference in the diet of P. pec tor alls compared to other
species of Peromyscus was the importance of fruits in the seasonal
diet. Peromyscus pectoralis began to eat friuts and seeds in
summer as they matured with highest consumption of these foods in
autumn extending into winter. Ashe juniper and green sumac
berries were important components of the autumn through winter
diets when abundance and availabilty was at a maximum in the
habitat of P. pectoralis. Residual hackberries in the environment
were important in the early spring through summer. Prickly pear
fruits became impotant in summer through autumn with maturation
of fruits. All of these berries and fruits became abundant and
important in the diet of P. pectoralis based on their maturation and
abundance in the environment (Table 4). Although the electivity
index for Ashe juniper berries indicated a slight preference by
white-ankled mice, the index value did not indicate the importance
of this food in the diet. The high percent frequency of occurrence
in the environment and percent volume in stomachs confirm the
importance of this food in the seasonal diet and opportunism of P.
pectoralis. Ashe juniper berries were most abundant between 15
December and 25 February. This period of high abundance of Ashe
juniper berries overlapped the time when juvenile mice were most
common in the population. Green sumac berries also had high
abundance during this period. The high use of Ashe juniper and
green sumac berries may have been the result of young white-
ankled mice learning to eat this food in their natal environment and
conditioning of adults to associate with habitats where berries are
abundant (Drickamer 1970; 1976).

114

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

The abundance of green foliage was highest in spring through
autumn for most herbaceous species except Stipa leucotricha that
had highest availability in winter. The phenology for most herba¬
ceous vegetation is invigorated growth during spring (Schmidly
2004). Peromysciis pectoralis had the highest consumption of the
green foliage of herbaceous vegetation in early and late springwhen
availability was highest. When available, green vegetation was a
very common food of P. maniculatiis (Kritzman 1974), moderately
common in the diets of P. californicus and P. truei (Merritt 1974),
but rarely consumed by P. leucopus (Whitaker 1966) and P.
eremicus (Meserve 1976).

In a comparison of the size of food items consumed, larger
berries and seeds had a positive electivity index; whereas, minute
seeds had negative values (i.e., compare hackberry and green sumac
berries vs. Johnsongrass and King Ranch bluestem grass seeds).
Kantak (1983) found northern populations of both wild caught
Peromyscus maniculatus bairdii and Peromyscus leucopus
noveboracensis preferred larger size grass seeds of Andropogon.
Moriarty (1977) suggested that size and selections of food items
were critical expenses related to search time incurred by an animal
while foraging. Both Ashe juniper and sumac berries were readily
available seasonally, and foraging and handling times necessary to
encounter and eat these items were probably minimal (Moriarity
1977). Otherwise, the diversity of food items consumed by P.
pectoralis and the implied trophic niche was similar to other species
of Peromyscus,

In one of the first food habits of a southern species of
Peromyscus, this study delineated by an Index of Change a five
season (winter, early spring, late spring, summer, and autumn)
pattern of trophic ecology for Peromyscus pectoralis in central
Texas. The species had considerable variation in the seasonal diet,
and is primarily a frugivorous/granivorous herbivore with
omnivorous tendencies reflecting opportunistic feeding habits .

BACCUS ET AL.

115

Acknowledgments

Thanks are extended to R. W. Manning, M. F. Small, J. G.
Brant, T. R. Simpson, and an anonymous reviewer for critical
reviews of the manuscript. A special thanks is extended to E.
Longcope for permission to collect on his property. Funding for
this study came from a Texas State University Faculty
Enhancement grant to J. T. Baccus.

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JTB at: jb02@txstate.edu

TEXAS J. OF SCI. 61(2): 119-130

MAY, 2009

CHANGES IN VEGETATION PATTERNS AND THEIR EFFECT ON
TEXAS KANGAROO RATS (DIPODOMYS ELATOR)

Allan D. Nelson, Jim R. Goetze*, Elizabeth Watson
and Mark Nelson

Department of Biological Sciences, Box T-OlOO,

Tar! et on State University, Stephenville, Texas 76401 and
^Science Department, Laredo Community College
Laredo, Texas 78040

Abstract-Investigations of vegetation in Wichita County, Texas indicate that
changes in patterns of grazing and the introduction of non-native plant species may
affect populations of the Texas kangaroo rat. Intensely and moderately grazed areas
were compared to each other and to a previous investigation involving an ungrazed
pasture dominated by introduced Japanese brome (Bromiis japoniciis). Thirty
Dipodomys elator were trapped at the intensely and moderately grazed sites, whereas
only two animals were caught on the periphery of the ungrazed site in Wichita
County. In addition, the moderately grazed site was compared to the intensely grazed
site and no significant differences in vegetative richness or percentages grass and forb
were found between sites. Height of vegetation, percentage bare ground and woody
species coverage were significantly different in comparisons between the two grazed
sites. Because the two sites contained populations of D. elator, it appears that they
can use moderately to heavily grazed habitats as burrow locations and can tolerate
significant differences in vegetation height and amount of bare ground and woody
vegetation. They rarely use ungrazed sites as habitat and, in a previous investigation,
an ungrazed site was significantly different from the grazed sites in vegetational
height, percentage bare ground, and percentage grass coverage. Grazing regimes,
amount of bare ground coverage, and introduction of tall, dense-growing grasses may
be important eonsiderations in managing habitat for Texas kangaroo rats. Moderately
to heavily grazed sites may provide better habitat for these state-threatened mammals.

Dipodomys elator (Merriam 1894) is a state threatened species.
The International Union for the Conservation of Nature (1986) lists
habitat loss and degradation resulting from agricultural and
infrastructure development as major threats. Though degradation
such as fragmentation and loss of habitat have played important
roles, changes in vegetation patterns may also be important. Much
historic range of the Texas kangaroo rat has been fragmented by
extensive cultivation within the Rolling Plains Region of Texas and
adjacent regions of Oklahoma (Correll & Johnston 1970).

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Cultivation fragmented the grasslands and only those areas
unsuitable for cultivation were left in their natural state. These
fragments were fenced by ranchers and grazed by cattle and
provided some habitat for the Texas kangaroo rat. However,
because of a decline in ranching, some of these pasturelands are no
longer grazed and have been invaded by introduced species such as
Japanese brome. Additionally, lack of fire has allowed the increase
of woody vegetation such as honey mesquite (Prosopis glandulosa)
and lotebush (Ziziphus obtiisifolia). In some cases, Texas kangaroo
rats used these woody species as burrow sites because the plants
collected wind-blown soil and the Texas kangaroo rats dig burrows
at their bases. However, as mesquites mature, their shade changes
the vegetation composition sometimes favoring introduced grasses
like Japanese brome, which grows densely and changes the habitat
so that it is more suitable for other types of small mammals.

Mesquite forest was not seen as a problem by the Texas Parks
and Wildlife Department (TPWD) because of the assumption that
this habitat was required for Texas kangaroo rats. However,
perceptions regarding threats to the species and ideas about future
management have changed. Research has suggested that (1)
mesquite is not a critical component of D. elator habitat (Stangl et
al. 1992; Goetze et al. 2007), (2) grazing may benefit D. elator
(Stangl et al. 1992; Stasey 2005; Goetze et al. 2007), and (3) Texas
kangaroo rats opportunistically use human structures that collect
friable soils as burrow sites (Stangl et al. 1992; Stasey 2005; Goetze
et al. 2007). Schmidly (2004), which is used by TPWD as their
main source of small mammal data for the state, states that heavily
grazed rangeland and the eroded sites of rangeland roadways may
provide optimum habitat. NatureServe (2008) states that vegetation
has become overgrown and that rangeland practices that result in
dense growth of grasses or invasion of non-native grasses have
degraded habitat because the Texas kangaroo rat thrives in heavily
grazed or otherwise disturbed conditions. NatureServe (2008)
further states that habitat for D. elator consists of sparsely vegetated
areas that may or may not include honey mesquite, including

NELSON ET AL.

121

heavily grazed land, disturbed areas, and areas along fencerows
adjacent to cultivated fields and roads. These current statements are
quite different from Davis & Schmidly (1994), who stated that
Texas kangaroo rat burrows invariably entered the ground at the
base of a mesquite and the primary threat contributing to the rarity
of the species was the clearing of mesquite brush.

Currently, the only relatively large populations of Texas
kangaroo rats known in Wichita County occur in pastures with
small or scattered mesquite, and burrows are often not associated
with mesquite at all, but rather with lotebush, prairie mounds
(natural, elevated, and relatively bare areas possibly uplifted by clay
soils swelling in cracks; Diggs et al. 1999), or in areas where man¬
made berms occur due to road, fence, and oilfield construction, or
in association with old (>30 years), unbumed brush piles where
wood has decayed leaving a mound of loose friable soil (Stangl et
al. 1992; Goetze et al. 2007). Stangl et al. (1992) hypothesized that
grazing bison and prairie dogs, along with fire, historically
maintained the type of disturbances needed by the Texas kangaroo
rat. Also, prolonged drought likely played an important role in fire
frequency and maintaining short vegetation with intermittent bare
patches of soil. Natural prairie heterogeneity such as prairie
mounds (Diggs et al. 1999; Goetze et al. 2007) appear to be
important in providing the type of habitat needed by the Texas
kangaroo rat before cattle grazing and human mediated
disturbances were used opportunistically as burrow sites (Stangl et
al. 1992). In areas where cattle no longer graze or at sites where
native vegetation has been replaced by introduced species, it
appears that populations of Texas kangaroo rats have declined.

Ecological characterization of burrows in situations that lacked
grazing as a component are rare (Martin & Matocha 1992; Stasey
2005). Martin & Matocha (1992) trapped for 10 trap nights and
characterized a burrow where a single Texas kangaroo rat was
trapped in association with a fence row adjacent to a gravel county
road in Motley County, Texas. No other mammals were captured.
This capture site contained vegetation characteristic of a disturbed

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

site and was adjacent to a field of Sudan grass. At the capture site
there was 30.2% bare ground, 65% grasses and 4.8% forb (Martin
& Matocha 1992).

Stasey (2005) trapped for 972 trap nights in a mesquite forest
with an understory dominated by Japanese brome. His site was
within 4.0 km of a large and persistent Texas kangaroo rat
population in Wichita County where the grazing regime is heavy
(Goetze et al. 2007). During his investigation, he caught only two
kangaroo rats on the periphery of the mesquite forest habitat. One
was caught in a friable clay soil that had blown in and accumulated
around the comer posts that supported the gate leading into the
pasture and the other was captured along the fence separating this
pasture from a wheat field where a berm had accumulated due to
plowing next to the fence line. Stasey (2005) caught Sigmodon
hispidus, Peromysciis leucopus, and P. maniculatiis in the core of
the ungrazed site. In the core of the ungrazed pastureland, where
no Texas kangaroo rats were captured, seven quadrats were
sampled to assess vegetation characteristics. Percent bare ground
had a mean of 10.9%, grasses 63.7%, and forbs 16.1%, whereas the
mean average herbaceous height was 49.0 cm (Stasey et al. 2005).

Because identifying habitat critical to the survival of the species
is a research priority (Jones et al. 1988), the purpose of this
investigation was to compare a moderately grazed site to that of a
heavily grazed site, both of which have populations of kangaroo
rats. These data were then compared to data from ungrazed sites
dominated by Japanese brome (Stasey 2005) and vegetation
associated with a disturbed roadside (Martin & Matocha 1992).

Materials and Methods

The study area is in Wichita County on the east side of the
intersection of highways 1739 and 2384 and is a privately owned
ranch that is moderately grazed pasture (0.30 head per ha).
Coordinates at the entrance of the ranch are 34.05423 N, 98.81721
W. The pastureland is fenced, has small mesquite (less than 2 m. in
height), and has several old oil field storage sites. Mesquite density

NELSON ET AL.

123

at the site is 168/ha. This site was compared to a nearby population
of Texas kangaroo rat known in Wichita County and its locality and
history have previously been described (Stangl et al. 1992; Goetze
et al. 2007). Grazing at this site is intense (0.81 head/ha) and it has
about 54 small (less than 2m in height with most under 1 m)
mesquite/ha (Goetze et al. 2007).

All trapping was done using 7.5 by 8.8 by 30 cm Sherman traps
with rolled oats as bait. Traps were set just before dark and
checked early the next day. Based on parameters set by Stangl et
al. (1992) and Stasey (2005) regarding burrow entrance diameter,
angle of entry, and vegetation, suspected burrows were selected at
the sites and three traps were placed around each burrow entrance.

All vegetation data was quantified in May so that direct
comparisons could be made, thus eliminating seasonal vegetative
changes. For burrows where at least one Texas kangaroo rat was
caught, one square meter quadrats were centered around burrow
entrances and percentage cover, grass, forb, bare ground, and
woody vegetation (when present) was recorded (Goetze et al.
2007). Vegetative richness and height were measured and the
dominant grass, forb, and woody species of each quadrat were
identified (Goetze et al. 2007). Quantitative data was compared
using SPSS 14.0 (SPSS, Inc. 2005). A Mann Whitney test was
used to test for significant differences in richness and percentages
of grasses, forbs, bare ground, and woody vegetation between the
two grazed sites. In addition, data from a previous study (Stasey
2005) at a site that was ungrazed was included in a Kruskal-Wallis
analysis to test for significant differences between the grazed and
ungrazed sites.

Dominant vegetation at the site was identified using floras for
the state and for north central Texas (Correll & Johnson 1970;
Diggs et al. 1999). Voucher specimens are deposited in the
Tarleton State University Herbarium (TAG).

The specific location of each burrow was recorded in decimal
degrees using a Garmin GPS- 12 unit and burrows were classified as

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

being associated with human-mediated disturbances such as old
brush piles and fence rows, or other available natural habitats that
included prairie mounds, which are elevated, open areas formed by
clay soil shrinkage and swelling (Diggs et al. 1999), or
accumulation of soil at the base of lotebushes or honey mesquites
(Goetze et al. 2007; Table 1).

Results

At the moderately grazed site, three traps were placed around
each of 12 burrows which resulted in the capture of 10 D. elator.
No other rodents were captured. Three of the animals were caught
in each of the traps placed at one burrow and two were caught in
the three traps surrounding another burrow site. The dominant
grass at five of the seven burrows was little barley (Hordeum
pusilhim) with buffalograss (Buchloe dactyloides) and rescue grass
{Bromus catharticus) being dominant at the other two burrows
(Table 1). The dominant forb at six of the seven burrows was
Texas broom weed {Gutierrezia texana) with Virginia pepperweed
(Lepidium virginicum) being dominat at the other burrow (Table 1).
Five of the seven quadrats had honey mesquite (Prosopis
glandulosa) as the dominant woody vegetation and two contained
no woody vegetation (Table 1). Two burrows were associated with
fence rows and five with honey mesquite (Table 1).

At the heavily grazed site, three traps were placed around each
of 22 burrows which resulted in the capture of 1 8 D. elator. Of
these, 10 burrow sites were analyzed for vegetation and burrow
associations. At the heavily grazed site, little barley was always the
dominant grass and most quadrats contained Virginia pepperweed
as the dominant forb (Table 1). Other herbaceous dominants
included Texas broomweed, hog potato {Hoffmannseggia glaiica),
and western ragweed {Ambrosia psilostachya) (Table 1). Woody
vegetation was evenly distributed between lotebush and honey
mesquite. At the heavily grazed site, five burrows were associated
with old brush piles, two with prairie mounds, one with a fence
row, and one each with lotebush and honey mesquite (Table 1).

NELSON ET AL.

125

Table 1. Dominant vegetation and burrow classifications at the two study sites.
Dominant forbs are broomweed {Gutierrezia texana), hog potato {Hoffmanmegia
glauca), pepperweed (Lepidhim virginicum), and ragweed {Ambrosia psilostachy^a).
Dominant grasses are barley {Hordeum pmillum), buffalo grass {Biichloe
dactyloides), and rescue grass {Bromiis catharticm). Woody vegetation includes
lotebush {Zizyphus obtiisifolia) and mesquite (Prosopis glandiilosa). Biurow
classifications included in this table are defined in the methods section of this paper.

Moderately grazed burrows:

Burrow #

FORBS

Broomweed

Pepperweed

GRASS

Barley

Buffalo

Rescue

WOODY VEG.

Mesquite

None

Burrow assoc.

Fence

Mesquite

Heavily grazed burrows:

Burrow #

7 8 9 10

FORBS

Broomweed

Hog potato

Pepperweed

Ragweed

Unknown

GRASS

Barley

WOODY VEG.
Lotebush
Mesquite
None

Burrow assoc.
Fence
Lotebush
Mesquite
Old brush pile
Prairie mound

XXX

X X

X X X X X X

X X

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

Average herbaceous height, percentage bare ground, and
percentage woody vegetation were significantly different between
heavily and moderately grazed sites (Table 2). Percentage forbs,
grasses, and other categories such as rocks and stumps, as well as
richness were not significantly different between sites (Table 2).

Average herbaceous height (P = 0.001), percentage bare ground
(P = 0.009) and grasses (P = 0.009) were significantly different
when comparing the moderately and heavily grazed sites as well as
the ungrazed site examined by Stasey (2005).

Discussion

Little barley, Texas broomweed, hog potato, Virginia
pepperweed, and western ragweed occur in disturbed habitats
(Diggs et al. 1999). These plants were dominant species associated
with burrows of Texas kangaroo rats and their occurrence is likely
caused by disturbances such as grazing of cattle and rodent activity
around the burrows. Habitat of D. elator was dominated by short,
herbaceous vegetation (2.0 - 40.0 cm in height). There is general
agreement that D. elator requires a sparse, short-grassland habitat
(Dalquest & Collier 1964; Roberts & Packard 1973; Carter et al.
1985; Stangl et al. 1992), and findings from this current study
support this conclusion. These findings also indicate that grazing
may be important in maintaining sparse, short grassland habitat.
When comparing two grazed sites and an ungrazed site (Stasey
2005), the only significant differences were in average herbaceous
vegetation height and percentage bare ground and grass. Grazing
can change these three parameters, which appear to be important in
maintaining Texas kangaroo rat habitat. As previously discussed,
this is complicated by the dominance of the introduced grass,
Bromus japonicas at the ungrazed site and additional studies need
to be conducted at ungrazed sites containing native vegetation.
However, based on the lack of D. elator at this site and the relative
abundance at the grazed sites, it appears that grazing plays a role in
maintaining suitable habitat for Texas kangaroo rats. Lack of
grazing significantly increased vegetation height at the ungrazed

NELSON ET AL.

127

Table 2. Comparison of heavily and moderately grazed sites for average herbaceous
height, percentage coverage of bare ground, forbs, grasses, woody, other (rocks,
stumps, posts, etc.), and riclmess. Comparisons are made using means, standard
deviations (in parentheses), and ranges [in brackets] and evaluated using Wilcoxon
Mann- Whitney test. Significant differences at P < 0.05 are denoted by an asterick.

Heavily Grazed

Moderately Grazed

P-value

Avg. Herb. Height

7.1 (±7.9) [2.0-40.0]

24.5 (±12.7) [9.0-29.4]

0.0001*

% Bare Ground

49.9 (±24.0) [0.0-80.0]

20.7 (±18.1) [5.0-60.0]

0.024*

% Forbs

16.5 (±13.1) [1.0-35.0]

33.7 (±22.6) [15.0-67.0]

0.133

% Grasses

24.60 (±18.9) [1.0-55.0]

20.1 (±15.6) [1.0-45.0]

0.623

% Woody

6.0 (±15.8) [0.0-50.0]

26.1 (±26.5) [0.0-60.0]

0.037*

% Other

2.0 (±4.2) [0.0-2.0]

0.0 (±0.0) [O.O-O.O]

0.222

Riclmess

6.2 (±2.4) [3.0-10.0]

7.6 (±1.8) [6.0-10.0]

0.137

site, which on average was double the moderately grazed site and
seven times greater than the heavily grazed site. Percentage of
grasses was about three times greater on average at the ungrazed
site when compared to grazed sites. Percentage bare ground, on
average was about one-half that of the moderately grazed site and
about one-fourth that of the heavily grazed site. This tall, dense
coverage by grasses may impede Texas kangaroo rat movement,
inhibit their ability to see potential predators, and may make burrow
construction difficult. Lack of bare ground likely inhibits their dust
bathing activities.

Burrows at the grazed site compare favorably to the burrow
ecological characteristics reported from Motley County (Martin &
Matocha 1992). In the classification system used for burrows and
described in the Materials and Methods, this burrow would have
been a fence line association and the animals were likely using soil
that accumulated at the base of the fence. Although no grazing was
reported at this site, the value reported for bare ground percentage
(30.2%) compares favorably with the grazed sites in this
investigation but forb percentage (4.8%) was low and grass
percentage was high (65%). The location of this burrow in a fence
row berm may have provided the friable soil preferred for burrow
development. Its location at the edge of a sudan field adjacent to a
gravel road (Martin & Matocha 1992) may have provided enough

128

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

disturbance to maintain bare patches for dust bathing and for
foraging trails that still allow the rats to spot predators.

Of the 17 burrows examined in this investigation, 47% were
associated with human disturbances including old brush piles and
fence rows. The other burrows were associated with more natural
sites such as shrubs and prairie mounds. In the heavily grazed site
most burrows were associated with 30 year old brush piles, while at
the moderately grazed site most were associated with honey
mesquite. Second in number of burrow associations at the heavily
grazed site was prairie mounds and none of this association was
observed at the moderately grazed site. Heavy grazing might make
prairie mounds more suitable as burrow sites, since that was the
only place this habitat association occurred. At the heavily grazed
site, lotebush and honey mesquite associations were equal in
number while no lotebush associations were observed at the
moderately grazed site. This is likely because the moderately
grazed site had woody vegetation dominated by honey mesquite
whereas the heavily grazed site had some lotebush available. The
type of shrub is probably not as important as is the accumulation of
loose, friable soil at the base of the shrubs.

Extrinsic disturbance caused by grazing, fire, or drought, and
natural landscape heterogeneity such as prairie mounds probably is
important for burrow site selection in Texas kangaroo rats. The
slight elevation of prairie mounds may provide more bare ground
because of a drier microclimate and better drainage. Also, the
animals can dig their more characteristically horizontal openings
into the sides of these mounds. Opportunistic use of any natural or
manmade disturbance where friable soil accumulates such as
around shrubs, the bases of rocks, fence lines, cattle pens, pasture
and oil field roads, abandoned equipment, old, unbumed brush piles
with most of the wood decayed have been observed (Stangl et al.
1992; Goetze et al. 2007). Almost one-half of the burrows in this
investigation may be characterized as such, supporting hypotheses
of opportunistic use made by others (Stangl et al. 1992; Stasey

NELSON ET AL.

129

2005; Goetze et al. 2007). These manmade disturbances likely
mimic natural prairie heterogeneity.

Investigations of Texas kangaroo rats that examine symbiosis
with prairie dogs, effects of fire in maintaining habitat, the role of
drought on habitat, and additional research into the influence of
natural prairie heterogeneity and grazing regimes are critical for
understanding the animal’s niche. Surveys need to be conducted at
Buffalo Creek Reservoir and Lake Arrowhead State Park to
ascertain if D, elator occurs in any protected natural areas (pnas) in
Wichita County or if not, if suitable habitat is available in protected
natural areas. Also, based on the results of this investigation,
managers of pnas may need to consider grazing as a management
practice to promote the development of habitat for the Texas
kangaroo rat and other organisms that require grazing as a
disturbance.

Acknowledgments

We would like to thank Oscar and Edith Goetze for allowing
access to their properties in Wichita County and for room and board
while conducting fieldwork. Tarleton State University Organized
Faculty Research provided funding for parts of this project. This
study was conducted under Texas Parks and Wildlife permit SPR-
0496-775.

Literature Cited

Carter, D. C., W. D. Webster, J. K. Jones, Jr., C. Jones & R. D, Suttkus. 1985.

Dipodomys elator. Mammalian Species, 232: 1-3.

Correll, D, S. & M. C. Johnston. 1970. Manual of the vascular plants of Texas.

Texas Research Foundation. Renner, Texas. 1083 pp.

Dalquest, W. W. & G. Collier. 1964. Notes on Dipodomys elator, a rare kangaroo
rat. Southwestern Nat, 9:146-150.

Davis, W. B. & D. J. Schmidly. 1994. The mammals of Texas. Texas Parks and
Wildlife Press. Austin, TX, 338 pp.

Diggs, G. M., B. L. Lipscomb & R. J. O’Kennon. 1999. Shinners & Mahler’s
Illustrated Flora of North Central Texas. Botanical Research Institute of Texas.
Fort Worth, Texas, 1626 pp.

130

THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 2, 2009

Goetze J. R, W. C. Stasey, A. D. Nelson, & P. D. Sudman. 2007. Habitat attributes
and population size of Texas kangaroo rats on an intensely grazed pasture in
Wichita County, Texas. The Texas J. of Sci., 59(1):1 1-22.

International Union for Consei*vation of Nature and Natural Resources. 1986. 1986
lUCN red List of threatened animals. lUCN. Cambridge, U. K., 105 pp.

Jones, C., M. A. Bogan & L. M. Mount. 1988. Status of the Texas kangaroo rat
{Dipodomys elator). The Texas J. of Sci., 40(3):249-258.

Martin, R. E. & K. G. Matocha. 1991. The Texas kangaroo rat, Dipodomys elator,
from Motley County, Texas, with notes on habitat attributes. Southwestern Nat.,
36:354-356.

Merriam, C. H. 1894. Preliminaiy descriptions of eleven new kangaroo rats of the
genera Dipodomys and Perodipus. Proc. Biol. Soc. Washington, 9:109-1 16.

NatureServe. 2008. NatureServe Explorer: An online encyclopedia of life [web
application]. Version 7.0. NatureServe, Arlington, Virginia. Available
http://www.natureserve.org/explorer. (Accessed: September 4, 2008 ).

Roberts, J. D. & R. L. Packard. 1973. Comments on movements, home range and
ecology of the Texas kangaroo rat, Dipodomys elator Merriam. J. Mamm.,
54:957-962.

Schmidly, D. J. 2004. The mammals of Texas, revised edition. University of Texas
Press, Austin, Texas, 501 pp.

Stangl, F. B., Jr., T. S. Schafer, J. R. Goetze & W. Pinchak. 1992. Opportunistic use
of modified and disturbed habitat by the Texas kangaroo rat {Dipodomys elator).
The Texas J. of Sci., 44(l):25-35.

Stasey, W. C. 2005. An evaluation of Texas kangaroo rat {Dipodomys elator):
Biological habits and population estimation. Unpublished Masters Thesis.
Tarleton State University, 45 pp.

SPSS, Inc. SPSS 14.0 brief guide. 2005. Prentice Hall. Upper Saddle River, New
Jersey, 245 pp.

ADN at: nelson@tarleton.edu

TEXAS J. OF SCI. 61(2): 131-146

MAY, 2009

BREEDING BIOLOGY OF THE

BARN SWALLOW (HIRUNDO RUSTICA) IN NORTHEAST TEXAS
WITH TEMPORAL AND GEOGRAPHIC COMPARISONS TO
OTHER NORTH AMERICAN STUDIES

K. T. Turner and J. G. Kopachena

Department of Biological and Environmental Sciences
Texas A&M University-Commerce, Commerce, Texas 75429

Abstract.-The reproductive output of American bam swallows (Hirimdo nistica
erythrogaster) was studied over a five year period at a colony in northeast Texas.
Clutch size, hatching success, nestling success, and nesting success were all found to
vary significantly among years suggesting that the swallows were influenced by
annual changes in enviromnental conditions. However, the observed annual
variations in clutch size, hatching success, and nesting success were well within the
range observed among 14 other studies in North America. The greatest sources of
mortality were hatch failure, followed by infanticide and ectoparasites. These
sources of mortality were also common in other studies. Despite the wide geographic
range of bam swallows in North America, there were no latitudinal trends in clutch
size, hatching success, or nesting success. Lastly, in the 44 years spanned by the
analysis, there was no evidence of changes in reproductive output, suggesting that
American bam swallows have not yet been obviously impacted by global climate
change.

Barn swallows (Hirundo nistica) have been well studied,
especially in Europe. They have been used to study the adaptive
value of colonial nesting (Snapp 1976; Shields & Crook 1987), the
effects of parasitism (Saino et al. 1998; Barclay 1988; Kopachena et
al. 2000; Merino et al. 2000; Moller 2000), sexual selection (Saino
& Moller 1996; Kose et al. 1999; Perrier et al. 2002; Saino et al.

2002) and immunocompetence (Saino & Moller 1996; Saino et al.
1997; Merino et al. 2000; Hasselquist et al. 2001; Saino et al.

2003) . In North America studies on the breeding biology of barn
swallows have been conducted in West Virginia (Samuel 1971),
New York (Ramstack et al. 1998), and Kansas (Thompson 1961;
Anthony & Ely 1976). Two studies on breeding biology were
performed in Texas, one in Brazos County (Barr 1979) and one in
south central Texas (Martin 1974). A third study in Texas examined
the effect of a selenium-contaminated lake on the reproductive

132

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

success of bam swallows in Rusk County (King et al. 1994).

Because of their broad geographical distribution and ease of
study, bam swallows should make a good model species to study
temporal and geographic patterns in reproductive success and
behavior. Such studies can be useful in elucidating how species
with broad distributions can adapt to variations in climate and
resource availability.

This study documented the breeding biology of bam swallows at
a colony in northeast Texas for a five-year period. The data are
used to provide baseline data on this species in this region and to
document temporal variation in breeding parameters across years.
These data are used to compare with breeding parameters described
in the literature from other locations in North America. Lastly, a
comparison of temporal and spatial variation is used to determine if
there are geographic trends in breeding output.

Materials and Methods

During the breeding seasons of 1998 through 2002, 369 nests
were studied at a colony near Commerce, Texas. The nests were
located under two parallel two-lane bridges located 2.3 km south of
the junction of State Highway 50 and Loop 178. The nests were
checked at least once a day from April 1 through July 31 of each
year. In 1998, nests were checked between 14:00h and 17:00h
CDT. For the remaining years, the nests were checked between
08:00h and 10:30h CDT. Nest initiation date was defined as the
date on which an egg first appeared in a nest. Each nest was given
a unique number upon initiation.

Nestlings in each nest were marked with a unique combination
of toenail clippings on the day of hatching and again on the sixth
day post-hatching (St. Louis et al. 1989). Nests were checked daily
until fledging. If a nestling died, the cause of death, where
possible, was noted. Causes of death were categorized as follows.
Eggs that failed to hatch were noted as failed to hatch. Barn

TURNER & KOPACHENA

133

swallows in both Europe and North America practice infanticide
(Crook & Shields 1985; Moller 1994). In the current study,
infanticide was identified by the presence of eggs on the ground
below the nest or the presence of chicks on the ground below the
nest. In the latter case, most chicks were dead or dying and usually
had evidence of trauma on the abdomen. Chicks found on the
ground that were over 12 days of age and which were otherwise
healthy were considered to have fledged early. Infanticide was
stratified into infanticide during the incubation period and
infanticide during the nestling stage. Many nests were infected
with swallow bugs {Oeciaciis vicarius), tropical fowl mites
{Ornithonyssus bursa), or both. In heavily infested nests the chicks
were noticeably exsanguinated and jaundiced. Deaths that occurred
in these nests were attributed to the parasites. In some nests,
nestlings showed signs of decline in the absence of obvious parasite
infections, while in other cases nestlings died shortly after hatching
without apparent cause. In either case the cause of death was
assigned as unknown. Similarly, nestlings that disappeared prior to
the twelfth day were assumed to have died from unknown causes.

Each nestling’s mass and right primary feather length was
measured at twelve days post hatching. A 50g spring scale was
used to measure nestling mass to the nearest O.lg. Digital calipers
were used to measure the primary feather length to the nearest
0.01mm. Clutch size, brood size at hatch, brood size on day 12,
and per capita mortality rate for whole broods were calculated.

The temporal pattern of nests containing eggs showed a
distinctly bimodal distribution for all years, indicating that the
majority of pairs were double-brooded. Since adult pairs were not
marked, this distribution was used to define first and second broods.

To compare the data collected in northeast Texas with other
studies a literature survey was conducted. Data using the same
measures of mean clutch size, hatching success, and fledging
success as used in the current study were obtained from 14 studies

134

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

in which location and year of study were provided. For each study,
the latitude of the study area was recorded. Along with the current
study, data spanning 44 years and over 19® of latitude were
collected. These data were used to examine the extent of variation
in reproductive output among studies and to determine whether
reproductive output has changed over time or varies relative to
latitude.

Statistical analyses were done using SAS (Version 9.1). Where
parameters were not normally distributed, non-parametric statistics
were used.

Results

In all years the modal clutch size was five (Fig. 1). However,
the average clutch size varied among years, being highest in 1999
and lowest in 1998 (Kruskal- Wallis Test, Chi-Square Approxi¬
mation, X^= 11.44, df = 4, p = 0.0220). Clutch size also varied
between first and second broods. Five-egg clutches were most
common in first broods, whereas four-egg clutches were more
common in second broods (Fig. 2) (Chi-Square, 2 by 2 Contingency
Table, = 29.91, df = \, p < 0.0001). Clutch size varied
significantly among years for first broods (Kruskal- Wallis Test,
Chi-Square Approximation, X^= 17.36, df^ 4, /? = 0.0016), but not
for second broods (Kruskal-Wallis Test, Chi-Square
Approximation, 3.81, df= 4,p = 0.4317).

Hatching success for all years combined is summarized in Table 1.
It was measured as the percentage of eggs that hatched and as the
percentage of nests experiencing complete hatch failure (whole
clutch loss. Table 1). The percentage of eggs hatching varied
significantly among years (Chi-Square, 2 by 5 Contingency Table,
X^ = 23.20, df = 4, p < 0.0001) whereas there was little annual
variation in the number of whole clutches lost (Chi-Square, 2 by 5
Contingency Table, X^ = 5.96, df= 4, /7 = 0.2022). The proportion
of eggs hatching did not differ between first and second broods

TURNER & KOPACHENA

135

Clutch Size

Figure 1. Distribution of clutch sizes for each year from 1998 through 2002. Number of
clutches monitored each year were as follows: 1998 = 86, 1999 = 105, 2000 = 42,
2001 =55, 2002 = 81.

Clutch Size

Figure 2. Distribution of clutch sizes for first and second broods.

Table 1. Breeding success of bam swallows at a colony in northeast Texas from 1998 tln ough 2002. Means are expressed as
mean + standard deviation. Numbers in parentheses are sample sizes.

136

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

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TURNER & KOPACHENA

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(Chi”Square, 2 by 2 Contingency Table, = 1.60, df = p =
0.2059). Similarly, there was no statistical difference between first
and second broods relative to whole clutch losses (Chi-Square, 2 by
2 Contingency Table, 1.68, df= \,p = 0,1949).

Nestling success for all years combined is also summarized in
Table 1. It was measured as the percent of nestlings that hatched
and survived to day twelve and by the percent of nests with eggs
that hatched that experienced whole brood loss during the nestling
period. Survivorship of nestlings varied significantly from year to
year for second broods (Chi-Square, 2 by 5 Contingency Table,

= 21.95, df= A,p = 0.0002), but not for first broods (Chi-Square, 2
by 5 Contingency Table, X^ = 7.53, df = 4, p = 0.1104). First
broods had higher survival than second broods (Chi-Square, 2 by 2
Contingency Table, X^ = 21.88, df ^ 1, /? < 0.0001). However,
during the nestling period, whole brood losses did not differ
significantly between first and second broods (Chi-Square, 2 by 2
Contingency Table, X^ = 0.03, df= \,p = 0.8625) or among years
(Chi-Square, 2 by 5 Contingency Table, X^ = 6.74, df = 4, p =
0.1503).

Total nesting success (nesting success) was determined as the
number of eggs laid that resulted in chicks surviving to day twelve
(Table 1). Percent survival of nestlings until day 12 varied
significantly among years (Chi-Square, 2 by 5 Contingency Table,
X^== 30.92, df= 4,p < 0.0001) and was higher for first broods than
for second broods (Chi-Square, 2 by 2 Contingency Table, X^ =
5.80, df= 4,p = 0.016). The percentage of whole brood loss varied
significantly among years (Chi-Square, 2 by 5 Contingency Table,
X^ = 13.93, df= 4, p = 0.0075), but did not differ overall between
first and second broods (Chi-Square, 2 by 2 Contingency Table, X^
= 0.99,#= 1,/? = 0,3197).

Measurements of mass and primary feather length were taken on
the 12^^ day after hatching (Table 1). First broods generally had
higher mean mass than did second broods (Wilcoxon Two-Sample

138

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

Failed to Infanticide Infanticide Parasites Unknown
Hatch Eggs Nestlings

Figure 3. Causes of mortality for all cases observed from 1998 through 2002. For more
detail on causes of mortality, see text.

Test, Normal Approximation, -6.9451, n = 162, 110, p < 0.0001)
and mass also varied among years for both first (Kruskal-Wallis
Test, Chi-Square Approximation, = 21.91, df= 4, p = 0.0002)
and second broods (Kruskal-Wallis Test, Chi-Square
Approximation, X^= 33.32, df= 4,p< 0.0001). On the other hand,
there was no difference in primary feather length among years
(Kruskal-Wallis Test, Chi-Square Approximation, = 5.12, df= 4,
p = 0.2748) or between first and second broods (Wilcoxon Two-
Sample Test, Normal Approximation, 1.5635, 162, 110, =

0.1179).

For both first and second broods the most common source of
mortality was hatch failure; more than a third of all mortalities were
hatch failures (Fig. 3). However, with the exception of infanticide
during the nestling period, the frequency for all causes of mortality
varied significantly between first broods and second broods (Chi-
Square, 2 by 5 Contingency Table, = 61.11, df= 4, p < 0.0001)
(Fig. 3). Hatch failure, infanticide during incubation, and mortality

TURNER & KOPACHENA

139

due to parasites were all higher for first broods than for second
broods. The number of unknown mortalities was higher for second
broods than for first broods.

The values for clutch size^ hatching success, and fledging
success (mean number of chicks surviving to day 12, Table 1) were
well within the range of values obtained in the 14 studies used for
comparison (Table 2). The geographic variance in clutch size,
hatching success, and fledging success observed among these
studies did not differ from the among year variance of these same
measures observed in this study (Folded F-test for Equality of
Variance; clutch size, /= 2.42, df= 12, 4, /? = 0.4073; hatching
success, /= 2.79, df= 4,7,p = 0.2244; fledging success, /= 2.49, df
= 8, 4,/? = 0.3954).

The data were used to determine if reproductive success varied
relative to latitude or the year of study. There was no statistical
correlation between clutch size, hatching success, or fledging
success and latitude (Pearson’s r: clutch size, r = 0.35, n= 14, /? =
0.2201; hatching success, r = 0.157, = 9, /? = 0.2163; fledging

success, r = 0.046, n = 10, /? = 0.8991). Similarly, there was no
significant correlation between clutch size, hatching success, or
fledging success and year of study (Pearson’s r: clutch size, r =
0.199, n = 14, jf? = 0.4955; hatching success, r = -0.133, n = 9, p =
0.7330; fledging success, r = -0.320, n=\0,p = 0.3667).

Discussion

Clutch size, hatching success, nestling success and total nesting
success of barn swallows at the Texas colony were found to vary
considerably between years. This likely reflects variations in
resource availability, which, in turn, is a product of variations in
climatic variables (Stenseth et al. 2002). In northeast Texas,
temperatures are lower and precipitation is higher in May and June
than in July and August. Such conditions are likely to contribute to
higher nesting success in first broods than in second broods.
Furthermore, as conditions become hotter and drier in July and

Table 2. Breeding data collected from other studies conducted in North America.

140

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

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TURNER & KOPACHENA

141

August, the annual reproductive output in bam swallows becomes
more variable, suggesting that the birds are more sensitive at this
time to fluctuations in resource availability. The tendency for first
broods to do better than second broods is a trend that is common in
temperate zone nesting birds (Perrins 1970). However, among bam
swallows deviations from this pattern have been documented in
Kansas (Anthony & Ely 1976) and West Virginia (Samuel 1971)

Measures of clutch size, hatching success, and fledging success
also varied considerably among the geographic locations
documented in the literature. However, the variation among
geographic locations did not differ from the variation among years
in the current study. Given that there was also no trend in breeding
output relative to latitude or date, and given that most of the cited
studies were short term (one or two years), it seems unlikely that
there is substantial difference among geographic localities in
overall, long term reproductive performance among bam swallow
populations in North America.

Hatch failure was the single greatest source of mortality in the
current study, followed closely by infanticide during the egg and
nestling period. Hatch failure is apparently rather common in many
bam swallow populations (Brown & Brown 1999) and appears to
be mainly due to infertility (Moller 1994). Infertility was likely the
main source of hatch failure in the current study as well, since
unhatched eggs seldom showed signs of development when they
were examined.

Infanticide was cited as the major source of nestling mortality in
Denmark (Moller 1994) and as the second most frequent cause of
mortality in New York (Sheilds & Crook 1987). Other studies in
North America, make no mention of infanticide. There are two
possible reasons for this. First, both the Sheilds & Crook (1987)
study and the current study were conducted on large colonies and
the rate of infanticide increases with colony size and population
density (Moller 2004). If other studies were conducted at much

142

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

smaller colonies, then infanticide may be less common and less
easily detected. Secondly, the other studies may not have
considered the possibility that bam swallows exercise infanticide.
In these cases, cases of infanticide may have simply been
categorized as egg loss or nestling loss. Infanticide in North
American populations of bam swallows remains poorly
documented and is in need of further study.

Bam swallows are well known to harbor a variety of
ectoparasites (Brown & Brown 1999). Ectoparasites were con¬
sidered to be the single greatest source of mortality in New York
(Shield & Crook 1987) and in Manitoba (Barclay 1988) and this
may be tme in many other populations as well (Brown & Brown
1999). However, the types of ectoparasites seem to vary
considerably from one area to another. For example, the sole
ectoparasite observed associated with bam swallow nests in
Manitoba were hematophagous mites (Barclay 1988). In New
York, nests were infected with blow flies (Protocalliphora sp.). In
the current study, bam swallow nests were found to be infected
with both tropical fowl mites {Ornithonyssus bursa) and swallow
bugs (Oeciacus vicarius). Swallow bugs have apparently spread
from associations with cliff swallows and are now widespread in
bam swallow colonies in northeast Texas (Kopachena et al. 2007).

Interestingly no latitudinal trend in clutch size was found for
North American bam swallows. In Europe and Asia, Bam Swallow
clutches are smaller at low latitudes and larger at high latitudes
(Moller 1994). Similar trends occur in other species in both
western and eastern hemispheres. Thus increasing clutch size with
increasing latitude has been found in house wrens nesting from
South America through Canada (Young 1994), eastern bluebirds
nesting in the United States (Dhondt et al. 2002), and great tits
nesting in Spain, Eurasia, North Africa, and the Netherlands (Sanz
1998). However, some species do not conform to this pattern. For
example Red-breasted Sapsuckers (Sphyrapicus ruber) and Red-
naped Sapsuckers {Sphyrapicus nuchalis) show no latitudinal trends

TURNER & KOPACHENA

143

in clutch size (Walters et* al 2002). Sanz (1997) found a variation
in clutch size in Pied Flycatchers relative to latitude in a study in
central Spain, the Netherlands, and Great Britain, but Jarvinen
(1989) did not find this to be true in Finland. In the case of North
American bam swallows there does not appear to be sufficient
latitudinal stratification of resource availability and constraints on
reproduction to warrant latitudinal trends in clutch size.

Climate change has been shown to affect reproductive ecology
of birds in recent years (Gordo et al. 2004; Both et al. 2006; Visser
et al. 2006). However, we found that clutch size, hatching success,
and fledging success among barn swallows has not varied
substantially from 1956 to 2000. Thus, there is no evidence that
global warming is affecting North American bam swallows based
on this rather crude analysis. This result may not be surprising
given that bam swallows, as an adaptable generalist species, seem
well adapted to accommodating environmental fluctuations and
novel conditions.

Acknowledgements

The research was supported by three Texas A&M University-
Commerce Mini Grants and a Texas A&M Univerisity-Commerce
faculty research grant. The field assistance of Dr. Tony Buckley,
Greg Potts, and Sigrid Slemp is gratefully acknowledged.

Literature Cited

Altemus, E. L. 1977. Bam swallow nesting data. Cassinia, 56:24-25.

Anthony, L. W. & C. A. Ely. 1976. Breeding biology of bam swallows in west-
central Kansas. Kansas Om. Soc. Bull., 27(4):37-43.

Barclay, R. M, R. 1988. Variation in the costs, benefits, and frequency of nest reuse
by bam swallows {Hirundo rustica). Auk, 105:53-60.

Barr, A. L, 1979. Nesting success, growth rates, and recruitment of bam swallows
Hirundo rustica in Brazos County, Texas. Unpublished MS Thesis, Texas A&M
University, College Station, Texas, 50pp.

Both, C., S. Bouwhuis, C. M. Lessells & M. E. Visser. 2006. Climate change and
population declines in a long-distance migratoiy bird. Nature, 44:81- 83.

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Brown, C. R. & M. B. Brown. 1999. Bam swallow {Hirundo rusticd). Pp. 1-32 in
The Birds of North America, No. 452 (A. Poole & F. Gill, eds.). The Birds of
North America, Inc., Philadelphia, PA.

Crook, J. R. & W. M. Shields. 1985. Sexually selected infanticide by adult male
bam swallows. Anim. Behav., 33:754-761.

Dhondt, A. A., T. L. Kast & P. E. Allen. 2002. Geographical differences in seasonal
clutch size variation in multi-brooded bird species. Ibis, 144:646-651.

Goodman, S. M. 1982. A test of nest cup volume and reproductive success in the
bam swallow. Jack-Pine Warbler, 60: 107-1 12.

Gordo, O., L. Brotons, X. Ferrer & P. Comas. 2004. Do changes in climate patterns
in wintering areas affect the timing of the spring arrival of trans-Saharan migrant
birds? Global Change Biology, 11: 12-21.

Gme, C. E., T. J. O’Shey & D. J. Hoffman. 1984. Lead concentrations and
reproduction in highway-nesting bam swallows. Condor, 86:383-389.

Hasselquist, D., M. F. Wasson & D. W. Winkler. 2001. Humoral
immunocompetence con’elates with date of egg-laying and reflects work load in
female tree swallows. Behav. Ecol., 12:93-97.

Jarvinen, A. 1989. Clutch-size variation in the Pied Flycatcher Ficedula hypoleuca.
Ibis, 131:572-577.

Johnsgard, P. A. 1979. Birds of the Great Plains: the breeding species and their
distribution. Univ. Nebraska Press, Lincoln, xlv+539 pp.

Johnston, R. F. 1960. Directory to the bird-life of Kansas. Univ. Kansas Mus. Nat.
Hist. Misc Publ., 23:1-69.

King, K. A., T. W. Custer & D. A. Weaver. 1994. Reproductive success of bam
swallows nesting near a selenium-contaminated lake in East Texas, USA. Env.
Poll., 84:53-58.

Kopachena, Jeffrey G., Anthony J. Buckley & Greg A. Potts. 2000. Effects of the
American swallow bug {Oeciacus vicarius) on reproductive success of bam
swallow (Hinmdo rustica). Tex. J. Sci., 52(1): 33-47.

Kopachena,. J. G., B. L. Cochran & T. B. Nichols. 2007. The incidence of American
swallow bugs {Oeciacus vicarius) in bam swallow {Hirundo rustica) colonies in
northeast Texas. J. Vector Ecol., 32: 280-284.

Kose, M., R. Mand & A. P. Moller. 1999. Sexual selection for white tail spots in the
bam swallow in relation to habitat choice by feather lice. Anim. Behav.,
58:1201-1205.

Lohoefener, R. 1980. Comparative breeding biology and ethology of colonial and
solitary nesting bam swallows {Hirundo rustica) in east-central Mississippi.
Ph.D. Dissertation, Mississippi State University, Mississippi, ix+72 pp.

Martin, R. F. 1974. Syntoptic culvert nesting of cave and bam swallows in Texas.
Auk, 91:776-782.

McGinn, D. B. & H. Clark. 1978. Some measurements of swallow breeding biology
in lowland Scotland. Bird Study, 25:109-1 18.

Merino, S., A. P. Moller, & F. de Lope. 2000. Seasonal changes in cell-mediated
immunocompetence and mass gain in nestling bam swallows: a parasite-mediated
effect? Oikos, 90:327-332.

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Moller, A. P. 1991. Clutch size, nest predation, and distribution of avian unequal
competitors in a patchy environment. Ecology, 72:1336-1349.

Moller, A. P. 1994. Sexual selection and the bam swallow. Oxford Univ. Press,

Oxford, x+365pp.

Moller, A. P. 2000. Survival and reproductive rate of mites in relation to resistance
of their bam swallow hosts. Oecologia, 124:351-357.

Moller, A. P. 2004. Rapid temporal change in frequency of infanticide in a passerine
bid associated with change in population density and body condition. Behav.
Ecol., 15:462-468.

Perrier, C., F. de Lope, A. P. Moller & P. Ninni. 2002. Stmctural coloration and
sexual selection in the bam swallow Hirimdo rustica. Behav. Ecol., 13:728-736.

Peixins, C. M. 1970. The timing of birds’ breeding seasons. Ibis, 112:242-255.

Ramstack, J. M., M. T. Murphy, & M. R. Palmer. 1998. Comparative reproductive
biology of three species of swallows in a common environment. Wilson Bull.,
110:233-243.

Saino, N., A. M. Bolzem & A. P. Moller. 1997. Immunocompetence,
ornamentation, and viability of male bam swallows {Hirundo rustica). Proc. Nat.
Acad. Sci., 94:549-552.

Saino, N., S. Calza & A. P. Moller. 1998. Effects of a dipteran ectoparasite on
immune response and growth trade-offs in bam swallow, Hirimdo rustica,
nestlings. Oikos, 81:217-228.

Saino, N. & A. P. Moller. 1996. Sexual ornamentation and immunocompetence in
the bam swallow. Behav. Ecol., 7:227-232.

Saino, N., M. Incagli, R.Martinelli, & A. P. Moller. 2002. Immune response of male
bam swallows in relation to parental effort, corticosterone plasma levels, and
sexual ornamentation. Behav. Ecol., 13:169-174.

Saino, N., R. Ambrossini, R. Martinelli, P. Ninni, & A. P. Moller. 2003. Gape
coloration reliably reflects immunocompetence of bam swallow {Hirimdo
rustica) nestlings. Behav. Ecol., 14:16-22.

Samuel, D. E. 1971. The breeding biology of bam and cliff swallows in West
Virginia. The Wilson Bull., 83:284-301.

Sanz, J. J. 1997. Geographic variation in breeding parameters of the pied flycatcher
Ficedula hypoleuca. Ibis, 139: 107-1 14.

Sanz, J. J. 1998. Effeets of geographic location and habitat on breeding parameters
of great tits. Auk, 1 15:1034-1051.

Shields, W. H. & J. R. Crook. 1987. Bam Swallow coloniality: A net cost for group
breeding in the Adirondacks? Ecology, 68:1373-1386.

Smith, H. G. & R. Montgomerie. 1991, Sexual selection and the tail ornaments of
North American bam swallows. Behav. Ecol. and Sociobiol., 28:195-201.

Snapp, B. D. 1976. Colonial breeding in the bam swallow {Hirundo rustica) and its
adaptive significance. Condor, 78:471-480.

St. Louis, V. L., J. C. Barlow & J. R. A. Sweerts. 1989. Toenail-clipping: a simple
technique for marking individual chicks. J. Field Omithol., 60:21 1-215.

Stenseth, N., A. Mysetemd, G. Ottersen, J. W. Hurrell, K. Chan & M. Lima. 2002.
Ecological effects of climate fluctuations. Science, 297:1292-1296.

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Thompson, M. C. 1961. Observations on the nesting success of the Bam Swallow in
South-Central Kansas. Kansas Om. Soc. Bull., 13(2):9-11.

Visser, M. E., L. J. M. Holleman, & P. Gienapp. 2006. Shifts in caterpillar biomass
phenology due to climate change and its impact on the breeding biology of an
insectivorous bird. Oecologia, 147:164-172.

Walters, E. L., E. H. Miller, & P. E. Lowther. 2002. Red-breasted Sapsucker
{Sphyrapicus ruber) and Red-naped Sapsucker {Sphyrapicus nuchalis). Pp. 1-32
in The Birds of North America, No. 663 (A. Poole & F. Gill, eds.). The Birds of
North America, Inc., Philadelphia, PA.

Young, B. E. 1993. Geographical and seasonal patterns of clutch-size variation in
House Wrens. The Auk, 1 1 1:545-555.

JGK at: Jeff_Kopachena@tamu-commerce.edu

TEXAS J. SCI. 61(2), MAY, 2009

147

GENERAL NOTES

REPRODUCTIVE CYCLE OF
THE CENTRAL AMERICAN MABUYA,
MABUYA UN MARGIN ATA (SQUAMATA: SCINCIDAE)
FROM COSTA RICA

Stephen R. Goldberg

Department of Biology’, Whittier College, PO Box 634
Whittier, California 90608

The Central American mabuya, Mabuya iinimarginata is a
common diurnal skink ranging from Colima and Veracruz, Mexico
to Panama at 1-1,500 m where it inhabits a variety of habitats
(Savage 2002). Information on its reproduction is in Webb (1958),
McCoy (1966), Fitch (1973), Alvarez del Toro (1982), Savage
(2002) and Guyer & Donnelly (2005). Females give birth to live
young. The purpose of this paper is to add to the knowledge of the
reproductive biology of M unimarginata from a histological
analysis of gonadal material. The first information on the M
unimarginata testicular cycle is presented. New minimum sizes for
reproduction in males and females of M unimarginata are reported.

A total of 25 adult males (mean snout-vent length, SVL = 64.8
mm ± 7.4 SD, range = 51-77 mm) and 30 females (SVL = 70.0 mm
± 11.0 SD, range = 50-92 mm) from Costa Rica were examined
from the herpetology collection of the Natural History Museum of
Los Angeles County (LACM), Los Angeles, California.

The left testis was removed from males and the left ovary was
removed from females for histological examination. Tissues were
embedded in paraffin, sectioned at 5 pm and stained with
Harris’hematoxylin followed by eosin counterstain (Presnell &
Schreibman 1997). Histology slides are deposited in LACM. An
unpaired Mest was used to compare male and female mean body
sizes and the relationship between female SVL and litter size was

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examined by linear regression analysis using Instat, vers. 3.0b,
Graphpad Software, San Diego, CA.

Material examined -1\\q following specimens of M
unimarginata from Costa Rica (by province) were examined:
Guanacaste: {n = 22) LACM 166201, 166202, 166204, 166211,
166214, 166219, 166226, 166228, 166230, 166231, 166234-

166236, 166240, 166244, 166245, 166249-166251, 166257,

166260, 166264. Heredia: = 1) LACM 166252. Limon: {n=\2)
LACM 166203, 166210, 166215, 166218, 166220, 166227,

166243, 166253, 166262, 166265, 166266, 166277. Puntarenas: {n
= 20) LACM 166200, 166213, 166216, 166224, 166225, 166238,
166239, 166241, 166242, 166248, 166255, 166258, 166261,

166269, 166271-166275, 166279.

Three stages in the testicular cycle (Table 1) were present: (1)
regressed (= quiescence), seminiferous tubules contain Sertoli cells
and spermatogonia; (2) recrudescence (= recovery), proliferation of
cells in the germinal epithelium is in progress, primary, secondary
spermatocytes and, in some cases, spermatids are present; (3)
spermiogenesis (= sperm production), lumina of seminiferous
tubules lined by spermatozoa; clusters of metamorphosing
spermatids are present. The smallest reproductively active male
(spermiogenesis in progress) measured 52 mm SVL (LACM

166279) and was collected in February. This likely represents a
new minimum size for maturity of M. unimarginata males.
Spermiogenesis occurred in all months except August (only 3 males
examined), suggesting that sperm production may be continuous.

The mean body size of M. unimarginata females was
significantly larger than that of males (unpaired t test, t = 2.0, df=
53, P = 0.05). Females ovulate tiny eggs which develop to live
young. The smallest reproductively active female measured 50 mm
SVL (2 mm diameter eggs in oviducts) and was from May (LACM
166234). This likely represents a new minimum size for maturity
of M. unimarginata females. Reproductive activity (Table 2) was

TEXAS I SCI 61(2), MAY, 2009

149

recorded in all months sampled, suggesting reproduction may be
continuous. One female from February (LACM 166231) contained
corpora lutea and convoluted oviducts indicating parturition had
recently occurred. Females with discernible embryos were seen in
April and June to September. Linear regression analysis indicated a
significant positive correlation between female body size (SVL)
and developing eggs and/or embryos: n = 30, r = 0.48, P = 0.007, Y
= ~0.13 + 0.07X.

The finding of males exhibiting recrudescence from June to
September, and one male each from July and September with
regressed testes, suggests there may be some seasonality in the
testicular cycle of M unimarginata. In contrast, only spermiogenic
males were reported for the sympatric gecko, Lepidoblepharis
xanthostigma, by Goldberg (2008).

Fitch (1973) reported two females from March that contained
well-developed embryos and suggested births may be concentrated
around March. The finding of one female from February that had
recently given birth supports the hypothesis of Fitch (1973). In
contrast, Webb (1958) reported that M unimarginata (as Mabuya
mabouya) from southern Mexico gave birth to 4-6 young in June
and July. The size for sexual maturity in males was estimated at 56
mm SVL and for females at 62 mm, with a suggestion that females
as small as 56 mm SVL were mature (Webb, 1958). Alvarez del
Toro (1982) reported M unimarginata (as Mabuya brachypoda)
gave birth to 4-6 young in June through August in Chiapas, Mexico.
McCoy (1966) reported M unimarginata (as Mabuya brachypoda)
from southern El Peten Province, Guatemala contained 6-9
embryos. Somma & Brooks (1976) reported M. unimarginata (as
Mabuya mabouya) from Dominica collected in February, August,
September and December contained embryos.

While it is apparent that M unimarginata from Costa Rica has
an extended reproductive cycle with activity exhibited in all
months, subsequent work is needed to elucidate details of the

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Table 1. Monthly conditions in testicular cycle of 25 Mabuya iinimarginata from Costa
Rica. Values are the numbers of males exhibitmg each of the three conditions.

Month

Regression

Recrudescent

Spermiogenesis

January

February

March

May

June

July

August

September

October

Table 2. Monthly stages in the ovarian cycle of 30 Mabuya iinimarginata from Costa
Rica. Sizes of ovulated follicles are in mm. The post-partum female from February
contained corpora lutea and convoluted oviducts.

Month

2 mm

3-5 nun

>6 nun

Post-partum

February

April

May

June

July

August

September

October

ovarian cycle (e.g., seasonal and geographic variation in timing of
parturition).

Acknowledgment

I thank Christine Thacker (Natural History Museum of Los
Angeles County) for permission to examine M unimarginata.
Specimens are part of the CRE (Costa Rica Expeditions) collection
donated to LACM by Jay Savage in 1998.

Literature Cited

Alvarez del Toro, M. 1982. Los Reptiles de Chiapas, Tercera Edic. Coleccion Libros de
Chiapas, Tuxtla Gutienez, Chiapas, 248 pp.

TEXAS I SCL 61(2), MAY, 2009

151

Fitch, H. S. 1973. A field study of Costa Rican lizards. Univ. Kansas Sci. Bull., 50:39-
126.

Goldberg, S. R. 2008. Notes on the reproductive biology of the Costa Rica scaly-eyed
gecko, Lepidoblepharis xanthostigma (Squamata: Gekkonidae), from Costa Rica.
Bull. Chicago Herpetol, Soc., 43:130-131.

Guyer, C. & M. A. Donnelly. 2005. Amphibians and Reptiles of La Selva, Costa Rica,
and the Caribbean Slope. A Comprehensive Guide. University of California Press,
Berkeley, viii + 299 pp.

McCoy, C. J. 1966. Additions to the herpetofauna of southern El Peten, Guatemala.
Herpetologica, 22:306-308.

Presnell, J. K. & M. P. Schreibman. 1997. Humason’s Animal Tissue Techniques. 5*
Ed., Johns Hopkins Press, Baltimore, xix + 572 pp.

Savage, J. M. 2002. The Amphibians and Reptiles of Costa Rica. A Herpetofauna
Between two Continents, Between two Seas. The University of Chicago Press,
Chicago, 934 pp.

Somma, C. A. & G. R. Brooks. 1976. Reproduction in Anolis oculatm, Ameiva fmcata
and Mabuya mabouya from Dominica. Copeia, 1976:249-256.

Webb, R. G. 1958. The status of the Mexican lizards of the genus Mabuya. Univ.
Kansas Sci. Bulk, 38:1303-1313.

SRG at: sgoldberg@whittier.edu

NEW GEOGRAPHIC DISTRIBUTION RECORDS FOR
PARAJULID MILLIPEDS (DIPLOPODA: JULIDA),

IN ARKANSAS AND TEXAS

Chris T, McAllister and Henry W. Robison

RapidWrite, 102 Brown Street
Hot Springs National Park, Arkansas 71913 and
Department of Biology, Southern Arkansas University
Magnol ia, A rkansas 71754

The primary North American milliped family Parajulidae ranges
from Yakutut, Alaska, and James Bay, Ontario, to western El
Salvador (Shelley 2008), This large and complex assemblage of
taxa includes the genus Aniulus Chamberlin which, at present,
contains at least 22 species inhabiting various habitats from the
Atlantic Ocean to southwestern Utah, and north to south, from
southern Quebec to southern Texas and Arizona (Shelley 2001;
McAllister et ak 2009). Another group, the subgenus Hakiulus,

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ranges from the vicinity of the Canadian border in North Dakota
and Michigan to the Rio Grande in Texas and east to west,
extending from central Ohio to eastern Michigan to southwestern
Colorado (Shelley 2000). There are currently eight species of
Aniuliis within the subgenus Hakiulus; one of these, Aniuliis
(Hakiulus) amophor (Chamberlin), was described from material
collected along Turtle Creek, Kerr County, Texas (Chamberlin
1940). Since then, additional records of A. {Hakiulus) amophor
have been reported only from Texas (Chamberlin & Hoffman 1958;
Hoffman 1999; Shelley 2000). Herein, this study provides a
significant distributional record for A. {Hakiulus) amophor in
Arkansas, as well as 13 new county records for other
species/subspecies Aniuliis in Texas.

Between October 2002 and November 2007, millipeds were
collected from various sites in Arkansas and Texas. Collecting
techniques involved using a potato rake to move debris, turning
decaying logs, peeling bark off fallen trees, and moving leaf litter
and rocks; pitfall trapping was used at only one site. Following
preliminary identification, specimens were placed in vials
containing 70% ethanol and shipped to Rowland M. Shelley at the
North Carolina State Museum of Natural Sciences, Raleigh, North
Carolina (NCSM) for verification of identification. Voucher
specimens were subsequently deposited in the NCSM. Taxa
recovered are presented below along with distributional (state,
county, specific locality, number of millipeds and sexes, collection
date) information.

Annotated List of Species

Aniulus {Hakiulus) amophor (Chamberlin).“ARKANSAS:
Union Co., El Dorado (33.1244°N, 92.3957°W), S. 27 November
2007. A single male specimen was collected with a pitfall trap
from a yard in urban habitat. This is the first time this species has
been reported from a state outside of Texas. This milliped was
previously known from Bexar, Comal, DeWitt, Gonzales,
Guadalupe, Jasper, Jim Wells, Karnes, Kerr, Live Oak, San

TEXAS J. SCI. 61(2), MAY, 2009

153

Patricio, Wharton, and Wilson counties. The new disjunct site
reported herein is ca. 365 km NE of the most proximate locality of
A. (Hakiulus) amophor in Jasper, Jasper Co., Texas. Other
myriapods collected at this site included Eurymerodesmus
angularis Causey and Hemiscolopendra marginata (Say).

Compared to northern Arkansas where the late Nell B. Causey
(1910-1979) did most of her work (Causey 1950; 1951; and others),
little is known about the general milliped fauna of southern
Arkansas. New records for Arkansas millipeds were reported by
McAllister et al. (2002a; 2002b; 2003) and McAllister & Shelley
(2008). In addition, several miscellaneous taxonomic papers by
Shelley (1990), Shear (2003), Shelley & McAllister (2006) and
Shelley et al. (2006) reported millipeds from various southern
Arkansas counties. Obviously, additional collecting of millipeds
should be attempted in this part of the state as well as eastern
Arkansas where there are few records.

Aniulus (Hakiuhis) diversijrom diversifrons (Wood).-TEXAS:
Bowie Co., 11.3 km N DeKalb off CR (County Road) 3207, 2(S,
3$, 23 November 2004; Delta Co., Cooper Lake State Park,
Doctors Creek Unit, S, 3 January 2003; Hood Co., Fort Spunky off
CR 1120, 3(5', 49, 17 February 2005; Marion Co., Berea Six off
FM 728, (5^, $, 2 juvs., 23 October 2002; Titus Co., Argo, off FM
1993 at Snake Creek, 10 November 2003. These sites (Fig. 1)
document five new county records for A. (Hakiulus) diver sifi^ons
diversifrons. It has previously been reported from much of the
central United States, including Arkansas, Illinois, Michigan,
Minnesota, Missouri, North Dakota, Ohio, Oklahoma, and Texas
(Shelley 2000). In Texas, this milliped was known previously from
Anderson, Baylor, Brown, Cherokee, Colorado, Dallas, Erath,
Grayson, Hamilton, Hopkins, Hunt, Lavaca, Milam, Potter,
Randall, Smith, Stonewall, Travis, Val Verde, and Wilson counties
(Stewart 1969; Shelley 2000).

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Figure 1. Distribution of Aniuhis spp. in Texas (solid symbols = new records; open
symbols = literature records). A. (Hakiulns) diversifrom diversifrom (stars); A.
(Hakhilits) diversifrom neomexicamis (crosses); A. (Hakiulus) diversifrom
intergrades (diamonds); A. brazomis (dots); A. craterm craterus (squares); A.
craterus intergrades (triangles); A. fluviatilis (inverted triangles).

Aniiilus {Hakiulus) diversifrons neomexicanus (Chamberlin).-
TEXAS: Brown Co., Brownwood State Park, 20.9 km NNW
Brownwood, 27 November 2002; Limestone Co., Confederate
Reunion Grounds State Park, 6(5^, 189, 21 December 2002. This
milliped was previously known from Colorado and New Mexico,
and Hudspeth, Potter, and Randall counties, Texas (Shelley 2000).
Two new county records are documented herein (Fig. 1) that are
considerable distances east of previous Texas localities in the
Panhandle and Guadalupe Mountains areas of the state.

Aniulus (Hakiulus) diversifrons intergrades.-TEXAS: Coleman
Co., 3.2 km E Talpa off US 67, 2S, 29, juv., 23 December 2006;
Tom Green Co., San Angelo, 156 Las Lomas Trail, 6S, 79, 4 juvs.,
13 November 2005 & 24 December 2006. These specimens display
gonopodal characters intermediate between those of two races, A.
(Hakiulus) diversifrons diversifrons and A. (Hakiulus) diversifrons

TEXAS J. SCI. 61(2), MAY, 2009

155

neomexicanus (Chamberlin). Shelley (2000) previously reported
similar intergrades from Garza Co., Texas, and Larimer Co.,
Colorado, and the new sites (county records. Fig. 1) are a
considerable distance northwest and southeast, respectively, from
those localities.

Aniulus brazonus Chamberlin .-TEXAS: Dallas Co., Cedar Hill
State Park, Talaha Trail, S, 2$, 16 November 2002. This
represents only the second record of A. brazonus ever documented,
as the species was previously known only from the type locality in
Brazos County, Texas (Shelley 2001). These specimens were
collected underneath leaf litter and decaying logs in prairie habitat
dominated by mesquite, live oak, and ashe juniper; the new site
(Fig. 1) is ca. 240 km N of the type locality.

Aniulus craterus craterus Chamberlin.-TEXAS: Kimble Co.,
South Llano River State Park, 3.2 km S Junction, 6S, 2$, 22
February 2003; 1.6 km NE Telegraph off US 377 at Llano River,
3(?, 3$, 21 February 2004; 8.0 km SW Junction off US 377 at
Bailey Creek, c?, 21 February 2004. This milliped was previously
reported from Bandera (Loomis 1959) and Bexar and Kerr counties
(Shelley 2001). This study documents a new county record (Fig. 1)
in the Edwards Plateau (Hill Country), the northernmost localities
for^. craterus in the state.

Aniulus craterus intergrades.-TEXAS: Uvalde Co., Gamer
State Park, Crystal Cave, 2 March 2004. These specimens
display gonopodal characters intermediate between those of two
subspecies, A. craterus craterus and^. craterus fill Loomis. This is
the first time, to the author’s knowledge, that Aniulus millipeds
have been reported from Uvalde County (see Fig. 1).

Aniulus fluviatilis Chamberlin.-TEXAS: Hood Co., Fort
Spunky off CR 1 120, 3(5^, 20 February 2004. This represents a new
county record as A. fluviatilis was known previously from Brazos
and Polk counties (Causey 1952; Shelley 2001). The new site (Fig.

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1) is a considerable distance northwest and west of previously
reported localities into the Cross Timbers area of the state.
Specimens were collected underneath rocks in disturbed shortgrass
prairie habitat dominated by mesquite, ashe juniper, live oak, and
cottonwood.

Acknowledgments

We thank Dr. R. M. Shelley (NCSM) for specimen identification
and curatorial assistance, the Arkansas Game and Fish Commission
and Texas Parks and Wildlife Department for scientific collecting
permits issued to CTM, and J. T. McAllister, III, D. Moore, T.
Ratliff, and T. Sciara for assistance in collecting.

Literature Cited

Causey, N. B. 1950. Five new Arkansas millipeds of the genera Eiirymerodesmus and
Paresmiis (Xystodesmidae). Ohio J. Sci., 50:267-272.

Causey, N. B. 1951. On Eurymerodesmidae, a new family of Diplopoda
(Strongylosomidae), and a new Arkansas species of Eurymerodesmus. Proc.
Arkansas Acad. Sci., 4:69-71.

Causey, N. B. 1952. New species and records of paraiulid millipeds from Texas. Texas
J. Sci., 4(2):200-203.

Chamberlin, R. V. 1940. New genera and species of North American Paraiulidae. Bull.
Univ. Utah, 30 [Biol. Ser. 5]: 1-39.

Chamberlm, R. V. & R. L. Hoffean. 1958. Checklist of the millipeds of North America.
U.S. Nat. Mus. Bull., 212:1-236.

Hoffman, R. L. 1999. Checklist of the millipeds of North and Middle America. Virginia
Mus. Nat. Hist. Spec. Publ., 8:1-584.

Loomis, H. F. 1959. Millipeds collected enroute from Florida to San Antonio, Texas,
and vicinity. J. Washington Acad. Sci., 49:157-163.

McAllister, C. T. & R. M. Shelley. 2008. New records of eurymerodesmid millipeds
(Diplopoda: Polydesmida) from Arkansas, Kansas, Louisiana, Oklalioma, and Texas.
J. Arkansas Acad. Sci., 62:155-158.

McAllister, C. T., R. M. Shelley & J. T. McAllister, III. 2002a. Millipeds (Artluopoda:
Diplopoda) of the Ark-La-Tex. 11. Distributional records for some species of western
and central Arkansas and eastern and southeastern Oklahoma. J. Arkansas Acad.
Sci., 56:95-98.

McAllister, C. T., R. M. Shelley & J. T. McAllister, III. 2003. Millipeds (Arthiopoda:
Diplopoda) of the Ark-La-Tex. III. Additional records from Arkansas. J. Arkansas
Acad. Sci., 57:115-121.

McAllister, C. T., R. M. Shelley & S. E, Trauth. 2009. Aniuhis garius (Chamberlin,
1912), a widespread milliped in central and eastern North America (Julida:
Parajulidae: Aniulini). Spec. Publ. Virgmia Mus. Nat. Hist., 16:229-238.

TEXAS J. SCI. 61(2), MAY, 2009

157

McAllister, C. T., C. S. Harris, R. M. Shelley & J. T. McAllister, III. 2002b. Millipeds
(Arthropoda: Diplopoda) of the Ark-La-Tex. 1. New distributional and state records
for seven counties of the west Gulf Coastal Plain of Arkansas. J. Arkansas Acad.
Sci., 56:91-94.

Shear, W. A. 2003. Branneria bonocuhis n. sp., a second species in the North American
milliped family Brarmeriidae (Diplopoda: Chordeumatida: Brannerioidea). Zootaxa,
233:1-7.

Shelley, R. M. 1990. Revision of the milliped family Eurymerodesmidae (Polydesmida:

Chelodesmidea). Mem. American Entomol. Soc., 37:1-112.

Shelley, R. M. 2000. Parajulid studies II. The subgenus Hakwlus Chamberlin (Julida:

Parajulidae: Parajulinae: Aniulmi). Myriapodologica, 6:121-145.

Shelley, R. M. 2001. A synopsis of the milliped genus Aniidiis Chamberlin (Julida:

Parajulidae: Parajulinae: Aniulini). Texas Mem. Mus., Speleol. Monogr., 5:73-94.
Shelley, R. M. 2008. Way down south: the milliped family Parajulidae (Julida:
Parajulini) in Mexico and Central America; first records from El Salvador and the
Baja California peninsula. Zootaxa, 1893:1-37.

Shelley, R. M. & C. T. McAllister. 2006. Composition and distribution of the milliped
tribe Pachydesmini west of the Mississippi River (Polydesmida: Xystodesmidae). W.
North Amer. Nat., 66:45-54.

Shelley, R. M., C. T. McAllister & M. F. Medrano. 2006. Distribution of the milliped
genus Narceus Rafinesque, 1820 (Spirobolida: Spirobolidae): Occurrences in New
England and west of the Mississippi River: A summary of peripheral localities; and
first records from Connecticut, Delaware, Maine, and Minnesota. W. North Amer.
Nat., 66:374-389.

Stewart, T. C. 1969. Records of millipeds in twenty five northeast Texas counties.
Texas J. Sci., 20(4):383-385.

CTM at: drctmcallister@aol.com

NOTEWORTHY RECORDS OF
DRAGONFLIES (ODONATA: ANISOPTERA) FROM
JONES AND TAYLOR COUNTIES OF CENTRAL TEXAS

Thomas E. Lee, Jr., Amisha J. Patel, Benjamin W. Johnson
and Roy C. Vogtsberger*

Department of Biology, Abilene Christian University
Abilene, Texas 79601 and
^Department of Biology, Midwestern State University
Wichita Falls, Texas 76308

There is growing interest in the biology of dragonflies (Odonata:
Anisoptera) among both amateur naturalists and professional
biologists (e.g., Dunkle 2000; Kondratieff 2000; Merritt & Cummins

158

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

1996; Milne & Milne 1994; Needham & Westfall 1955; Needham et
al. 2000; Silsby 2001). Little is known of the dragonfly fauna of the
Southern Rolling Plains (Abbott 2001), although Abbott et al. (2003)
provided a recent synopsis of dragonflies from the Texas Panhandle.

Reported below are the partial results of recent collecting efforts
from the contiguous central Texas counties of Jones and Taylor. This
study examined the holdings from Abilene Christian University
Natural History Collection (ACUNHC) and Hardin- Simmons
University Invertebrate Collection (HSUIC), which provided records
for 23 species of anisopterans, of which eight are from the families
Aeshnidae, Corduliidae, and Libellulidae. Eight taxa were geo¬
graphically noteworthy, and are listed below. The majority of the
specimens were collected between 1966 and 1977, although some
specimens were taken as recently as 2001. Identifications were based
on Merritt & Cummins (1996) and Dunkle (2000). Kondratieff
(2000) and Reece & McIntyre (2008) provided the basis for
detennination of geographic significance.

Family Aeshnidae

Rhionaeschna multicolor (Hagen I861).-Taylor County: un¬
specified locality, 1 (ACUNHC 0956). The specimen represents a
county record, with the most proximate record from adjoining Jones
County.

Family Corduliidae

Didymops transversa (Say 1839).-Taylor County: Abilene, 1
(HSUIC 4). This specimen represents a county record and range
extension of approximately 200 km to the north from Kimble County.

Epitheca costalis (Selys 1871).“Taylor County: unspecified
locality, 3 (ACUNHC 0998, ACUNHC 0999, and ACUNHC 01000);
Abilene, 1 (HSUIC 2). These specimens represent a county record
and range extension of approximately 200 km to the north from
Kimble County.

TEXAS J. SCI. 61(2), MAY, 2009

159

N eurocordulia xanthosoma (Williamson 1908) -Jones County:
unspecified locality, 1 (ACUNHC 01003). This specimen represents
a county record and range extension of approximately 200 km to the
northwest from San Saba County.

Family Libellulidae

Libellula luctuosa (Burmeister 1839) —Jones County: Hawley
Farm, Clear Fork of the Brazos River, 3 (ACUNHC 0489, ACUNHC
0736, and ACUNHC 0737); Lake Fort Phantom Hill, 1 (ACUNHC
0545). These specimens represent a county record, with the most
proximate record from adjoining Taylor County.

Pachydiplax longipennis Burmeister 1839. -Jones County: Hawley
Farm, Clear Fork of the Brazos River, 1 (ACUNHC 0589). This
specimen represents a county record, with the most proximate record
from adjoining Taylor County.

Plathemis lydia (Drury 1773). -Jones County: Hawley Fanu, Clear
Fork of the Brazos River, 1 (ACUNHC 0740); Lake Fort Phantom
Hill, 2 (ACUNHC 0516 and ACUNHC 0517). These specimens
represent a county record, with the most proximate record from
adjoining Taylor County.

Tramea lacerata Hagen 1861. -Jones County: Hawley Farm, Clear
Fork of the Brazos River, 1 (ACUNHC 0495). Taylor County:
Abilene, 1 (HSUIC 13). These specimens represent county records,
with the most proximate record from approximately 80 km to the
north from Coleman County.

Literature Cited

Abbott, J. C. 2001. Distribution of dragonflies and damselflies (Odonata) in Texas.

Trans. American Entomol. Soc., 127:189-228.

Abbott, J. C., R. A. Belirstock & R. R. Larsen. 2003. Notes on the distribution of
Odonata in the Texas Panhandle, with a summary of new state and county records.
Southwestern Nat., 48(3):444-449.

Dunkle, S. W. 2000. Dragonflies Through Binoculars: A Field Guide to Dragonflies of
North America. Oxford Univ. Press, New York, 266 pp.

160

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 2, 2009

Kondratieff, B. C. 2000. Dragonflies and Damselflies (Odonata) of the U. S.
Jamestown, North Dakota; Northern Prairie Wildlife Research Center, US Geological
Survey (government web page).

Merritt, R. W. & K. W. Cummins (eds.). 1996. An hitroduction to the Aquatic Insects of
North America, ed. Kendall/Hunt Publ. Co., Dubuque, Iowa, 862 pp.

Milne, L. & M. Milne. 1994. National Audubon Society Field Guide to North American
Insects and Spiders. Alfred A. Knopf, New York, 989 pp.

Needliam, J. G. & M. J. Westfall, Jr. 1955. A Manual of the Dragonflies of North
America. Univ. of California Press, Berkeley, xii + 615 pp.

Needham, J. G., M. J. Westfall, Jr. & M. L. May. 2000. Dragonflies of North America.

Scientific Publishers, Gainesville, Florida, 615 pp.

Reece, B. A. & N. E. McIntyre. 2008. Dragonfly (Odonata: Anisoptera) Holdings of the
Museum of Texas Tech University. Occasional Papers, Museum of Texas Tech
University, 279:1-13.

Silsby, J. 2001. Dragonflies of the World. Smithsonian Institution Press, Washington,

216 pp.

TEL at: lee@biology.acu.edu

THE TEXAS ACADEMY OF SCIENCE, 2009-2010

OFFICERS

William J. Quinn, St. Edward’s University
Benjamin A. Pierce, Southwestern University
Romi L. Burks, Southwestern University
Raymond C. Mathews, Jr., Texas Water Dev. Board
Fred Stevens, Schreiner University
Diane B. Hyatt, Texas Water Development Board
Ned E. Strenth, Angelo State University
Frederick B. Stangl, Jr., Midwestern State University
John A. Ward, Brooke Army Medical Center
AAAS Council Representative: James W. Westgate, Lamar University
International Coordinator: Armando J. Contreras, Universidad Autonoma de N.L.

DIRECTORS

2007 Renard L. Thomas, Texas Southern University
Bob Murphy, Texas Parks and Wildlife Department

2008 Christopher M. Ritzi, Sul Ross State University
Andrew C. Kasner, Audubon Texas

2009 Ana B. Christensen, Lamar University
Thomas L. Arsuffi, Texas Tech at Junction

SECTIONAL CHAIRPERSONS

Anthropology: Raymond Mauldin, University of Texas at San Antonio
Biomedical: G. Scott Weston, University of the Incarnate Word
Botany: David Lemke, Texas State University

Cell and Molecular Biology: Magaly Rincon-Zachary, Midwestern State University

Chemistry and Biochemistry: J. D. Lewis, St. Edward’s University

Computer Science: James McGuffee, St. Edward’s University

Conservation Ecology: Wendi Moran, Hardin-Simmons University

Environmental Science: Kenneth R. Summy, University of Texas-Pan American

Freshwater Sciences: Matt Chumchal, Texas Christion University

Geosciences: Chris Barken, Stephen F. Austin State University

Marine Sciences: Larry D. McKinney, Harte Research Institute

Mathematics: Elsie M. Campbell, Angelo State University

Physics: David L. Bixler, Angelo State University

Science Education: Patricia Ritschel-Trifilo, Harden-Simmons University

Systematics and Evolutionary Biology: Tara Maginnis, St. Edward’s University

Terrestrial Ecology and Management: Richard Patrock, St. Edward’s University

COUNSELORS

Collegiate Academy: David S. Marsh, Angelo State University
Junior Academy: Vince Schielack, Texas A&M University

President:

President Elect:
Vice-President:

Immediate Past President:
Executive Secretary:
Corresponding Secretary:
Managing Editor:
Manuscript Editor:
Treasurer:

PERIODICALS

THE TEXAS JOURNAL OF SCIENCE
Texas Academy of Science
CMB 6252
Schreiner University
Kerrville, Texas 78028-5697

^T-^X
^ ft

THE

TEXAS JOURNAL

SCIENCE

GENERAL INFORMATION

www.texasacademyofscience.org

Dr. Andrew C. Kasner
The Texas Academy of Science
Wayland Baptist University
1900 West V* Street - CMB 629
Plainview, Texas 79072
E-mail: kasnera@wbu.edu

The Texas Journal of Science (ISSN 0040-4403) is published quarterly at Lawrence, Kansas
(Allen Press), U.S.A. Periodicals postage paid at San Angelo, Texas and additional mailing
offices. POSTMASTER: Send address changes and returned copies to The Texas Journal of
Science, Dr. Andrew C. Kasner, 1900 West 7*'’ Street - CMB 629, Wayland Baptist University,
Plainview, Texas 79072, U.S.A. The known office of publication for The Texas Journal of
Science is the Department of Biology, Angelo State University, San Angelo, Texas 76909; Dr.
Ned E. Strenth, Managing Editor.

THE TEXAS JOURNAL OF SCIENCE

Volume 61, No. 3 August, 2009

CONTENTS

Nesting Ecology of Golden Mice {Ochrotomys nuttalli) in Eastern Texas.

By Cody W Edwards and Andy P. Bradstreet . . . . . 163

The Sand Dollar Periarchus lyelli (Echinoidea: Clypeasteroida: Scutelliformes)
in the Caddell Formation (Upper Eocene) of Texas.

By Louis G. Zachos . . . . . . . 181

Mixed Infections of Nasopharyngeal Bots, Cephenemyia spp. (Oestridae) in

White-Tailed Deer {Odocoileus virginianus) and Mule Deer
{Odocoileus hemionus) of Texas.

By Samuel W. Kelley . . . 187

Karyotype Diversity Among and Within Avian Taxa: A Simple Test in R.

By Michael F. Small, Michael R. J. Forstner and John T, Baccus . 195

The Arkansas Endemic Fauna: An Update with Additions, Deletions,

A Synthesis of New Distributional Records, and Changes in Nomenclature.

Chris T. McAllister, Henry W. Robison and Michael E. Slay . . . 203

Selection of Available Post-Fire Substrate by the Ground Skink,

Scincella lateralis (Squamata: Scincidae).

By Charles M. Watson . . . . . . . .....219

General Notes

Reproduction in Smith’s Green-Eyed Gecko, Gekko smithii
(Squamata: Gekkonidae).

By Stephen R. Goldberg . . . . . 225

The Long-Tailed Weasel Mustela frenata (Mammalia: Mustelidae)
in Baja California, Mexico.

By Gorgonio Riiiz-Campos, Roberto Martlnez-Gallardo,

Salvador Gonzdlez-Guzmdn and Jorge Alaniz-Garcla . . . 229

Stomach Contents of Calidris minutilla (Charadriiformes: Scolopacidae)
Wintering at a Freshwater Reservoir in West-Central Texas.

By Andrew C. Kasner, Randall H. Ruddick, and Terry C. Maxwell . . . 233

Recognition of Special Members . . . . . 240

THE TEXAS JOURNAL OF SCIENCE
EDITORIAL STAFF

Managing Editor:

Ned E. Strenth, Angelo State University
Manuscript Editor:

Frederick B. Stangl, Jr., Midwestern State University
Associate Editors:

Allan D. Nelson, Tarleton State University
Jim R. Goetze, Laredo Community College
Associate Editor for Botany:

Janis K. Bush, The University of Texas at San Antonio
Associate Editor for Chemistry:

John R. Villarreal, The University of Texas-Pan American
Associate Editor for Computer Science:

Nelson Passos, Midwestern State University
Associate Editor for Geology:

Ernest L. Lundelius, University of Texas at Austin
Associate Editor for Mathematics and Statistics:

E. Donice McCune, Stephen F. Austin State University

Manuscripts intended for publication in the Journal should be submitted in
TRIPLICATE to:

Dr. Allan D. Nelson
Department of Biological Sciences
Tarleton State University
Box T-OlOO

Stephenville, Texas 76402
nelson@tarleton.edu

www.texasacademyofscience.org

AFFILIATED ORGANIZATIONS
American Association for the Advancement of Science,

TEXAS T OF SCL 6 1(3): 163- 180

AUGUST, 2009

NESTING ECOLOGY OF GOLDEN MICE
{OCHROTOMYS NUTTALLF) m EASTERN TEXAS

Cody W* Edwards and Aady P* Bradstreet

Department of Environmental Science and Policy, George Mason University
4400 University Drive, MSN 5F2, Fairfax, Virginia 22030 and
Department of Biology, Stephen F. Austin State University
Nacogdoches, Texas 75961

Abstract -Golden mice {Ochrotomys nuttalli) construct distinctive arboreal nests
throughout much of their range. Little is known of the arboreal nature of golden mice
in eastern Texas and previous studies have produced conflicting results. Live-
trapping (7,150 trap nights) was conducted in four East Texas counties: Angelina,
Houston, Nacogdoches, and San Augustine. Golden mice were trapped at two
locations in Nacogdoches County and one location in San Augustine County. Results
indicate viable populations of golden mice remain in eastern Texas, although
additional surveys are needed. Fourteen golden mice were radio-tracked at the
Stephen F. Austin State University Experimental Forest (Nacogdoches County,
Texas) to determine nest site selection during two seasons. No evidence of arboreal
nesting in golden mice was found during the summer. However, significant levels of
arboreal nest use were recorded during autumn. The lack of ground-level structural
diversity in eastern Texas forests in conjunction with seasonal flooding may cause
terrestrial nest locations to become a limiting resource among small mammals during
autumn.

Resumen.-Los ratones dorados {Ochrotomys nuttalli) construyen nidos arboreos
muy distintivos a lo largo de todo su ambito. Sin embargo, se conoce muy poco de la
naturaleza arborea de los ratones dorados en el Este de Texas, y los estudios previos
han obtenido resultados divergentes. Se distribuyeron trampas (7,150 trap nights) en
cuatro distritos del Este de Texas: Angelina, Houston, Nacogdoches y San Augustine,
capturando ratones dorados en dos localidades en el distrito de Nacogdoches y en una
localidad en el distrito San Augustine. Nuestro estudio indica que aun existen
poblaciones viables de ratones dorados en el Este de Texas, aunque resulta necesario
obtener mas registros. Catorce ratones dorados fiieron monitoreados para determinar
preferencia del lugar de anidamiento durante dos estaciones en el Stephen F. Austin
State University Experimental Forest (Nacogdoches, Texas). No se encontro
evidencia de anidamiento arboreo durante el verano. Sin embargo, niveles
significativos de anidamiento arboreo fiieron registrados durante el otono. La
deficiencia en diversidad estructural de niveles bajos en el este de Texas, en conjunto
con inundaciones estacionales, puede ocasionar que las localidades para anidamiento
terrestre sean un recurso limitante para mamiferos pequenos durante el otono.

The habitats of golden mice {Ochrotomys nuttalli) vary from
densely forested lowlands and floodplains to sandy upland pine

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 3, 2009

communities (Linzey & Packard 1977). However, McCarley
(1958) found that isolated upland communities seldom contained
golden mice, suggesting floodplains may serve as focal points for
dispersal. The main factor influencing the ecological distribution of
this species is density of underbrush (McCarley 1958). The dense
understory of golden mouse habitat is often composed of various
plant species including honeysuckle (Lonicera sp.), greenbrier
(Smilax sp.), grapevine {Vitis sp.), and poison ivy {Toxicodendron
radicans) (Goodpaster & Hoffmeister 1954). Linzey (1968) found
that the predominant food items of golden mice were greenbrier and
blackberry {Rubus sp.) seeds and invertebrates.

Golden mice are considered highly arboreal and posses well
developed abdominal musculature, a semi-prehensile tail, and large
plantar tubercles indicative of an arboreal lifestyle. Several authors
(Barbour 1942; Goodpaster & Hoffmeister 1954; Packard & Gamer
1964; Linzey 1968; Linzey & Packard 1977; Frank & Layne 1992)
have described the nests and feeding platforms of golden mice.
Briefly, arboreal nests are globular stmctures averaging 150-200
mm long, 100-125 mm wide, 100-200 mm high, and weighing 10-
30 g. Arboreal nests are usually located 1. 5-4.5 m above the
ground and are interwoven in thickets of greenbrier, grapevine, and
honeysuckle. However, some have been found as high as 10 m
above ground in pine {Pinus sp.^ and cedar {Juniperus sp.) trees.
Feeding platforms are similar to arboreal nests, but are bulkier and
incomplete, often being open ended with only a shallow covering.
No distinct feeding platforms have been observed other than those
found by Barbour (1942) and Goodpaster & Hoffmeister (1954) in
Kentucky. Further, Barrett (2007) questioned the use of feeding
platforms by golden mice.

Arboreal nesting has been documented in varying degrees
throughout the range of golden mice. Several authors (Klein &
Layne 1978; Wagner et al. 2000; Morzillo et al. 2003) suggested
arboreal nests could be used to avoid predation. Arboreal nesting
may also provide protection during extreme weather conditions.
Arboreal and semi-arboreal small mammals show high survival and

EDWARDS & BRADSTREET

165

retention of previous home range following short-term flooding
events (Stickel 1948; McCarley 1959; Packard & Gamer 1964).
McCarley (1959) noted that a one- week inundation resulted in a
differential rate of mortality between cotton mice {Peromyscus
gossypinus) and golden mice. About 60% of cotton mice remained
following recession of the water, whereas 84% of golden mice were
recaptured on the plot within previously established home ranges.
Packard & Gamer (1964) noted that the habitat shift from terrestrial
to arboreal nests took place each year well in advance of the
autumn rains indicating that this behavior may have an ecological
and climatic basis.

Goodpaster & Hoffmeister (1954) documented the use of ground
nests by golden mice after finding only empty arboreal nests during
the summer. McCarley (1958) and Pearson (1953) failed to locate
arboreal nests in eastern Texas and northern Florida indicating
exclusive use of ground nest locations. Subsequent investigations
by Easterla (1968) and Frank & Layne (1992) revealed that ground
nests are located just below the leaf litter or under logs and are
similar in composition to arboreal nests. The mechanisms
regulating nest-site selection are not known and the use of ground
nests is likely underestimated (Frank & Layne 1992).

Despite evidence of continuous arboreal nest occupation in some
populations of golden mice (Morzillo et al. 2003), no researcher has
shown exclusive use of arboreal nests. Compared to arboreal nests,
ground refugia may require significantly less investment of time
and energy to constmct and maintain. Terrestrial refuge locations
also provide better temperature moderation, providing a
thermoregulatory advantage (Goodpaster & Hoffmeister 1954;
Klein & Layne 1978; Frank & Layne 1992) and may provide better
protection from predators. Packer & Layne (1991) showed that
golden mice forage extensively on the ground. Foraging at ground
level has been indicated throughout their range as evinced by
numerous captures from live traps placed on the ground. While
foraging on the ground, terrestrial locations may provide a quicker

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 3, 2009

route of escape during attempted predation events (Morzillo et al.
2003).

Simultaneous refuge use is a common occurrence in golden mice
and has been used to indicate sociality (Linzey & Packard 1977;
Dietz & Barrett 1992; Morzillo et al. 2003). However, most
support of group nesting in golden mice was obtained from studies
conducted in the northern portions of its geographic range (Barbour
1942; Goodpaster & Hoffmeister 1954; Blus 1966; Dietz & Barrett
1992; Morzillo et al. 2003). Furthermore, much of these data were
collected during autumn. The thermoregulatory benefits of nesting
and huddling have been well documented in golden mice
(Sealander 1952; Knuth & Barrett 1984; Peles & Barrett 2007).
Because of the geographical and seasonal aspects of previous work,
group nesting in golden mice could be little more than an artifact of
thermoregulatory huddling. Frank & Layne (1992) reported
simultaneous refuge use in 8.2% of golden mice and group sizes of
no more than two.

Golden mice are considered only apparently secure (AS) in
Texas (Feldhamer & Morzillo 2007). Very few golden mice have
been captured in Texas during the past two decades and there is a
need to assess and monitor their population status (Schmidly 2004).

One hypothesis for the potential decline of golden mice in
eastern Texas is changing land use patterns. Because of its affinity
for mid-successional stage habitats, golden mice are dependent
upon regular disturbance (cutting, wind damage, etc.). Fire,
whether natural or prescribed, creates a landscape mosaic favorable
to golden mouse occupation. Absence of a regular fire regime has
led to the loss of mid-successional understory vegetation in much of
eastern Texas. Additional conservation concerns of golden mice
are discussed by Feldhamer & Morzillo (2007).

Objectives were to: (1) determine the population status

(presence/absence) of golden mice in four eastern Texas counties

EDWARDS & BRADSTREET

167

(Angelina, Houston, Nacogdoches, and San Augustine; Fig. 1), and
(2) describe the nesting ecology of golden mice during two distinct
seasons at the western extent of their geographic range. Specific
questions addressed included: (1) Determine the microhabitat
characteristics at nesting locations of golden mice in summer and
autumn. (2) Determine the extent of simultaneous nest use and its
potential implications for social behavior.

Materials and methods

Study area.SmwQys were conducted for golden mice in four
eastern Texas counties (Angelina, Houston, Nacogdoches, and San
Augustine; Fig. 1). Sites were located in early successional sapling
forests, mid-successional mixed pine-hardwood forests, and mature
pine forests. The surveys were concentrated in mesic, mid-
successional forests due to O, nutt allies known affinity for these
habitats in eastern Texas (McCarley 1958; Packard & Gamer 1964;
Schmidly 2004).

The Stephen F. Austin Experimental Forest (SFAEF) is a 1,036
ha tract located 15 km southwest of Nacogdoches, Texas (Fig. 1).
This area is part of the Angelina National Forest and is
administered through the Southern Research Station in cooperation
with Stephen F. Austin State University. The SFAEF consists of
approximately 730 ha of bottomland hardwood forest with the
remainder being upland pine and mixed pine-hardwood forests. A
2.5 ha grid of 144 stations spaced at 10 m intervals was established
in an ecotone area of mixed pine-hardwood forest. Dominant
woody plant species included slash pine {Pinus elliottii), loblolly
pine (P. taeda), sweetgum {Liquidambar styraciflua), greenbrier,
blackberry, and grapevine.

Trapping procedure -GQnQml population surveys were
conducted from April-August 2003 and Febmary-July 2004. The
SFAEF study plot was surveyed July, September, and November
2004. Population surveys were conducted by setting transects of 50
Sherman live traps (H.B. Sherman, Tallahassee, Florida) spaced 15

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 3, 2009

Fig. 1. Locations of counties surveyed during golden mice research project.

m apart. Traps were placed in areas of suitable golden mouse
habitat (on ground and above ground in vines) with 100-300 total
traps per survey location. Sampling of the study plot involved
placing traps within 1 m of designated trap stations. Trapping was
conducted for 2-5 consecutive nights using a bait mixture of wild
birdseed, rolled oats, peanut butter, and raisins. All captured
individuals were identified to species, sexed, and uniquely marked
by toe-clipping (Kumar 1979). Age class was defined as either
adult or juvenile on the basis of relative size, weight, and pelage
coloration (Layne 1960; Linzey & Linzey 1967). Males with
descended testes and females that were either pregnant, lactating, or

EDWARDS & BRADSTREET

169

had perforate vulva were considered to be in reproductive
condition. Mice were released at the point of capture.

Radiotelemetry and daytime refugia of golden mice were
located using radiotelemetry during two distinct seasons. The
summer radiotelemetry was conducted from 25 July to 20
September 2004, whereas autumn radiotelemetry occurred from 15
November 2004 to 5 January 2005. Small mammal radio
transmitters (2.0 g; Blackburn Transmitters, Nacogdoches, Texas)
mounted on zip-ties were attached around the neck of each mouse.
Transmitters were designed to have a range of approximately 200 m
and a battery life of 60 days. The range was variable depending on
vegetation density and depth below ground, but highly directional
at close range and accurate within 0.5 m.

Mice were weighed and fitted with transmitters in the field and
released at the point of capture. At least 12 hours elapsed between
release and first relocation attempt. Mice were located through
standard radiotelemetry techniques (White & Garrott 1990;
Millspaugh & Marzluff 2001) using a 3 element Yagi antennae and
a R-1000 receiver (Communication Specialists Inc., Orange,
California). Locations were marked using nylon stake flags.

Characterization and analysis of microhabitat -Micro\idib\t2it
variables were measured at each ground location, arboreal location,
and random locations. Length, width, depth, height above ground,
and plant species used as substrate for construction were recorded
for arboreal nests. Six independent variables were measured at
each refuge and the randomly selected control site. Variables
measured included depth (cm) of soil litter layer, distance (m) to
nearest dead wood (log or stump) greater than 10 cm in diameter,
distance (m) to nearest vine suitable for climbing, distance (m) to
nearest tree with a DBH greater than 7.5 cm, mean percentage of
area blocked visually by vegetation from 0-1 m on a Im by Im
density board, read at 90° increments at a distance of 3 m from the
nest or random point, and mean percentage of area blocked visually

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by vegetation from 1-2 m on a Im by Im density board, read at 90°
increments at a distance of 3 m from the nest or random point.
Selected variables were modified from Dueser & Shugart (1978),
Frank & Layne (1992), and Morzillo et al. (2003) and were selected
to minimize correlation and redundancy between measurements.
Random sites were used to characterize the available microhabitat
conditions present on the study plot. Forty-eight locations were
selected using random number generation during each season.

Because microhabitat data were collected during different times
of the year, seasonal variation was possible. Non-parametric Mann-
Whitney tests of the independent variables were used to compare
microhabitat data collected during the two sampling periods. Only
random sites were used to test seasonality, as microhabitat
differences at nest sites could be caused by factors other than
seasonal variation. Forward stepwise discriminant function
analysis (DFA) was used to evaluate which microhabitat variables
were best for predicting nest site location (Table 1). Model
selection was evaluated at a significance level of 0.10.

All data were checked for univariate normality using the
Shapiro- Wilks’ Test and for multivariate normality using the
Mardia Test. Statistical analyses were performed in JMP 5.0.1
(SAS Institute Inc., Cary N.C., 1989-2002) and SAS 9.1 (SAS
Institute Inc., Cary, N.C., 2002-2003).

Results

Population survey efforts consisted of 7,150 trap
nights and resulted in the capture of 297 individuals representing
eight genera (Table 2). Overall trap success was 4.2%.
Peromyscus sp. {n = 156) comprised 52.5% of individuals captured.
Twenty-six golden mice were captured in Nacogdoches and San
Augustine Counties. Four individuals were captured in San
Augustine County, whereas six and 16 individuals, respectively,
were captured from two locations in Nacogdoches County.

EDWARDS & BRADSTREET

171

Table 1. Dependent variables used in discriminant function analysis (DFA).

Statistical Analysis

Dependent Variables

DFA 1

Golden mice ground nests

Golden mice arboreal nests

DFA 2

Random locations

Ground nest locations

DFA 3

Random locations

Arboreal nest locations

Table 2. Total captures of individuals during population survey.

Species

# of Captures

% of Captures

Peromyscus spp.

156

52.5

Reithrodontomys fulvescens

14.1

Neotoma floridana

11.1

Ochrotomys nuttalli

8.8

Blarina carolinensis

7.7

Sigmodon hispidus

3.7

Cryptotis parva

1.4

Cardinalus cardinalus

0.7

TOTAL

297

100

Radiotelemetry -Vom golden mice (two males and two females)
were radio-tracked during the summer resulting in 95 locations at
13 unique refugia. All located refugia were terrestrial. A thorough
search of the plot failed to yield any evidence of arboreal nesting.
Additionally, six males and four females were radio-tracked during
the autumn resulting in 185 locations at 26 unique refugia. Of the
26 unique refugia, 18 were located at terrestrial sites and eight
arboreal nests were found. By additional searching of the
surrounding area, three additional arboreal nests were discovered
that appeared to be in use.

Golden mice utilized an average of 3.0 refugia per individual
during autumn and 4.7 refugia per individual during summer.

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Golden mice used significantly fewer refugia in the autumn than in
the summer {U= 35.5, P = 0.02).

Sequential use, defined as the same location used by multiple
individuals at different times, was observed in 40% of golden mice
refugia. No seasonal differences were present in sequential refuge
use and sequential use was not documented at any arboreal
locations. Simultaneous use (group nesting), defined as the use of
the same refuge site by different individuals at the same time, was
not documented.

Arboreal nests were used only in autumn. Nests ranged from
0.6-2. 1 m above ground and were constructed in a variety of
substrates (Table 3). Eight of the 10 radio-collared mice were
documented using arboreal nests, but not exclusively. The number
of ground refugia used by arboreal nesting individuals ranged from
1-4. All four females were tracked to an arboreal location and four
of the six males were found in arboreal nests. Two males used
ground nests exclusively.

Microhabitat use.-No seasonal variation was found in any of the
variables (P > 0.20). Due to the lack of seasonal affects in available
habitat, seasons were pooled for subsequent analysis.

Distance to nearest vine, distance to nearest coarse woody
debris, and distance to nearest tree were retained using the stepwise
DFA classification procedure as important variables distinguishing
golden mice ground refugia from arboreal nests. The model
constructed using these variables correctly classified 100% of
observations. Stepwise logistic regression retained the same three
variables. This model was highly significant (X^ = 57.14, P <
0.0001, = 1.00). A greater distance to coarse woody debris, less

distance to nearest vine, and less distance to nearest tree were
positively associated with arboreal nest locations.

EDWARDS & BRADSTREET

173

Table 3. Arboreal nest locations and dimensions.

Nest Dimensions

Substrate

Height

(m)

Length

(cm)

Width

(cm)

Depth

(cm)

Sweetgum*

1.2

Sweetgum

1.9

Sweetgum

1.1

Devil’s Walking Stick
(Aralia spinosa)

1.4

Privet

{Ligustrum japonicum)

0.9

Loblolly Pine Sapling

1.3

Greenbrier

0.8

Greenbrier

0.6

Greenbrier*

2.1

Greenbrier

0.8

Yaupon*

{Ilex vomitoria)

1.4

* Active nests discovered, but not documented through radiotelemetry.

Distance to nearest coarse woody debris, distance to nearest
vine, and depth of litter layer were retained using the stepwise DFA
classification procedure as important variables distinguishing
ground refugia from random locations. The model constructed
using these variables correctly classified 89% of observations.
Stepwise logistic regression retained the same three variables as
significant. This model was statistically significant = 156.14, P
< 0.0001, = 0.59). A greater depth of litter layer, a greater

distance to coarse woody debris, and a greater distance to nearest
vine were positively associated with random locations.

Distance to coarse woody debris and distance to nearest vine
were retained using the stepwise DFA classification procedure as
important variables distinguishing arboreal nests from random
locations. The model constructed using these two variables
correctly classified 97% of observations. Stepwise logistic
regression retained the same two variables as significant. This
model was statistically significant (X^ = 56.76, P < 0.0001, =

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1.00). A greater distance to coarse woody debris and a greater
distance to the nearest vine were associated positively with random
locations.

Discussion

Feldhamer & May croft (1992) suggest behavioral traits of O,
nuttalli may result in underestimation of home range and population
density of the species. Although all mice captured in this study
were captured on the ground, their arboreal nature may make
trapping through conventional means less successful. Golden mice
are also known to be more trap shy than many sympatric small
mammals (Feldhamer & Maycroft 1992). This was evident in the
present study, as only five of a total of 26 golden mice were
captured during the first night at a trapping location. Seven golden
mice were captured on the second night, while nine were captured
on the third night at a given location. It is suggested that adequate
sampling techniques should include both arboreal and terrestrial
placement of traps and multiple trap nights at each location.

Historically, golden mice have been considered a habitat
specialist (Dueser & Shugart 1979; Dueser & Hallett 1980). Most
mice captured during this study were captured in dense understory
vegetation. However, golden mice were also found in areas
considered suboptimal for occupation, including a pine sapling
monoculture and mature pine forest. Both of these locations
possessed little to no understory vegetation. This is consistent with
researcher’s results from southern Illinois, where golden mice were
found in a variety of habitats (Blus 1966; Morzillo et al. 2003).
Whiting & Fleet (1987) described the small mammal composition
within four different types of even-age pine stands in East Texas:
seedling, sapling, pole, and sawtimber. Golden mice were captured
in each forest type with the largest percentage of total captures
occurring in the sapling stands {n = 42%) and the smallest
percentage of captures occurring in sawtimber {n = 16%). These
data suggest that golden mice may occur in a broader range of

EDWARDS & BRADSTREET

175

habitats in eastern Texas. However, the relationship between
habitat type and fitness has yet to be investigated.

The common occurrence of golden mice in early and mid-
successional stages suggests that regular disturbance may be an
important component in maintaining populations and enhancing
habitat. Periodic disturbance creates a mosaic habitat and increases
early successional seed-producing species important to golden mice
(Morzillo et al. 2003).

Refuge use~J\\Q use of daytime refugia and nests are vital to
small mammals for protection from extreme environmental
conditions, predator avoidance, and security for offspring (Frank &
Layne 1992). Results from this study indicate golden mice in
eastern Texas generally use fewer refugia and exhibited greater
refuge fidelity during the autumn sampling period.

Sequential use was observed in 40% of golden mice refugia and
might indicate a familiarity with the refuge locations of
conspecifics and close social grouping. Further, golden mice have
a high degree of home range overlap and exhibit little to no
evidence of territoriality (McCarley 1958; Morzillo et al. 2003).

Although no simultaneous refuge use was documented, not all
mice on the study grid were outfitted with radiotransmitters. It is
possible that untagged individuals were occupying the same refuge
site, but were not detected. However, it is likely that researchers
would have observed such groupings had aggregations occurred.
Frank & Layne (1992) recorded simultaneous refuge use in 8.2% of
golden mice locations in Florida. They found refuge use was
higher in the autumn and involved only two individuals.
Conversely, group nesting in golden mice has been reported
commonly from their northern range, with as many as eight
individuals occupying the same nest (Barbour 1942; Goodpaster &
Hoffmeister 1954; Howell 1954; Dunaway 1955; Layne 1958).
The lack of group nesting in eastern Texas supports the hypothesis
that group nesting in golden mice may be a thermoregulatory

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mechanism to conserve energy at low temperatures and not
necessarily a difference in social behavior (Frank & Layne 1992).

Refuge site microhabitat. -Duq to the presence or absence of
leaves, vegetation density measurements generally vary between
seasons. No statistical difference was found among variables
between seasons. The lack of seasonal variation is likely due to a
high abundance of woody stems within the understory vegetation.

Neither measurement of understory density was retained in the
discriminant function models. This is surprising because of the
known affinities of golden mice for dense understory habitats.
However, these measurements were not retained primarily due to an
abundance of dense understory vegetation throughout the study
plot. Natural forests in eastern Texas are a mosaic matrix of
isolated patches of dense vegetation separated by intervening
regions of open forest floor (McCarley 1958). Given the abundance
of understory vegetation on the study plot, constructed models
could not discriminate between true refugia and random locations.
Additionally, there is no means of assessing how many of the
random sites were “near” a ground nest.

Importance of woody .-Distance to coarse woody debris

was the most important variable in determining the location of
ground refugia. Woody debris, including fallen logs, snags, and
stumps, is important in the distribution and occurrence of many
small mammalian species (Strecker & Williams 1929; Ivey 1949;
Easterla 1968; Graves et al. 1988). Coarse woody debris serves as
habitat for invertebrates and fungi eaten by small mammals, retains
moisture, and provides important navigational cues (McCay 2000).
The SFAEF has little ground-level structural diversity. Therefore,
ground nesting species have few alternatives to the use of stumps,
root boles, and their associated decomposing root systems in which
to seek refuge. Rock outcroppings are not present and burrows of
larger vertebrates and standing snags were rare on the study plot.
The finite number of suitable ground refuge locations creates an
environment in which competition could play an important role.

EDWARDS & BRADSTREET

111

Distance to coarse woody debris was retained as an important
variable in the microhabitat of golden mice arboreal nests.
Arboreal nests of golden mice typically were closer to woody
debris than would be predicted from random locations. The use of
ground refiigia, in addition to arboreal nests, may provide
alternative means of escape when faced with predation. Packard &
Gamer (1964) reported flushing an adult male golden mouse from a
nest. After 10 min of harassment, the mouse leaped from the vines
and vanished into an underground burrow system from which it
could not be extracted.

Importance of climbing vine-\i was expected that distance to
climbing vines would be an important variable retained in the
discrimination of arboreal nests. Vines provide a common substrate
in which arboreal nests are constmcted (Linzey & Packard 1977).
With one exception, arboreal nests found in this study were either
constmcted entirely within vines or used vines as anchoring support
to a tree or shmb. Vines provide mice with multiple escape
pathways when faced with predation pressure.

Distance to nearest vine suitable for climbing was also retained
in the discrimination model of ground refugia. Ground refugia
were closer to climbing vines than would be expected from random
locations. Golden mice are able climbers and commonly seek
refuge in trees or bushes when released from live-traps (Linzey &
Packard 1977). As mentioned with arboreal nests, constmcting
terrestrial refugia within close proximity to climbing vines may
provide alternative options of escape.

Prior to this research, there have been two studies of golden
mice nesting behavior in eastern Texas. McCarley (1958) found no
evidence of arboreal nesting in eastern Texas, whereas Packard &
Gamer (1964) discovered numerous arboreal nests at two sampling
localities in Nacogdoches County, Texas. It was noted that from
1955 to 1957, Nacogdoches County, Texas experienced the worst
drought on record for this area. Since 1901, the average annual
rainfall reported for Nacogdoches County, Texas is 122 cm. In

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contrast, the average annual rainfall recorded from 1955 to 1957
was 81 cm. Rainfall totals recorded during 1961 and 2004 (the year
this research was conducted) were above average at 137 cm and
173 cm, respectively. Further, approximately 30% of the study plot
established at the SFAEF had standing water for much of the
autumn monitoring season. These results support those of Packard
& Gamer (1964). Golden mice in eastern Texas seemingly use
arboreal nests only during the autumn. The lack of ground-level
stmctural diversity in eastern Texas forests in conjunction with
seasonal flooding may cause terrestrial nest locations to become a
limiting resource for golden mice during autumn. It should be
noted that supplemental nest boxes were provided by McCarley
(1958). The timing of occupation of these nest boxes closely
coincides with the timing of the shift from terrestrial to arboreal
habitats reported by Packard and Gamer (1964) and by this study.
Arboreal nesting seems to be of adaptive value to golden mice
populations in eastern Texas.

Acknowledgments

We gratefully acknowledge the assistance of S. Johnson and S.
Williams for help during fieldwork. We thank P. Blackburn for the
constmction and design of all radio transmitters used in this study.
T. Henry and S. Johnson assisted in production of figures and
tables. For critical reviews of this manuscript, we thank G. A.
Feldhamer, C. Jones, E. McTavish, R. D. Stevens, and Jim Goetze.
Partial funding was provided by a Grants-In-Aid of Research award
from The American Society of Mammalogists (to APB) and by the
Department of Biology at Stephen F. Austin State University.

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CWE at: cedward7@gmu.edu

TEXAS J. OF SCI. 61(3):181-186

AUGUST, 2009

THE SAND DOLLAR PERIARCHUS LYELLl
(ECHINOIDEA: CLYPEASTEROIDA: SCUTELLIFORMES)

IN THE CADDELL FORMATION (UPPER EOCENE) OF TEXAS

Louis G. Zachos

Department of Paleobiology MRC~121
National Museum of Natural History, Smithsonian Institution
P.O.Box 37012, Washington, DC 20013-7012

Abstract. -The occurrence of the sand dollar echinoid Periarchus lyelli in the
Upper Eocene Caddell Formation of Texas is confirmed. This extends the geo¬
graphic range of this Gulf Coast index fossil west from Mississippi to Texas.

The sand dollar echinoid Periarchus lyelli (Conrad) is a
distinctive and well-known element of the fauna of Upper Eocene
(Jackson) marls and limestones across the eastern Gulf of Mexico
and lower Atlantic coastal plain. Hurricane Rita struck the Texas
coast in September, 2005, and caused significant erosion on the
northern shore of the Sam Rayburn Reservoir and created fresh
exposure of sandstones and shales of the Upper Eocene Caddell
Formation in the immediate vicinity of the type area. A number of
specimens of Periarchus lyelli were collected from these exposures
following the storm. This report documents a geographic range
extension of the species west from Mississippi to Texas.

Study Area and Methods

The Caddell Formation was named by Dumble (1915) after the
town of Caddell in southern San Augustine County, Texas (Figure
1). The town site was inundated by the Sam Rayburn Reservoir in
the 1960s, although the cemetery remains. The type section was
determined by Eargle (1959) and the locality was redescribed by
the Gulf Coast Section of Economic Paleontologists and
Mineralogists (1966) before inundation by the reservoir. The old
town site is near the Harvey Creek Recreation Area at the end of
FM2390 about 10 miles (by road) from Broaddus. There are poorly
preserved fragments of sand dollar echinoids collected from this
area in the Rio Bravo Collection (Molineux 2008) at the Texas

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Periarchus lyelli

5 10 15 Kilometers

Fig. 1. Study Area. (A) Gulf of Mexico and Atlantic coastal plain showing Upper Eocene
occurrences of Periarchus lyelli. Each circle represents a county with one or more
localities where P. lyelli has been reported. (B) San Augustine County, Texas. The
Harvey Creek Recreation area is marked by a box located at 31°12’30” N, 94°15’45’'
W. (C) Harvey Creek Recreation Area on the northeast shore of Sam Rayburn
Reservoir. Type locality for the Caddell Formation, which outcrops along the
outlined shoreline. Each triangle marks a location from which P. lyelli was collected
in situ. (The formation dips towards the southeast).

Natural Science Center. The original collecting locality for these
specimens, as recorded by specimen labels more than century old,
was in a southward- facing slope of the valley of the Angelina River
at Caddell. Loose cobbles and boulders of indurated sandstone
contained fragments of sand dollars. In September, 2005, the eye
of Hurricane Rita crossed the area and caused significant erosion on
the northern shore of Sam Rayburn Reservoir. This erosion
exposed fresh sections of sandstone and shale of the Caddell

ZACHOS

183

Formation. Outcrops along the irregular shoreline are now best
accessed by boat, and the exposure of the formation extends from
the boat ramp at the Harvey Creek Recreation Area to
approximately 3 km southeastward, exposing a stratigraphic
interval about 35 m thick. The lowest interval of the section is
composed of interbedded argillaceous, iron oxide impregnated
sandstone and sandy clay, with trace fossils and poorly preserved
(moldic) megafauna. This is overlain by a fine-grained unit with
calcareous lenses several meters in diameter containing the large
oyster Crassostrea gigantissima (Finch). A thin layer of
calcareous, very shelly sand less than a meter thick overlies the
shale, grading upwards into indurated sandstone which weathers
into large concretionary masses. Well-preserved fragments (Fig.
2a) of Periarchus lyelli are found in the thin calcareous unit and
relatively whole specimens (Fig, 2b) occur on the weathered
surfaces of the upper sandstone. Both occurrences are associated
with unidentified nummulitid foraminifera.

Material Examined

All fossil material collected has been cataloged in the collections
of the Nonvertebrate Paleontology Laboratory (NPL) of the Texas
Natural Science Center (TNSC), on the Pickle Research Campus of
the University of Texas at Austin.

Discussion

Sand dollar echinoids, members of the order Clypeasteroida,
first appeared in the fossil record of the Gulf of Mexico coastal
plain in the early part of the Middle Eocene. The oldest known
sand dollars from this region are the putative species Protoscutella
mississippiensis found near the top of the Tallahatta Formation in
western Alabama. Various nominal species of Protoscutella are
found in abundance in slightly younger deposits from Texas to
North Carolina. Periarchus lyelli first appeared in the Middle
Eocene and by the Late Eocene it became the predominant species
of sand dollar. Oddly, although the oldest occurrence of P. lyelli is

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Fig. 2. Periarchus lyelli from the Caddell Formation, Sam Rayburn Reservoir. (A)

TNSC NPL19819, from shelly calcareous bed. (B,C) TNSC NPL19815, aboral and

left side, from concretionary sandstone boulder. Scale bars 1 5 mm.

in the Cook Mountain Formation in east Texas, the species was not
definitely known from the Upper Eocene Jackson Group in Texas.
Fragmentary evidence in the TNSC Rio Bravo Collection was
suggestive of the occurrence of the species, but is not definitive.

Periarchus lyelli was first described by Conrad (1834) from
(probably) the Moodys Branch Formation below Claiborne Bluff on
the Alabama River (Monroe County, Alabama). The species is
characterized by significant morphologic variability and widespread
geographic distribution. This has resulted in several closely related
species being described then variously reduced to synonyms or
subspecies of P. lyelli by different authors. The species is especially
abundant in the Moodys Branch Formation and equivalents from
Mississippi east and north to North Carolina and as far south as
central Florida. It has not been found in the Moodys Branch
Formation exposures along the Quachita River in central Louisiana
(Huner 1939), but Cooke (1942; 1959) reported that fragments of sand

ZACHOS

185

dollars attributable to this species were found by T,W. Vaughan in the
Moody s Branch Formation exposed at Montgomery Landing (Creole
Bluff) on the Red River in Louisiana (now inundated by water
dammed behind navigational locks on the river). The species is not
listed in the fauna from Montgomery Landing in Schiebout & van den
Bold (1986), but Vaughan’s specimens (27 small fragments) are in the
Smithsonian paleobiology collections, catalog number USNM
164680, and do appear to be attributable to P. lyelli. Clark &
Twitchell (1915) reported some occurrences of P. lyelli in Texas, but
all of these were misidentifications of Protoscutella mississippiensis.
The occurrence of P. lyelli noted by Cooke (1959) is from a Cook
Mountain Formation locality in Sabine County, Texas (see Zachos &
Molineux 2003). A probable identification of P. lyelli for the material
in the Rio Bravo collection was reported by Zachos & Molineux
(2003), but the latest collections reported here were taken in situ from
the Caddell Formation and confirm the identification.

Better preserved specimens collected from the Caddell Formation
show the conical “Chinese haf ’ profile characteristic of the putative
subspecies P. lyelli pileussinensis (Figure 2c). This variety is
generally found in Jackson (Bartonian) deposits and is evidence that
the Texas occurrence is of equivalent age to the Moody s Branch
Formation or younger deposits in the eastern Gulf of Mexico coastal
plain.

Carter et al. (1989) showed that Periarchus lyelli in all its variants
preferred a sand substrate. Carter & McKinney (1992) presented
evidence that distribution patterns in Upper Eocene echinoid faunas
were related to facies patterns, particularly in relation to sand/mud
ratios. The extension of the geographic range westward into Texas
demonstrates that P. lyelli had a preference for sandy substrate but
was tolerant of composition (quartz/glauconite vs. calcite/aragonite)
and minor variation in water temperature represented by differences in
environmental regimes that included the Atlantic coast of the
Carolinas and the carbonate platform of Florida.

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Acknowledgments

Appreciation is extended to A. Molineux, Texas Natural Science
Center, Austin for assistance with the collections of the Non-
Vertebrate Paleontology Laboratory, and to C. Ciampaglio and T.
Yancey for their critical review of the manuscript. This work was
supported by the Geology Foundation of the Jackson School of
Geosciences at the University of Texas at Austin., and by the
Smithsonian Institution Fellowship Program.

Literature Cited

Carter, B. D., T. H. Beisel, W. B. Branch & C. M. Mashbum. 1989. Substrate
preferences of Late Eocene (Priabonian/Jacksonian) echinoids of the eastern Gulf
Coast. Jour. Paleo., 63:495-503.

Carter, B. D. & M. L. McKinney, 1992. Eocene echinoids, the Suwannee Strait, and
biogeographic taphonomy. PaleobioL, 18:299-325.

Clark, W. B. & M. W. Twitchell. 1915. The Mesozoic and Cenozoic Echinodermata of
the United States. U.S. Geol. Surv. Monograph 54, 341 pp.

Conrad, T. A. 1834. Descriptions of new Tertiary fossils from the southern states. Jour.

Acad. Nat. Sci. Philadelphia, Ser. 1, v. 7:130-157.

Cooke, C. W. 1942. Cenozoic irregular echinoids of eastern United States. Jour. Paleo.,
16:1-62.

Cooke, C. W. 1959. Cenozoic echinoids of eastern United States. U.S. Geol. Surv. Prof
Paper 321, Hi + 106 pp.

Dumble, E. T. 1915. Problem of the Texas Tertiary sands. Geol. Soc. Am. Bull. 26:447-
476.

Eargle, D. H. 1959. Stratigraphy of Jackson Group (Eocene), south-central Texas.

Amer. Assoc. Petrol. Geol. Bull. 43:2623-2635.

Gulf Coast Section of the Society of Economic Paleontologists and Mineralogists. 1966.
Caddell Formation (U. Eocene) Type Locality, GCS of SEPM Type Localities
Project Unit IV. Gulf Coast Assoc. Geol. Soc. Trans, 16:393-394.

Huner, J. 1939. Geology of Caldwell and Winn Parishes. Louisiana Geol. Surv. Bull.
15,356 pp.

Molineux, A. 2008. The Rio Bravo Collection: Preserving a unique collection for future
research in the Gulf Coast section. Gulf Coast Assoc. Geol. Soc. Trans. 58:699-700.
Schiebout, J. A. & W. van den Bold. 1986. Montgomery Landing Site, Marine Eocene
(Jackson) of Central Louisiana, Thirty Sixth Annual Meeting of the Gulf Coast
Association of Geological Societies, Baton Rouge, vi + 238pp.

Zachos, L. G. & A. Molineux. 2003. Eocene echinoids of Texas. Jour. Paleo. 77:491-
508.

LZ at: lg_zachos@alumni.utexas.net

TEXAS T OF SCI. 6 1(3): 187 A 94

AUGUST, 2009

MIXED INFECTIONS OF NASOPHARYNGEAL BOTS,
CEPHENEMYIA SPP. (OESTRIDAE) IN WHITEMAILED DEER
{ODOCOILEUS VIRGINIANUS) AND MULE DEER
(ODOCOILEUS HEMONUS) OF TEXAS

Samuel W. KeOey

U.S. Geological Survey
Wichita Falls, Texas 76308

Ahsir^ti -Cephenemyia spp. are widespread oestrid flies whose larvae parasitize
various cervid hosts, yet attempts to pathologize their significance as well as
delineate their taxonomy, dispersal potential, and distribution remain ambiguous.
This report provides new records of mixed Cephenemyia spp. infections in both mule
deer (Odocoileus hemionus) and white-tailed deer (Odocoileus virginianus) in west
and north-central Texas, including C, jellisoni, C, phobifer, and C pratti, thereby
filling a substantial void in previous southern distribution reports. Possible
ecological and pathological implications are discussed including the need for a
taxonomic review of the genus Cephenemyia plus dispersal, hybridization, and vector
potential for chronic wasting disease in cervid hosts.

Members of the genus Cephenemyia (Diptera: Oestridae) are
stout bee-like flies (Fig. 1) whose larvae parasitize the nasal
passages and pharyngeal regions of a variety of cervid hosts within
the Holarctic ecozone. Gravid females of most North American
Cephenemyia species are believed to larviposit into the nostrils of
their hosts (Golini et al. 1968; Anderson 2001), where larvae
migrate, feed, and grow within various nasopharyngeal regions of
the host including the retropharyngeal pouches, soft palate, under
the tongue, glottis, and throughout the nasal passages. Mature third
instars travel out of the nostrils and pupate in the soil with adult
emergent times variable and purportedly dependent upon ambient
temperatures (Bennett 1962; Hair et al. 1969; Nilssen 1997).

While Cephenemyia spp. infections are known to cause
deleterious effects within cervid hosts, their total impact upon
cervid ecology remains unclear. Dated reports indicate that the bots
may cause severe pathogenesis including death of the hosts (Walker
1929; Bruce 1931; Cowan 1946); however, more recent studies
report gadding and physiological trauma but downplay the risk of

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 3, 2009

Figure 1. Adult Cephenemyia jellisoni reared from a larva found in a mule deer

{Odocoileus hemionus) from Jeff Davis County, Texas.

host death or debilitation (Cogley 1987; McMahon & Bunch 1989;
but see Johnson et al. 1983). A recent study by Lupi (2005) posited
that Cephenemyia spp. could serve as vectors for chronic wasting
disease (CWD) to cervids due to their proximity to host structures
potentially rich in prion rods and their ability to replicate and
express prion proteins. Ingestion of similar encephalopathic-
infected fly larvae and pupae has caused rodent host infections
within a previously infection-free sample (Post et al. 1999).

Four species of Cephenemyia have been documented in the
south-central and southeast-central regions of Texas (Fig. 2): C.
pratti Hunter 1915, C phobifer (cf. Van Volkenberg & Nicholson
1943; Bennett & Sabrosky 1962), C. jellisoni (cf. Bennett &
Sabrosky 1962), and C. albina Taber & Fleenor 2004; Fleenor &
Taber 2007. Samuel & Trainer (1971) also described larvae

KELLEY

189

Figure 2. Regional distribution by state and county of Cephenemyia spp. in Texas,
Oklahoma, and New Mexico based on published records. Cephenemyia spp. from
Wilbarger and Jeff Davis counties, Texas represent new locality records in the state.

encountered in the southern Gulf Coast region of the state but could
not positively identify the species. Adjoining state records for
Cephenemyia include C. jelUsoni from eastern Oklahoma (Hair et
al. 1969) and C. jelUsoni plus C. pratti from New Mexico
(Townsend 1941; Bennett & Sabrosky 1962). Cephenemyia
phobifer occurs in Louisiana (Bennett & Sabrosky 1962; Kellogg et
al. 1971), yet none were found in a large sample of deer {n= 151)
examined from Arkansas (Kellogg et al. 1971). Weber (1992)
states that C. pratti is the only known species from Mexico, though
other Cephenemyia species are thought to occur in northern Mexico

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 3, 2009

as well. This study reports the first records of Cephenemyia spp.
from north-central and west Texas.

Methods

In February 2008 a Cephenemyia-infQciQd white-tailed deer
{Odocoileus virginianus) was found dead 13 km S of Vernon,
Wilbarger County, Texas (34^01’ 12” N, 99^ 14' 31” W; elev. 372
m) in an area predominately used for cattle rangeland and crop
cultivation. Cause of death was unknown, as it was estimated to
have been dead for two days and was heavily scavenged.

Subsequently, in December 2008, a Cephenemyia-mfQCtQd mule
deer {Odocoileus hemionus) was shot in the Davis Mountains, 21.7
km SE of Ft. Davis, Jeff Davis, County, Texas (30° 31' 01" N, 103°
44' 16" W; elev. 1557 m) in an area primarily reserved for hunting
activities.

Both male deer were aged following traditional tooth wear
methods (Severinghaus 1949). Live third instar larvae from both
cervid hosts were placed into separate glass containers half full of
sand, covered with light leaf litter, and kept indoors (21 C) for 48
h to facilitate pupariation. Larvae failing to undergo pupariation
were identified using available keys (Bennett & Sabrosky 1962)
and stored in 80 percent ethanol. Larval specimens were deposited
in the U.S. National Parasite Collection (USNPC), Beltsville,
Maryland (101515-101517). An emergent adult specimen was
identified and deposited in the Midwestern State University insect
collection, Wichita Falls, Texas.

Results

The infected white-tail deer’s estimated age was 8.5 yrs.
Eighteen Cephenemyia spp. larvae were found within the nasal
passages and back of the pharynx in addition to one puparia found
within the nasal passages. The mule deer was aged at 9.5 yrs, and

KELLEY

191

three third instar Cephenemyia larvae were noticed crawling out of
the nose after the deer had been in cold storage for two days. A
subsequent examination revealed no additional larvae, and it is
unknown if any larvae escaped before the head was examined.
Regarding bot larvae from the white-tail deer, none successfially
pupated except the puparia initially discovered within the nasal
passages. Dissection of the whitetail-host puparia after 10 months
revealed a moldy, degenerated, unidentifiable specimen. One of
three mule deer-host larvae successfully pupated, and the adult
emerged 27 days later.

The emergent mule deer-host bot fly was identified as C.
jellisoni. The other two mule deer-host larvae were identified as C.
pratti, indicating a mixed infection. Of the 18 larvae from the
white-tail deer in Wilbarger County, 15 were identified as C.
jellisoni (83%), and three as C phobifer (17%), providing another
example of mixed Cephenemyia spp. infection within the same
host. All specimens of Cephenemyia jellisoni, C. pratti, and C
phobifer represent new geographic records within Texas (Fig. 2).

Discussion

Distributions of several species within the genus Cephenemyia
overlap, and mixed infections among cervid hosts may be common,
but comprehensive ranges of Cephenemyia spp. remain unknown.
A major hindrance to the study of Cephenemyia spp. distributions
and their ecology stems from the paucity of captured adult
specimens. Adult botflies have morphological characteristics (hair
patterns, density, and color, plus wing infuscation, and arista
coloration) that allow for easier identification than larvae. Larvae
are encountered far more frequently in dead cervid hosts but are
practically impossible to rear to maturity if they fail to pupariate
soon after removal from the host. Identification of third instar
larvae can prove troublesome as key larval characteristics (spine
counts, anterior spiracle morphology) sometimes vary or overlap
among described species; moreover, a single larva is often
inadequate for species identification (Bennett & Sabrosky 1962).

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Nevertheless, larval specimens encountered in this study aligned
closely on average with descriptions in available keys.

Previously published Cephenemyia spp. vouchers provide vague
zonal boundaries for tentative identifications; however, dispersal
potential of these volant insects and their cursorial-saltatorial hosts
is large. Dispersal of bots may be compounded by the frequency of
exotic cervid spp. introductions and relocations. Additionally,
hybridization among sympatric Cephenemyia spp. may be possible
as it is in other dipterans (Stevens & Wall 1996; Dos Santos et al.
2001). It is unknown what morphological effect such hybridi¬
zations could have on both larval and adult specimens, though
phenotypic intergrades seem likely. Mixed infections of
Cephenemyia spp. in cervids may also confound some efforts at
larval species identification. Hence, the genus Cephenemyia
remains in a confused taxonomic state (Cogley 1991).

Taxonomically, the genus Cephenemyia could benefit greatly
from a thorough phenotypic and genotypic review, and additional
molecular studies may generate new information regarding
differences among species. Furthermore, field studies could be
conducted regarding cervid host ingestion of Cephenemyia larvae
and pupae exposed to CWD prions or larvipositing within CWD-
free deer by adult botfly females that formerly parasitized CWD-
infected deer as larvae. Such research may reveal what role, if any,
Cephenemyia spp. play in the transmission of CWD. Presently, a
control plan for wild herds is unfeasible, yet Ivermectin® has
proved 100% effective in Cephenemyia removal from captured
white-tailed deer in Mexico (Weber 1992) and may prove a useful
treatment in epidemiological areas among small captive herds.

Acknowledgments

A special thanks is given to Roy Vogtsberger of Midwestern
State University for all of his helpful commentary.

KELLEY

193

Literature Cited

Anderson, J. R. 2001. Larviposition by nasopharyngeal bot fly parasites of
Columbian black-tailed deer: a correction. Med. Vet. EntomoL, 15:438-442.
Bennett, G. F. & C. W. Sabrosky. 1962. The nearctic species of the genus
Cephenemyia (Diptera, Oestridae). Can. J. ZooL, 40:431-448.

Bennett, G. F, 1962. On the biology of Cephenemyia phobifera (Diptera: Oestridae),
the pharyngeal bot of the white-tailed deer, Odocoileus virginianus. Can. J.
ZooL, 40:1195-1210.

Bruce, E. A. 1931. Records of insects affecting wildlife in B. C. Rep. Vet. Dir. Gen.,
Can. Dep. Agr. Append., 5:71-73.

Cogley, T. P. 1987. Effects of Cephenemyia spp. (Diptera: Oestridae) on the
nasopharynx of black-tailed deer {Odocoileus hemionus columbianus). J. Wildl.

Dis., 23:596-605.

Cogley, T. P. 1991. Distribution of bot larvae, Cephenemyia spp. (Diptera:
Oestridae) from white-tailed deer {Odocoileus virginianus) in Florida. Florida
EntomoL, 74:479-481.

Cowan, I. M. 1946. Parasites, diseases, injuries, and anomalies of the Columbian
black-tailed deer, Odocoileus hemionus columbianus (Richardson) in British
Columbia. Can. J. Res., 24:71-103.

Dos Santos, P., K. Uramoto & S. R. Matioli. 2001. Experimental hybridization
among Anastrepha species (Diptera: Tephritidae): production and morphological
characterization of Fi hybrids. Ann. EntomoL Soc. Am., 94:717-725.

Fleenor, S. B. & S. W. Taber. 2007. Description of the female and first-instar larva
of a new bot fly species from central Texas. Southwest. EntomoL, 32:37-41.
Golini, V. L, S. M. Smith & D. M. Davies. 1968. Probable larviposition by
Cephenemyia phobifer (Clark) (Diptera: Oestridae). Can. J, Zoo., 46:809-814.
Hair, J. A., D. E. Howell, C. E. Rogers & J. Fletcher. 1969. Occurrence of the
pharyngeal bot Cephenemyia jellisoni in Oklahoma white-tailed deer, Odocoileus
virginianus. Ann. EntomoL Soc. Am., 62:1208-1210.

Hunter, W. D. 1915. A new species of from the United States. Proc.

EntomoL Soc. Wash., 17:169-173.

Johnson, J. L., J. B. Cambell, A. R. Doster, G. Nason & R. J. Cagne. 1983. Cerebral
abscess and Cephenemyia phobifer in a mule deer in central Nebraska. J. Wildl.
Dis., 19:279-280.

Kellogg, F. E., T. P. Kistner, R. K. Strickland & R. R. Gerrish. 1971. Arthropod
parasites collected from white-tailed deer. J. Med. Ent., 8:495-498.

Lupi, O. 2005. Risk analysis of ectoparasites acting as vectors for chronic wasting
disease. Med. Hypotheses, 65:47-54.

McMahon, D. C. & T. D. Bunch. 1989. Bot fly larvae {Cephenemyia spp.,
Oestridae) in mule deer {Odocoileus hemionus) from Utah. J. Wildl. Dis.,

25:636-638.

Nilssen, A. C. 1997. Effect of temperature on pupal development and eclosion dates
in the reindeer oestrids Hypoderma tarandi and Cephenemyia trompe (Diptera:
Oestridae). Environ. EntomoL, 26:296-306.

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Post, K., D. Riesner, V. Walldorf & R. Melhom. 1999. Fly larvae and pupae as
vectors for scrapie. Lancet 122:199-204.

Samuel, W. M. & D. O. Trainer. 1971. Pharyngeal botfly larvae in white-tailed deer.
J. Wildl. Dis., 7:142-146.

Severinghaus, C. W. 1949. Tooth development and wear as criteria of age in white¬
tailed deer. J. Wildl. Manage., 13:195-216.

Stevens, J. & R. Wall. 1996. Species, sub-species and hybrid populations of the
blowflies Lucilia cuprina and Lucilia sericata (Diptera: Calliphoridae). Proc. R.
Soc. Lond.B., 263:1335-1341.

Taber, S. W. & S. B. Fleenor. 2004. A new bot fly species (Diptera: Oestridae) from
central Texas. Great Lakes Ent., 37:76-80.

Townsend, C. H. 1941. An undescribed American Cephenemyia. J. N.Y. Entomol.
Soc., 49:161-163.

Van Volkenberg, H. L. & A. J. Nicholson. 1943. Parasitism and malnutrition of deer
in Texas. J. Wildl. Manage., 7:220-223.

Walker, C. R. 1929. Cephenomyia s^^.kxXXmgdLQox. Science 69:646-647.

Weber, W. 1992. Valoracion clinica del efecto de la Ivermectina contra
Cephenemyia spp en venados cola blanca. Vet. Mex., 23:239-242.

SWK at: skelley@usgs.gov

TEXAS J. SCL 61(3): 195-202

AUGUST, 2009

KARYOTYPE DIVERSITY AMONG AND WITHIN AVIAN TAXA:

A SIMPLE TEST IN R

Michael F. SmaO, Michael R. J. Forstner and John T. Baccus

Department of Biology, Wildlife Ecology Program
Texas State University - San Marcos
San Marcos, Texas 78666

Abstract -Taxonomic categories should be representative of natural groupings
based on a shared lineage. Approaches to implementing this tenet are numerous and
diverse with molecular methods such as gene sequencing being the most common
today. This study suggests that one approach, the use of diploid chromosome
characteristics, has been underutilized in avian species, particularly compared to other
higher taxa. It is further suggested that a primary cause of this lack of information is
because avian chromosomes are more difficult to study than other vertebrate groups.
Consequently, a common statistical method is demonstrated to determine if variation
in diploid chromosome number is greater between avian taxonomic groups than
within them, as would be expected. An example is demonstrated using the powerful,
free software package Program R. Implications of these results are also discussed.

Resumen.-Categorias taxonomicas deberian ser representatives de los grupos
sobre la base de un linaje compartido. Enfoques para la aplicacion de este principio
son multiples y con diversos metodos moleculares como la secuenciacion de genes es
el mas comun el dia de hoy. Sugerimos que un enfoque, el uso de cromosomas
diploides caracteristicas, ha sido subutilizado en las especies de aves, especialmente
en comparacion con otros taxa superiores. Ademas, sugerimos que una de las
principales causas de esta falta de informacion se debe a que los cromosomas aviar
son mas dificiles de estudiar que otros grupos de vertebrados. En consecuencia,
demostrar un metodo estadistico comun para determinar si la variacion en el numero
diploide de cromosomas aviar es mayor entre los grupos taxonomicos que dentro de
ellos, como era de esperar. Demostramos esta usando el gran alcance, libre del
paquete de software R. Programa Consecuencias de nuestros resultados son tambien
discutidos.

A karyotype is a characterization of an entire set of
chromosomes from an individual with regard to number, size, and
shape (Shields 1982). For more than 30 years researchers have
acknowledged that karyotypes represent part of an individuaTs
phenotype (Chiarelli & Capanna 1973) alongside its inherent link to
the genome. With more recent technological advances in
karyological techniques, this is even truer today.

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

The purpose of this paper is intended as both informative with
regard to application of a simple but efficient mechanism to
evaluate genomic diversity and attempts to address and integrate
two distinct issues that are believed to be of valuable as approaches
in avian karyology and statistical analysis that have been ignored
and often misrepresented. Foremost among concerns is emphasiz¬
ing the gap in karyological knowledge of avian chromosomes
versus other vertebrate taxa and second demonstrating the useful¬
ness and flexibility of the free statistical package R.

Avian chromosomes -OnQ critical and often overlooked aspect
of the advances in genomics has been, and remains, the dearth of
information for avian karyotypic and even general cytogenetic data.
Shields (1982) was among the first to point out that avian cyto¬
genetics research effort has lagged behind that of other vertebrate
taxa. The authors are in agreement, and believe this is still the case
for a very fundamental reason. Avian karyotypes are more difficult
to work with than other vertebrate taxa.

There are fewer studies of avian karyology than in other craniate
groups (De Boer & Van Brink 1984). From about 1970 to 1980 the
number of identified avian karyotypes doubled (Shields 1982).
However, at that time, only about 5-10% of the about 9,000 extant
species of birds had been karyotyped, many poorly (Shields 1983).
Although progress has undoubtedly been made in elucidating
karyotypes of avian species in recent years, to what degree remains
undocumented. Shields (1982) also pointed out that no compre¬
hensive cytogenetic analysis of a population of any wild bird
species has been undertaken. This is currently still correct.

The descriptions of bird karyotypes are inherently more proble¬
matic than those of other vertebrate groups because of higher
chromosome numbers, small numbers (8~15 pairs) of macro¬
chromosomes, and 30-40 pairs of microchromosomes. Makino et
al. (1956) described a diploid number of 80 as definitive for
mourning doves {Zenaida macroura), however, Benirschke & Hsu

SMALL, FORSTNER & BACCUS

197

(1971) found In = 78. Microchromosomes are difficult to accurate¬
ly count because of small size and morphological uniformity. It is
not uncommon for the diploid number to be reported within a range
because of differences in number of microchromosomes (usually ±
2) between cells. Small et al. (1993) reported finding three distinct
diploid numbers, not in a single species, but in the same tissue from
a single individual. This was not the result of difficulties in
technique, as numerous, well spread karyotypes were examined,
and meiotic karyotypes showed similar variability (Small 1991).
As the case in numerous species, the difference was in the number
of microchromosomes.

Consequently, in general, current cytogenetic studies of avian
chromosomes focus on specific macrochromosomes, which are
highly conserved (Stock & Mengden 1975, Derjusheva et al. 2004)
or sex chromosomes (Ellegren 2000). However, microchromo¬
somes are not unimportant components of the avian karyotype,
particularly given that nucleolus organizer regions in avian species
are located on microchromosomes (Small 1991). This is supported
by recent work on the chicken genome project which shows that
microchromosomes are gene rich and subsequently important
components of avian genomes (Douaud et al. 2008).

Program /^.-Program R is essentially a free, downloadable
version of S-Plus (Insightfiil Corporation). In our experience it is
an extremely powerful program for ecological analyses with
excellent graphing capabilities. However, despite recent commer¬
cial efforts (Dalgaard 2002, Maindonald & Braun 2006), no
comprehensive, downloadable user’s guide exists for Program R.
Consequently, this study presents a detailed example of application
of R to encourage researchers and students to make an effort to
learn and use this program.

Integrating R and ATaTjo/og^y.-Systematists have classified
species into discrete taxonomic groups in an effort to facilitate an
understanding of the structure and nature of diversity. Therefore,

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

classification of species into taxa has attempted to separate
organisms into natural groups. Additionally, taxonomy today
explicitly seeks to group taxa based on their evolutionary history
(i.e., phylogeny; Cracraft 1981). Consequently, the nature of
karyotypes makes it reasonable to expect them to reflect that natural
relationship. Thus, lower variation within taxonomic groups than
between them, in regard to karyotype should be expected.

Methods

Karyotype character variation can be analyzed using a linear
mixed effects (Ime) model with taxa (i.e., family, order, etc.) as a
random factor, karyotypic characteristic (i.e., diploid number,
fundamental number, etc.) as the response variable, and species as
replicates. This allows partitioning of sources of variation in an
unbalanced data set. Data can be analyzed using Program R
computer software (R Development Core Team 2005) with
maximum likelihood estimates in all analyses. Program R uses
restricted maximum likelihood estimates as the default for Ime
models, which may yield slightly different results than other
software if not taken into account. Unrestricted maximum
likelihood estimates are used in this example.

An example using Program R cor/e.-Below is an example model
code using diploid numbers from six avian families (this is for
illustration so sample sizes are small). Explanations are given in
italics bounded by # symbols:

> diploid.number<-scan()

# creates a database to store the diploid numbers #

# the number preceding the colon denotes the numerical level of the next entry #

1: 76 80 80 76 76 80 80 76 76 76 76 76 76 76 86 78 76 78 74 78 76 76 78 68

# Columbidiae #

25:86 78 74 76 76 76 76 80 80

# Troglodytidae #

34: 78 52 68 52 68 60 60 68 72 56

SMALL, FORSTNER & BACCUS

199

# Ciconiidae #

44: 78 92 80 82 82 80

# Anatidae #

50: 54 48 48 52 52 86

# Falconidiae #

56: 92 92 84 108 108 94 88 92 90 92

# Picidae #

66:

Read 65 items

# data is entered in separate strings for each taxa (in this case ‘family”) #

> family<-factor(rep(LETTERS[l:6],c(24,9,10,6,6,10)))

# creates a sequence of letters A-F designating families such that there are 24
“A ”s, 9 “B ”s, etc. #

> number<-data.frame(family,diploid.number)

# creates a matrix of families and diploid numbers #

> diploid<-

lme(diploid.number~l,data=number,random=~l I family, method="ML")

# creates a data file to store the model information; Ime tells R to use linear
mixed-effects model and method=”ML” tells R to use unrestricted maximum
likelihood #

> summary(diploid)

# displays output of the analysis #

Example data output:

Linear mixed-effects model fit by maximum likelihood

Data: number

AlC BIG
460.9535 467.4767

logLik

-227.4768

Random effects:

Formula: ~1 | family

(Intercept) Residual

StdDev: 11.95630 6.842518

Fixed effects: diploid.number ~ 1

Value Std.Error

(Intercept) 75.27702 5.009927

DF t-value p-value

59 15.02557 0

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Standardized Within-Group Residuals:

Min Q1 Med

-1.7211003 -0.4350450 -0.1427550

Q3 Max

0.3062634 4.1461398

Number of Observations: 65
Number of Groups: 6

>intervals(lme(diploid.number~l,data=number,random=~l|family,method="ML"))
# provides 95% Confidence Intervals #

Approximate 95% confidence intervals

Fixed effects:

lower

(Intercept) 65.3296
attr(, "label")

[1] "Fixed effects:"

Random Effects:

Level: family

lower

sd((Intercept)) 6.629736

est.

75.27702

upper

85.22445

est.

11.95630

upper

21.56240

# among group variance and 95%) confidence intervals #

Within-group standard error:

lower est. upper

5.712406 6.842518 8.196206

# within group variance and 95% confidence intervals #

Results

The output of this model gives intercept and residual standard
deviations. Squaring the intercept standard deviation gives among
taxa variance and squaring the residual standard deviation gives
within taxa variance. From these data, percent variance attributable
to each source can be calculated as well as confidence intervals. In
this example (11.95630^ [(11.95630^+(6.842518^)] gives percent
variance among families and 1-(1 1.95630^)/ [(11.95630^+

(6.842518^)] gives percent variance within familes, 75.3 and 24.
7%, respectively.

SMALL, FORSTNER & BACCUS

201

Discussion

One of the key difficulties in examining generalized trends in
karyotypic evolution is sample size. In the example presented here,
the vast majority of variance, 75%, is attributable to among family
variation, yet there is no significant difference at a < 0,05. Thus, a
power analysis may be warranted to demonstrate either a more
appropriate sample size or a-level.

Another consideration in using this test is determining which
karyotypic characteristic is appropriate for analysis. In groups with
highly conserved karyotypes, such as birds (Derjusheva et al.
2004), differences in diploid number likely result from centromeric
fissions and fusions (Burt 2002). In such cases, ftindamental
number may be a more biologically relevant measure of variance
than diploid number.

This example describes a relatively simple statistical test to
partition variation in karyotypes. This paper is intended as an
example of a potentially useful technique and is not intended to
address all considerations researchers should consider (e.g.,
homoplasy). It is not intended to suggest that karyology is a better
method than analysis of mitochondrial or nuclear DNA, but is
unique and complimentary, and certainly should be considered
when addressing issues of taxonomy. Consequently, as presented,
the methodology is applicable to any appropriate data set.

Acknowledgments

The authors thank F. W. Wekerly, B. S. Cade, G. V. Roslik, and
A. P. Kryukov, and an anonymous reviewer for helpful and
encouraging comments on drafts of this manuscript.

Literature Cited

Benirschke, K. & T. C. Hsu. 1971. Chromosome atlas: fish, amphibians, reptiles,

and birds. Springer- Verlag, New York, 208 pp.

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Burt, D. W. 2002. Origin and evolution of avian micrchromosomes. Cytogenet. and
Genome Res., 96:97-1 12.

Chiarelli, A. B. & E. Capanna. 1973. Cytotaxonomy and vertebrate evolution.
Academic Press, New York 783 pp.

Cracraft, J. 1981. Toward a phylogenetic classification of the recent birds of the
world (Class Aves). Auk, 98:681-714.

Dalgaard, P. 2002. Introductory statistics with R. Springer, New York, NY, 364 pp.

Douaud, M., K. Feve, M. Gams, V. Fillon, S. Bardes, D. Gourichon, D. A. Dawson,
O. Hanotte, T. Burke, F. Vignoles, M. Morrison, M. Tixier-Boichard, A. Vignal
& F. Pitel. 2008. Addition of the microchromosome GGA25 to the chicken
genome sequence assembly through radiation hybrid and genetic mapping. BMC
Genomics, 9: 129. Published online 2008 March 17. doi: 10.1186/1471-2164-9-
129.

De Boer, E. M. & J. M. Van Brink. 1982. Cytotaxonomy of the Ciconiiformes
(Aves), with eight species new to cytology. Cytogenet. Cell Genet., 34:19-34.

Derjusheva, S., A. Kuganova, F. Habermann & E. Gaganskya. 2004. High
chromosome conservation detected by comparative chromosome painting in
chicken, pigeon and passerine birds. Chromosome Res., 12: 715-723.

Ellegren, H. 2000. Evolution of the avian sex chromosomes and their role in sex
determination. Trends in Ecology & Evolution, 15:188-192,

Makino, S., T. Udagawa & Y. Yamashina. 1956. Karyot3T>ic studies in birds. 2: A
comparative study of chromosomes in the Columbidae. Caryologia, 8:275-293.

Maindonald, J. & J. Braun. 2006. Data Analysis and Graphics Using R: An
Example-based Approach. Cambridge University Press, Cambridge, U.K.

R Development Core Team (2005). R: A language and environment for statistical
computing. R Foundation for Statistical Computing, Vienna, Austria.
http://www.R-project.org.

Shields, G. F. 1982. Comparative avian C34ogenetics: a review. Condor, 84:45-58.

Shields, G. F. 1983. Bird chromosomes. Pp. 189-209, in Current Ornithology, (R.
F. Johnson, ed.). Plenum Press, New York. 1 : 1-402.

Small, M. F. 1991. The karyology of the white- winged dove {Zenaida asiatica) in
Texas. Unpublished M.S. thesis, Sul Ross State Univ., Alpine, Texas, 88 pp.

Small, M. F., K. M. Hogan & J. F. Scudday. 1993. The karyotype of the White¬
winged Dove. Condor, 95:1051-1053.

Stock, A. D. & G. A. Mengden. 1975. Chromosome banding pattern conservatism in
birds and nonhomology of chromosome banding patterns between birds, turtles,
snakes, and amphibians. Chromosoma, 50(l):69-77.

MFS at: ms81@txstate.edu

TEXAS I OF SCI. 61(3):203=-218

AUGUST, 2009

THE ARKANSAS ENDEMIC FAUNA:

AN UPDATE WITH ADDITIONS, DELETIONS,

A SYNTHESIS OF NEW DISTRIBUTIONAL RECORDS,

AND CHANGES IN NOMENCLATURE

Chris T* McAllister, Henry W. Robison and Michael E. Slay

RapidWrite, 102 Brown Street, Hot Springs National Park, Arkansas 71913
Department of Biology, Southern Arkansas University
Magnolia, Arkansas 71754 and
The Nature Conservancy, 601 North University Avenue
Little Rock, Arkansas 72205

Abstract -This study provides an update to the endemic biota of Arkansas by
adding 19 species to the state list, including two fungi, three gastropods, one araneid,
two opilionids, two pseudoscorpions, one diplopod, three collembolans, two
trichopterans, one coleopteran, one dipteran and one hymenopteran. In addition,
seven species (one pseudoscorpion, one collembolan, one bivalve, one
ephemeropteran, and three trichopterans) are removed from the state list and a
synthesis of new distributional records and changes in nomenclature are provided for
several species. This update brings to 126 the number of endemic species of the
state.

Robison et aL (2008) provided the most recent compilation on
the endemic biota of Arkansas. Their update brought to 113 (10
species of plants and 103 species/subspecies of animals) the total
number of Arkansas endemic flora and fauna. However, several
species were inadvertently overlooked. The following 19 species
are added to the list of Arkansas endemics: two fungi, three
gastropods, one araneid, two opilionids, two pseudoscorpions, one
diplopod, three collembolans, two trichopterans, one coleopteran,
one dipteran, and one hymenopteran. In addition, seven species
(one pseudoscorpion, one collembolan, one bivalve, one ephemer¬
opteran, and three trichopterans) are removed from the state list; a
synthesis of new distributional records are added for two
gastropods, one coleopteran, and one amphibian, and changes in
nomenclature are provided for three gastropods and two
coleopterans. This update brings to 126 the number of endemic
species of the state.

204 THE TEXAS JOURNAL OF SCIENCE-VOL. 6 1 , NO. 3, 2009

Table 1. Biota added to the state list of endemic species of Arkansas and counties of
occurrence.

Taxon

County/Counties

Reference

Fungi

Dictyostelium caveatum

Stone

Landolt et al. (2006)

Cryptovalsaria americana

Polk

Vasilyeva & Stephenson (2007)

Animalia

Daedalochila bisontes

Madison, Newton, Searcy

Coles & Walsh (2006)

Xolotrema occidentale

Independence, Stone

Walsh & Coles (2002)

Marstonia ozarkemis

Baxter

Hershler (1994)

Neoleptoneta arkansa

Stone

Gertsch (1974)

Crosbyella distincta

Boone

Goodnight & Goodnight (1942)

Crosbyella roeweri

Benton

Goodnight & Goodnight (1942)

Apochthonius diabolus

Washington

Muchmore (1967)

Apochthonius titanicus

Stone

Muchmore (1976)

Aliulus carrollus

Benton, Carroll, Searcy,

Washington

Hoffman (1999)

Typhlogastnira fousheensis

Independence

Christiansen & Wang (2006)

Pygmarrhopalites youngsteadti

Newton

Zeppelini et al. (2009)

Pygmarrhopalites buffaloensis

Newton

Zeppelini et al. (2009)

Cheumatopsyche robisoni

Garland, Montgomery, Polk Moulton & Stewart (1996)

Lepidostoma lescheni

Logan, Montgomery

Moulton et al. (1999)

Heterosternuta ouachitus

Howard, Izard, Newton,
Pike, Randolph, Searcy,

Sharp*

Longing & Hazzard (2009)

Atomosia arkansensis

Hempstead

Barnes (2008)

Idris leedsi

Johnson

Masner & Denis (1996)

* There are natural heritage records that also exist for Johnson and Pope counties
(NatureServe 2009).

List of Species

Material included following is a listing of the species
added (Table 1) and removed from the state endemic list, including
a synthesis of new distributional records and changes in
nomenclature for other endemic biota.

Additions to the State Endemic Fauna

Fungi, Mycetozoa, Dictyosteliaceae
Dictyostelium caveatum Waddell 1982

This cellular slime mold was described by Waddell (1982) from
a single isolate found on bat guano in total darkness in Blanchard
Springs Caverns, Stone County, Arkansas. It is considered to be a

MCALLISTER, ROBISON & SLAY

205

true Arkansas endemic found in a single Ozark cave to date
(Landolt et al. 2006).

Sordariomycetes

Cryptovalsaria americana Vasilyeva &l Stephenson 2007

This fungus was described by Vasilyeva & Stephenson (2007)
from specimens collected from the Ouachita Mountains Biological
Station, 6.5 km west of Big Fork, Polk County, Arkansas.
Collections of C americana were taken from the living bark of
hazel alder (Alnus serrulata).

Animalia, Mollusca, Gastropoda, Polygyridae

Daedalochila (syn. Millerelix) bisontes Coles & Walsh 2006

The Buffalo River liptooth, Daedalochila bisontes was
previously thought to be D. (Millerelix) peregrina from specimens
deposited in the Causey collection at the University of Arkansas-
Fayetteville and the Hubricht collection at the Field Museum of
Natural History-Chicago (Coles & Walsh 2006). It is considered
imperiled (G2) in rounded global status (NatureServe 2009). The
species inhabits limestone outcrops in the Ozarks, including
Madison, Newton, and Searcy counties (Walsh & Coles 2002;
Coles & Walsh 2006).

Xolotrema occidentale (Pilsbry & Ferriss 1907)

The Arkansas wedge, Xolotrema occidentale (syn. Triodopsis
occidentalis) is known only from Independence and Stone counties,
Arkansas (Pilsbry & Ferriss 1907; Walsh & Coles 2002). This snail
is considered critically imperiled (Gl) in rounded global status by
NatureServe (2009) and a species of special concern in the state by
the Arkansas Game and Fish Commission (Anonymous 2004).

Hydrobiidae

Marstonia ozarkensis (Hinkley 1915)

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

The Ozark pyrg, Marstonia ozarkensis (formerly Pyrgulopsis
ozarkensis) is known only from the type locality, the North Fork of
the White River above Norfolk, Baxter County, Arkansas (Hershler
1994). Thompson & Hershler (2002) re-evaluated eastern North
American species assigned to Pyrgulopsis and recognized them as
distinct species of the genus Marstonia. Interestingly, Wu et al.
(1997) lists a single site on the North Fork of the White River in
Ozark County, Missouri for M ozarkensis. However, efforts to
relocate this species beyond the single reported location in Missouri
have been unsuccessful (Natureserve 2009). This snail has also
likely been extirpated in Arkansas (Wu et al. 1997). It is
considered G1 in rounded global status (NatureServe 2009) and a
species of special concern in Arkansas (Anonymous 2004).

Arthropoda, Araneae, Leptonetidae

Neoleptoneta arkansa (Gertsch 1974)

This troglophilic spider was described by Gertsch (1974) from
Blanchard Springs Caverns, Stone County, Arkansas. Dorris
(1985) included the species as Leptoneta arkansa in her checklist of
Arkansas spiders.

Opiliones, Phalangodidae

Crosby ella distincta Goodnight & Goodnight 1942

This harvestman (an eyeless obligate cavemicole) was described
by Goodnight & Goodnight (1942) from specimens collected in
Wagler’s Cave near Harrison, Boone County, Arkansas. It is
considered critically imperiled (SI) in the state (NatureServe 2009).

Crosby ella roeweri Goodnight & Goodnight 1942

This harvestman species was described by Goodnight &
Goodnight (1942). Specimens were collected in Tom Danforth
Cave, Benton County, Arkansas. This is an eyed troglophile that is
considered SI in Arkansas (NatureServe 2009).

MCALLISTER, ROBISON & SLAY

207

Pseudoscorpiones, Chthoniidae

Apochthonius diabolus Muchmore 1967

Muchmore (1967) described A. diabolus from a single male
specimen from DeviFs Den Cave at DeviFs Den State Park,
Washington County, Arkansas. This obligate cavemicole is
considered SI in the state (NatureServe 2009).

Apochthonius titanicus Muchmore 1976

This pseudoscorpion was described by Muchmore (1976) from
individuals collected from Blanchard Springs Caverns, 5.6 km east
of Fifty Six, Stone County, Arkansas. Specimens were found under
a piece of paper near “The Titans”. This obligate cavemicole is
considered SI in Arkansas (NatureServe 2009).

Diplopoda, Julida, Parajulidae

Aliulus carrollus Causey 1950

This milliped was included as a state endemic by Robison &
Allen (1995) but inadvertently overlooked and not included by
Robison et al. (2008). The species was reported by Robison &
Allen (1995) from Carroll and Washington counties. However,
additional specimens have been reported from Benton and Searcy
counties (Hoffman 1999). The species may eventually be found in
adjacent states as the type locality (Blue Spring, Carroll County) is
just south of the Missouri line (Causey 1950) and sites in Benton
and Washington counties are close to eastern Oklahoma,

Hexapoda, Collembola, Hypogastmridae

Typhlogastrura fousheensis Christiansen & Wang 2006

This springtail species was described by Christiansen & Wang
(2006) from Foushee Cave, Independence County, Arkansas. A
single adult was collected by Norman and Jean Youngsteadt in May
1978. Additional adult specimens were collected 27 years later by
the same collectors from bat guano in the same cave on 18 March
2005 (Christiansen & Wang 2006).

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Arrhopalitidae

Pygmarrhopalites youngsteadtii Zeppelini, Taylor & Slay 2009

Specimens of this springtail species were collected from Tom
Barnes Cave, Newton County, Arkansas (Zeppelini et al. 2009).
This cave is located in the Ozarks within the Buffalo National
River.

Pygmarrhopalites buffaloensis Zeppelini, Taylor & Slay 2009

The holotype was collected from Walnut Cave, Newton County,
Arkansas (Zeppelini et al. 2009). This cave is located near the
Buffalo River, about 26 km upstream from the cave where P.
youngsteadtii was collected.

Trichoptera, Hydropsychidae

Cheumatopsyche robisoni Moulton & Stewart 1 996

This caddisfly was described by Moulton & Stewart (1996) from
specimens collected from Strawn Spring, 0.8 km east of Caddo
Gap, Montgomery County, Arkansas. Additional specimens of C
robisoni were collected from other sites in Garland, Montgomery
and Polk counties, Arkansas (Moulton & Stewart 1996). It appears
this species is endemic to small, spring-fed streams in the Ouachita
Mountain physiographic subregion. With additional collecting, C.
robisoni may be found just across the border in LeFlore County,
Oklahoma (along Rich Mountain) as specimens are available from
sites just to the east. This species is considered critically imperiled
(Gl) in rounded global status (NatureServe 2009).

Lepidostomatidae

Lepidostoma lescheni Bowles, Mathis & Weaver 1994

A single male L. lescheni was collected from Slocum Spring on
Mt. Magazine, Logan County, Arkansas, and described by Bowles
et al. (1994). Additional specimens (both males and females) were
collected from several sites in seep locations in the central Ouachita
Mountain region of the state in Montgomery County (Moulton et al.

MCALLISTER, ROBISON & SLAY

209

1999). A report (Weaver 2002) of the species from Missouri and
Oklahoma is erroneous (S. R. Moulton II pers. comm.). However,
additional collecting may reveal populations in similar seep areas of
eastern Oklahoma. This species is considered G1 in rounded global
status (NatureServe 2009).

Coleoptera, Dytiscidae

Heterosternuta ouachitus (Matta &Wolfe 1979)

The species was originally described as Hydroporus ouachitus
by Matta & Wolfe (1979). The subgenus Heterosternuta was
elevated to generic status by Matta & Wolfe (1981). The species
was originally reported from sites in the Ouachita Mountains (Matta
& Wolfe 1981). However, Pippenger & Harp (1985) reported the
range of H. ouachitus reaches into the Ozark Mountains (Janes
Creek, Randolph County). More recently, Harp & Robison (2006)
reported H. ouachitus from the Strawberry River system in Izard
and Sharp counties. Additional specimens were reported from
Long Creek (Searcy County), Beech Creek (Newton County), and
West Lafferty Creek (Izard County) by Longing & Haggard (2009).
Interestingly, Wolfe (2000) mentioned in couplets of keys to
Heterosternuta beetles, H, ouachitus probably occurs outside of
Arkansas; however, specimens have not yet been collected from
adjacent states or elsewhere (S. D. Longing, pers. comm.). The
species is considered imperiled (S2) in the state (NatureServe
2009).

Diptera, Asilidae

Atomosia arkansensis Barnes 2008

This robber fly was described by Barnes (2008) from specimens
collected in blackland prairie at Grandview Prairie Wildlife
Management Area near Columbus, Hempstead County, Arkansas.
The species is ranked SI in the state (NatureServe 2009) due to its
limited range.

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Hymenoptera, Scelionidae

Idris leedsi Masner & Denis 1996

A parasitoid wasp, Idris leedsi was described from a single
female collected using yellow pan traps from Baker Spring, 35.4
km NW of Clarksville, Johnson County, Arkansas (Masner & Denis
1996). Scelionids are solitary primary parasitoids of eggs of
various spiders (Johnson 1992).

Species Removed from the State Endemic Fauna

Bivalvia, Unionidae

Villosa arkansasensis (Lea 1862)

The Ouachita creekshell was reported to be an Arkansas
endemic by Robison & Allen (1995) from Clark, Garland, Howard,
Montgomery, Pike, Polk and Saline counties. In addition, it was
subsequently listed as an endemic by Robison et al. (2008).
However, Galbraith et al. (2008) report specimens of V.
arkansasensis from the Little River system of McCurtain County,
Oklahoma. It is a species of special concern in Arkansas
(Anonymous 2004).

Pseudoscorpiones, Neobisiidae

Tartarocreagris ozarkensis (Hoff 1945)

This pseudoscorpion was described as Microcreagis ozarkensis
by Hoff (1945) from specimens collected from Devil’s Den State
Park and Farmington, Washington County, Arkansas (Hoff 1945).
The species (=M ozarkensis) was included as an Arkansas endemic
by Allen (1988), Robison & Allen (1995) and Robison et al. (2008).
It is now known from additional localities in Arkansas (Clark and
Pulaski counties) and Latimer County, Oklahoma (Muchmore
2001), and is ranked SI in the state (NatureServe 2009).

MCALLISTER, ROBISON & SLAY

211

Collembola, Entomobryidae

Pseudosinella dubia Christiansen 1960

Christiansen (1960) described this troglobitic springtail from
specimens collected from Devil’s Den Kitchen Cave, Devil’s Den
Cave, and Granny Dean Cave, Washington County, Arkansas. It
was again reported from Devil’s Den Cave by Peck & Peck (1982).
Subsequently, the species was reported from a cave in Dent County,
Missouri, and a cave in Adair County, Oklahoma (Slay et al. 2009).

Ephemeroptera, Ephemerellidae

Dannella provonshai (McCafferty 1977)

This mayfly was originally described by McCafferty (1977)
from specimens collected on the Mulberry River, Johnson County,
Arkansas. Robison & Allen (1995) reported it was known only
from the type locality and Robison et al. (2008) included D.
provonshai in their list of endemics. However, the species has now
been reported from Alabama, Kentucky, New York, and Tennessee
(McCafferty & Webb 2006; NatureServe 2009; Ogden et al. 2009).
In Arkansas, D. provonshai is ranked SI (NatureServe 2009).

Trichoptera, Helicopsychidae

Helicopsyche limnella Ross 1938

Ross (1938) originally described this caddisfly from an unknown
Arkansas county. Unzicker et al. (1970) listed seven sites for H,
limnella in Benton, Crawford, Madison, and Washington counties.
Robison & Allen (1995) included Benton, Clark, Crawford,
Franklin, Garland, Hot Spring, Johnson, Madison, Montgomery,
Polk, Saline, and Washington counties in the range of H. limnella.
The species was also included in the Arkansas endemic biota list of
Robison et al. (2008). However, H. limnella has now been reported
from Missouri and Oklahoma (Moulton & Stewart 1996).

Hydroptilidae

Ochrotrichia robisoni Frazer & Harris 1991

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

This microcaddisfly species was described by Frazer & Harris
(1991) from specimens collected from Bear Creek at St, Hwy 7, 3.2
km south of Hollis, Perry County, Arkansas. The species is SI in
Arkansas (NatureServe 2009) and has been reported recently from
Oklahoma (Moulton & Stewart 1996).

Psychomyiidae

Paduniella nearctica Flint 1967

This caddisfly was originally described from specimens
collected from Devil’s Den State Park, Washington County,
Arkansas (Flint 1967); additional records include Johnson County,
Arkansas (Moulton & Stewart 1996). As such, it was included as a
state endemic species by Robison & Allen (1995) and Robison et
al. (2008). However, P. nearctica has now been reported from
southern Missouri (Moulton & Stewart 1996).

New Distributional Records and/or Changes in Nomenclature
Gastropoda, Polygyridae

Daedalochila (syn. Millerelix) peregrina (Rehder 1932)

The White Liptooth was reported as Polygyra peregrina in
Robison & Allen (1995) and Robison et al. (2008). However,
Coles & Walsh (2006) found that the diagnostic characters used to
define the genus Millerelix sensu Emberton (1995) were unreliable
and placed member species into the senior genus Daedalochila
Beck. The species is known from Izard, Marion, Newton, Searcy
and Stone counties (Robison & Smith 1982). Walsh & Coles
(2002) reported D. peregrina from Carroll County. This snail is G2
in rounded global status (NatureServe 2009) and a species of
special concern in Arkansas (Anonymous 2004).

MCALLISTER, ROBISON & SLAY

213

Patera clenchi (Rehder 1932)

The Calico Rock oval, P. clenchi was reported by Hubricht
(1972) only from a rock slide on Mt, Nebo, Yell County, Arkansas.
Robison & Smith (1982), Robison & Allen (1995) and Robison et
al. (2008) reported P. clenchi as Mesodon clenchi from Izard and
Yell counties. Walsh & Coles (2002) reported two new
distributional records for P. clenchi in Searcy and Scott counties.
It is considered G1 in rounded global status (NatureServe 2009) and
a species of special concern in the state (Anonymous 2004).

Inflectarius magazinensis (Pilsbry & Ferriss 1907)

This Magazine Mountain shagreen is only known to occur on the
north slope of Mt. Magazine in the Ozark National Forest of Logan
County, Arkansas (Pilsbry & Ferriss 1907). It was listed as an
Arkansas endemic by Robison & Smith (1982), Robison & Allen
(1995) and Robison et al. (2008) as Mesodon magazinensis.
Caldwell (1986) was unable to verify L magazinensis from the
south slope of Mt. Magazine; however, additional specimens were
reported from the north slope by Walsh & Coles (2002). Its limited
range makes it particularly sensitive to any habitat alteration and it
is therefore listed as SI in Arkansas (NatureServe 2009), as an
endangered species in the state (Anonymous 2004), and as a
threatened species by the U.S. Fish and Wildlife Service on 17
April 1989 (Anonymous 1989).

Coleoptera, Dytiscidae

Heterosternuta sulphuria (Malta & Wolfe 1979)

This predaceous diving beetle was originally described as
Hydroporus sulfurus by Malta & Wolfe (1979) and included as an
Arkansas endemic by Robison & Allen (1995). The subgenus
Heterosternuta was elevated to the generic level by Malta & Wolfe
(1981). Specimens of this endemic species were originally
collected from Sulphur Springs, Benton County, Arkansas (Malta &
Wolfe 1979). Additional historical records include sites in Izard,

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Newton, and Searcy counties. More recently, however. Longing &
Haggard (2009) reported new distributional records for H.
sulphuria in Benton, Newton, and Washington counties, including
the first report of the species from the entrance of a cave. With
additional collecting, this d3discid may eventually be found outside
of Arkansas in adjacent states (S. D. Longing, pers. comm.).

Chordata, Amphibia, Caudata, Plethodontidae

Plethodon caddoensis Pope & Pope 1951

The Caddo Mountain salamander, Plethodon caddoensis was
reported to be an Arkansas endemic in Howard, Montgomery, and
Polk counties (Robison & Allen 1995). Trauth & Wilhide (1999)
reported new geographic records for P. caddoensis from two sites
in Pike County. This salamander is considered a species of special
concern in the state (Anonymous 2004).

In summary, the present study brings to 126 species the number
of endemic biota of Arkansas. Nineteen species have been added to
the state list since the last update in 2008. In addition, seven
species (one pseudoscorption, one springtail, one bivalve, one
ephemeropteran, and three caddisflies) are removed from the state
list and a synthesis of new distributional records is added for two
endemic gastropods, one endemic coleopteran, and one endemic
amphibian. Changes in nomenclature are provided for three
endemic gastropods and two endemic coleopterans.

Acknowledgments

Appreciation is extended to D. Bowles (National Park Service),
G. L. Harp (Arkansas State University), G. Leeds (U. S. Forest
Service), S. D. Longing (UA-Fayetteville), J. C. Morse (Clemson
University), S. R. Moulton, II (U.S. Geological Survey), F. Spiegel
(UA-Fayetteville), S. Stephenson (UA-Fayetteville), and J. S.
Weaver, III (New Hampshire) for providing information on
Arkansas endemics. We also thank S. R. Moulton, II for critically
reviewing the manuscript. Funding for MBS was provided by the

MCALLISTER, ROBISON & SLAY

215

Arkansas Game and Fish Commission, The Nature Conservancy
(Arkansas Field Office), and U. S. Fish and Wildlife Service
(Arkansas Ecological Services).

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CTM at: drctmcallister@aol.com

TEXAS T OF SCI. 6 1(3):2 19-224

AUGUST, 2009

SELECTION OF AVAILABLE POST-FIRE SUBSTRATE
BY THE GROUND SKINK, SCINCELLA LATERALIS
(SQUAMATA: SCINCIDAE)

Charles M. Watson

Department of Biology, The University of Texas at Arlington
Arlington, Texas 76019

Abstract -Burning of the forest floor alters the structural components that
constitute the organic substrate. Many small animal species inhabit this layer, which
typically consists of leaf litter from surrounding trees. The availability of a species’
preferred substrate can be a factor in the rate of recolonization following a fire. Using
pair-wise choice trials within a controlled environment, preference of substrate
typically available after a bum by Scincella lateralis was determined. These skinks
primarily select hardwood leaf litter and secondarily choose pine needle litter and
pine bark slough. Bare ground was usually avoided. These findings indicate that S.
lateralis may not be able to completely recolonize a site until after the first seasonal
leaf fall following a fire.

The effects of fire on populations of various reptiles have been
well documented (Wilgers & Home 2006; Brown 2001; Granberry
et al. 1994). Braithewaite (1987) found that varied fire regimes
affect lizard populations differently, depending on such factors as
fire intensity, duration, and seasonality. Kahn (1960) determined
that the presence of unaltered refuge sites can buffer the effects of
fire on certain lizard populations. For many temperate leaf-litter
dwelling species, fire leaves behind limited and sparse refuge
(Watson 2004), The present study determines the selectivity of the
ground skink, Scincella lateralis, to four substrate components that
are available at varying degrees after a fire in a mixed
hardwood/pine forest. Scincella lateralis makes a sound
experimental subject due to the species’ abundance, small size, and
limited vagility. This animal is found in temperate wooded areas
where there is sufficient water, cover, and food (Brooks 1967).
However, the leaf-litter layer within the forest, or at forest’s edge, is
considered typical habitat for this species (Fitch & von Achen
1977).

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The ground skink depends on leaf litter and other structural
organic components of the forest floor for refuge (Brooks 1967).
The complete destruction of decaying leaf material may
dramatically affect recovery of this species within burned sites.
Conversely, if the ground skink will utilize pine needle litter or pine
bark slough, both of which are relatively abundant following a fire,
it may be able to persist and recolonize burned sites more rapidly.
This experiment aims to determine the selection of individual
ground skinks to four prominent substrate types found after a bum
in a mixed hardwood/pine forest ecosystem in eastern Texas.

Methods & Materials

Six experimental chambers were used, each measuring 32 cm by
77 cm at the base, providing 2464 cm^ of area. This area was
uniformly covered by approximately 3 cm of sifted topsoil.
Individual chambers contained two of the four treatments for
habitat preference, each making up half of the surface area.
Substrate treatments, with the exception of bare ground, were
loosely arranged aggregates of the treatment substrate approxi¬
mately 5 cm thick. These six chambers cover all of the possible
comparisons for these treatments. The four treatments are as
follows:

Hardwood leaf litter broad leaves, predominantly from

Quercus sp. This substrate type is generally destroyed in a bum
and will not be present until the following fall. The leaves that
make up this layer are broad and can serve as cover for S. lateralis.

Pine leaf /zY/er. -Needles from Finns taeda. This substrate type
will be present shortly after a bum and will continue to accumulate
throughout the year. This is due to the fact that the needles of pine
trees are persistent, and their leaf fall is not seasonnaly limited
(Vines 1990). The needles that make up this layer are thin and can
only serve as refuge in aggregate.

WATSON

221

Pine slough -T\{m pieces of scorched pine bark that accumulate
at the base of the tree in the months following a bum. The bark
pieces that make up this layer stmcturally resemble hardwood
leaves, providing broad areas of cover and stmcture. However,
these units are more dense than comparably sized leaves.

Bare ground large organic substrate present. This repre¬
sents the majority of a site’s substrate immediately following a
bum.

A total of 10 animals of unknown sex (five adults, snout-to-vent
length of 42-5 1mm; five juveniles, snout-to-vent length of 29-
34mm) were obtained from natural populations in Smith and Dallas
counties, Texas. The organic substrates were collected from sites
within Tyler State Park and heated to temperatures in excess of
65°C. The elevated temperatures ensured the elimination of prey
items from the material. This was verified by observation of
subsamples of this substrate under a stereomicroscope. No live
invertebrates were noted.

Trials were performed under fluorescent lighting at 2TC.
Individuals were each transferred to the center of the chamber, with
a treatment to either side of six experimental chambers. The animal
was then allowed a 10-min period of acclimation to the chamber.
After this period, the investigator approached the chamber and
marked which substrate type that the animal was in at time of initial
detection. The skink typically was visible when the chamber was
approached, and immediately retreated to cover. Each trial was
repeated 20 times for each animal.

Data were analyzed as recommended by Cherry (1998).
Confidence intervals were determined for the mean number of
times that a specimen was observed in each substrate, given equal
opportunity to choose between substrates. There were no
significant differences noted between age groups for any of the
treatments. Therefore, the values of both age classes were

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Figure 1. Mean (± 95% Cl) substrate choice values for Scincella lateralis given a
maximum of 60 potential observations for each type. HLL = Hardwood Leaf Litter,
PNL = Pine Needle Litter, PS = Pine Bark Slough, BG = Bare Ground.

combined into one sample to increase power. Significance was
determined between substrates if no values were shared within the
constructed 95% confidence intervals (Figure 1).

Results & Discussion

Bare ground exhibited the lowest mean frequency (x = 1.0, 95%
Cl : [0.3, 1.7]), and a significant preference was exhibited for
hardwood leaf litter over all three other treatments (x = 47.6 95%
Cl : [43.2, 52.0]). Pine slough (x = 34.3, 95% Cl : [30.6, 38.0])
and pine needle litter (x = 37.5, 95% Cl : [33.8, 41.2]) exhibited
overlap of the constructed confidence intervals, thereby exhibiting
no significant selection between these treatments.

The top layer of substrate on the forest floor is typically
comprised of dead and decaying organic material. Fire
dramatically alters the makeup of this substrate layer, thereby
altering the primary habitat of S. lateralis. Watson (2004) found,
from data gathered in Smith County, Texas, that the presence of
structural organic material is reduced by over 75% following a fire.
Preference for those substrates that are most decimated by fire may
prove to be a limiting factor in recolonization by this species.

WATSON

223

The preference of hardwood leaf litter for cover is consistent
with literature regarding other skink species (Fitch 1954).
However, the three other substrates are the most commonly
available in the period immediately following a bum and preceding
the annual leaf fall (Watson 2004). Potential costs to this animal as
related to these substrates may be a factor into substrate selection.
The probability of detection of the ground skink by avian predators
is increased in the absence of refiigia (Smith 1997). Pine needle
litter does not provide effective cover unless it is present in
aggregate and the density of the pine slough may not allow for free
movement of the lizard when it is arranged in a compact manner at
the base of a tree. Further reasons for the substrate preferences may
be dependant on the availability of prey. There is little doubt that
the acquisition of prey and the ability to hide from predators play
heavily in microhabitat selection.

The preferred microhabitat type for the ground skink is that
which is in shortest supply following a fire. Other factors, such as
prey availability and proximity to undisturbed habitat, may also
affect the recovery of this species following a fire. Invertebrate
communities, which constitute the prey of S. lateralis, are often
initially eradicated with the burning of leaf litter, further tying the
recovery of this species to the return of the leaf litter layer and
associated fauna (Abbot et al. 2002; Bird 1997). Therefore, ground
skinks may not be able to recolonize an area completely until
hardwood leaf litter is present in pre-bum amounts, which may take
over three years (Watson 2004). Furthermore, the community
stmcture of the leaf litter layer may not reach the pre-bum state for
many years afterwards, potentially limiting the fiill recovery of S.
lateralis populations. The recovery of this species following a bum
is nevertheless basically tied to substrate availability and its
timeline for recovery may be delayed for months following the
event, beginning with the first leaf fall and the associated
accumulation of hardwood leaf litter.

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THE TEXAS JOEfRNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Acknowledgments

I would like to thank Daniel Formanowicz and Laura Gough for
their input and encouragement over the duration of this project as
well as Jessie Meik, Brian Fontenot, and Rebbekah Watson for their
pre-submission editorial expertise.

Literature Cited

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Bird, S. 1997. The effects of silvicultural practices on soil and leaf litter arthropods in an
East Texas pine plantation. Unpublished Ph. D. dissertation. Texas A&M Univ.,
College Station, 193pp.

Braithwaite, R. W. 1987. Effects of fire regimes on lizards in the wet - dry tropics of
Australia. J. Trop. Ecol., 3(3):265-275.

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Fitch, H. S. 1954. Life History and Ecology of the Five-lined Skink, Eumeces fasciatus.
Univ. Kansas Press, Lawrence, KS, 156pp

Fitch, H. S. & P. L. von Achen. 1977. Spatial relationships and seasonality in the skinks
Eumeces fasciatus and Sciucella laterale in Northeastern Kansas. Herpetologica,
33(3):303-313.

Cranberry, C. H., D. G. Neary & L. D. Harris. 1994. Effects of high-intensity wildfire
and silvicultural treatments on reptile communities in Sand-Pine Scrub. Conservat.
Biol., 8(4):1047-1057.

Kahn, W. C. 1960. Observations on the effect of a bum on a population of Sceloporus
occidentalis. Ecology, 41(2):358-359

Smith, D. G. 1997. Ecological factors influencing the antipredator behaviors of the
ground skink, Scincella lateralis, Behav. Ecol., 8(6):622-629.

Vines, R. A. 1990. Trees, Shmbs, and Woody Vines of the Southeast. Univ. of Texas
Press, Austin, TX, 1 1 04pp

Watson, C. M. 2004. The Effects of Controlled Burning on Ground Skink Populations
in a Mixed Pine-Hardwood Habitat of East Texas. Unpublished M.S. Thesis., Univ.
of Texas at Arlington. 47pp.

Wilgers, D. J. & E. A. Home. 2006. Effects of different bum regimes on Tallgrass
Prairie herpetofaunal species diversity and community composition in the Flint Hills,
Kansas. J. Herpetol., 40(l):73-84.

CMW at: cwatson@uta.edu

TEXAS J. SCI. 61(3), AUGUST, 2009

225

GENERAL NOTES

REPRODUCTION IN SMITH’S GREEN-EYED GECKO,
GEKKO SMITHII (SQUAMAT A: GEKKONIDAE)

Stephen R. Goldberg

Department of Biology, Whittier College, PO Box 634,
Whittier, California 90608

Smith’s green-eyed gecko, Gekko smithii is known from
Sarawak, Sabah, Kalimantan, and Brunei (Borneo) and southern
Thailand, West Malaysia, Sumatra, Nias, Java and the Nicobar
Archipelago of India (Das 2007). It is the largest Bornean gecko
(Das 2007). There is information on egg laying of G. smithii in
Manthey & Grossmann (1997), Rogner (1997) and Das (2007).
The purpose of this note is to provide additional information on the
reproductive cycle of G. smithii from a histological examination of
museum specimens.

A sample of 43 G smithii consisting of 20 males (mean SVL =
162.3 mm ± 14.3 SD, range = 133-178 mm) and 23 females (mean
SVL - 153.4 mm ± 1 1.9 SD, range = 134-175 mm) collected 1960-
1990, from Borneo, Indonesia and Malaysia was borrowed from the
herpetology collection of the Field Museum of Natural History
(FMNH), Chicago, Illinois, USA.

For histological examination, the left testis was removed from
males and the left ovary was removed from females. Enlarged
follicles (> 5 mm length) or oviductal eggs were counted {in situ).
Tissues were embedded in paraffin and cut into sections of 5 pm.
Slides were stained with Harris hematoxylin followed by eosin
counterstain (Presnell & Schreibman 1997). Slides of testes were
examined to determine the stage of the spermatogenic cycle. Slides
of ovaries were examined for the presence of yolk deposition or
corpora lutea. Histology slides were deposited in the Field Museum
of Natural History (FMNH) herpetology collection. An unpaired t-

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

test was used to compare G. smithii male and female mean SVLs
using Instat (vers. 3.0b, Graphpad Software, San Diego, CA).

The following G. smithii were examined: Borneo, Sabah State,
Tawau Division, Lahad Datu District, FMNH 246237; Sarawak
State, Fourth Division, Miri District, FMNH 129494, 131521; Third
Division, Kapit District, FMNH 145834, 145835, Fourth Division,
Bintulu District, FMNH 148996-148998, 150731, 150733-150735,
150737-150739; Malaysia, Selangor State, Petaling District: FMNH
185107, 185111-185114, 185116, 185117, 185119-185126,
185133, 185135-185137, 185139, 185140-185142, 185144,
185145; Indonesia, North Sumatra, FMNH 209498- 209500,
209502.

Males of G. smithii were significantly larger than females
(unpaired Mest, df = A\, t = 2.22, P = 0.032). The only stage
observed in the testicular cycle was spermiogenesis in which the
lumina of the seminiferous tubules were lined by sperm or clusters
of metamorphosing spermatids. Spermiogenesis was noted in the
following males (sample size in parentheses): January (2); February
(1); March (7); April (7); May (1); September (1); November (1).
The smallest reproductively active (spermiogenic) male measured
133 mm SVL (FMNH 185137).

Four stages were noted in the ovarian cycle of G smithii (Table
1): (1) quiescent (no yolk deposition); (2) early yolk deposition,
basophilic yolk granules are present; (3) enlarged yolk-filled
ovarian follicles, > 5 mm diameter; and (4) oviductal eggs with
eggs in oviducts. The smallest reproductively active female
measured 141 mm SVL (FMNH 150733) contained two oviductal
eggs and was collected in November. Mean clutch size for 11
females was 2.0. A clutch size of two is typical for many
gekkonids (Vitt 1986) and was reported for G. smithii by Das
(2007).

TEXAS J. SCI. 61(3), AUGUST, 2009

227

Table 1. Monthly stages in the ovarian cycle of G. smithii.

Month

Quiescent

Early yolk
deposition

Enlarged follicles
> 5 mm

Oviductal

eggs

February

March

April

June

July

October

November

December

The presence of reproductively active females and males in all
months sampled suggests G. smithii exhibits year-round
reproductive activity. These samples are too small to ascertain
peaks in reproduction, if any exist. Rogner (1997) reported that a
captive G. smithii laid two clutches one week apart. Multiple
annual clutches of two eggs were reported by Manthey &
Grossmann (1997). McKeown (1996) reported the congener Gekko
gecko may lay a pair of eggs each month in Hawaii. No
histological evidence was found that multiple clutches were
produced as indicated by, for example, corpora lutea from a
previous clutch and concomitant yolk deposition for a subsequent
clutch. However, these data shows an extended period of
reproductive activity in natural populations, and along with reports
from Rogner (1997) and Manthey & Grossmann (1997), it is
expected that G. smithii produce several clutches of eggs each year.

Inger & Greenberg (1966) reported similar results in their
examination of the reproductive cycles of the gekkonids
Cyrtodactylus malayanus and C. pubisulcus from a Bornean rain
forest. In both species, males produced sperm throughout the year,
gravid females were present in most months, and no distinct
breeding season was evident. Similar results were also reported for
the geckos Hemidactylus platyurus {= Cosymbotus platyurus),

228

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Hemidactylus frenatus and Gehyra mutilata (= Peropus mutilatus)
from West Java, Indonesia by Church (1962) and Dixonius
siamensis from Thailand (Goldberg 2008). The above studies
suggest extended periods of reproduction may be typical for
gekkonid lizards from tropical southeast Asia. However, in view of
the great diversity of gekkonid species from southeast Asia (Uetz &
Hallermann 2009) subsequent investigations of the reproductive
cycle of many additional species will be needed to test this
hypothesis.

Acknowledgments

I thank Alan Resetar (Field museum of Natural History),
Chicago, Illinois for permission to examine G. smithii.

Literature Cited

Church, G. 1962. The reproductive cycles of the Javanese house geckos, Cosymbotus
platyurns, Hemidactylus frenatus, md Peropus mutilatus. Copeia, 1962:262-269.
Das, 1. 2007. Amphibians and Reptiles of Brunei. Natural History Publications

(Borneo) Kota Kinabalu Sabah, Malaysia. 200 pp.

Goldberg, S. R. 2008. Reproduction in the Siamese leaf-toed gecko, Dixonius siamensis
(Squamata: Gekkonidae) from Thailand. Texas J. Sci., 60(3):233-238.

Inger, R. F. & B. Greenberg. 1966. Annual reproductive patterns of lizards from a
Bornean rain forest. Ecology, 47 : 1 007-1 02 1 .

Manthey, U. & W. Grossmann. 1997. Amphibien & Reptilien Siidostasiens. Natur und
Tier Verlag, Munster, Germany, 512 pp.

McKeown, S. 1996. A Field Guide to Reptiles and Amphibians in the Hawaiian Islands.

Diamond Head Publishing, Inc., Los Osos, California, 172 pp.

Presnell, J. K. & M, P. Schreibman. 1997. Humason’s Animal Tissue Techniques. 5*^
Edit. The Johns Hopkins Press, Baltimore, 572 pp.

Rogner, M. 1997. Lizards, Vol. 1, Krieger Publishing Company, Malabar, Florida, 317

pp.

Uetz, P,, & J. Hallermann. 2009. TIGR Reptile Database: http:/www.reptile-
database.org. (accessed 7 December 2009).

Vitt, L. J. 1986. Reproductive tactics of sympatric gekkonid lizards with a comment on
the evolutionary and ecological consequences of invariant clutch size. Copeia,
1986:773-786.

SRG at: sgoldberg@whittier.edu

TEXAS J. SCL 61(3), AUGUST, 2009

229

THE LONG-TAILED WEASEL MUSTELA FRENATA
(MAMMALIA: MUSTELIDAE) m BAJA CALIFORNIA, MEXICO

Gorgonio Ruiz-Campos, Roberto Martmez-GaDardo,

Salvador Gonzalez-GuzmaE and Jorge Alaniz-Garcia

Facultad de Ciencias, Universidad Autonoma de Baja California,

Apdo. Postal 233, Ensenada, Baja California, 22800, Mexico

The long-tailed weasel Mustela frenata Lichtenstein 1831 has
the largest range of any mustelid in the Western Hemisphere, with a
known distribution extending from southern Canada throughout the
United States, Mexico, and Central America, and into northern
South America, Mustela frenata occurs in a variety of habitats
from alpine-artic to tropical, but does not inhabit arid biotopes or
deserts. Its favored habitats include brush land and open timber,
brushy borders of croplands, grasslands along creeks and lakes, and
swamps (Svendsen 2003),

The long-tailed weasel is considered a generalist predator that
consumes a wide variety of small vertebrates. Rodents and
immature rabbits are preferred prey, however shrews, moles, bats,
birds, bird eggs, snakes, invertebrates, and carrion may be
consumed as alternative prey (Sheffield 1999).

The density of the long-tailed weasel throughout its range is
generally low, and its presence varies from uncommon to rare.
Owing to low densities and periodic fluctuations of its populations
in North America, this species is listed as endangered, threatened,
rare, or of special concern in many states and provinces (Sheffield
1999), This species is not listed in any special category of
conservation in Mexico,

Of the ten subspecies of M frenata known to occur in Mexico
(Hall 1981), M frenata latirostra inhabits a small portion of
northwestern Baja California (Huey 1964). The first published
record of this subspecies in Baja California is referable to Federal

230

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Highway No. 1 near Rosarito (10 mi S Tijuana) (Huey 1964). The
second record as reported by Ralston & Clark (1971) at this same
highway near La Mision (32° 3’ N, 116° 54’ W) on 7 Apr 1968.
Additionally, there is one previously unreported voucher specimen
in the Museum of Vertebrate Zoology, University of Califomia-
Berkeley (MVZ- 148254), which was collected by W. E. Glanz 2 mi
WSW Maneadero (31° 42’ 0.0” N, 116° 35’ 24.0” W) on 13 Dec
1974.

Five recent records of M frenata latirostra are documented in
this paper for several localities within northwestern Baja California
(Fig. 1). Voucher specimens obtained during the study were
deposited in the Mammal Collection of the Universidad Autonoma
de Baja California (UABC) at Ensenada. The details of these
records are as follows: A male (UABC-093) was found dead in a
cultivated field at Valle de Maneadero, 14 km S Ensenada (3U 44'
37" N, 116° 35' 54" W), on 12 Aug 2000. Standard measurements
of this specimen were 405 mm, total length [TL]; 107 mm, tail
length [CL]; 44 mm, hind foot length [HFL]; weight [W], 288 g).
A female (U ABC-092) was collected in the Valle de Guadalupe, 36
km NE Ensenada, on 24 May 2005 (32° 03’ 30” N, 116° 35’ 09”
W). Standard measurements of UABC-092 were 360 mm TL, 150
mm CL, 36 mm HFL, 149 g W (Fig. 2). A second male (UABC-
107) was captured in the L.A. Cetto vineyard at Valle de Guadalupe
on 15 Jul 2003 (32° 07’30” N, 116° 28’ 40” W). The standard
measurements of the second male were 345 mm TL, 140 mm CL,
36 mm HFL, 100 g W. The first two specimens (UABC-092 and
093) were preserved as skin and skull vouchers, whereas the third
specimen was preserved in ethanol. Selected cranial measurements
in millimeters of UABC-092 and UABC-093 were respectively as
follows: zygomatic breadth 21.9 and 28.3, braincase height 14 and
17, upper jaw length 17 and 18, lower jaw length 24.7 and 30.4, and
least interorbital breadth 6.4 and 9.4.

TEXAS I SCI. 61(3), AUGUST, 2009

231

Fig. 1. Historical and recent records of Mustela frenata latirostra in Baja California,
Mexico. VR = visual record, and SR = specimen record.

Figure 2. Female Mustela frenata latirostra (UABC-092) collected at Valle de
Guadalupe, 36 km NE Ensenada, Baja California, on 24 May 2005. Photograph by
Gorgonio Ruiz-Campos.

A fourth record corresponds to a head in taxidermy belonging to
Ramon Femat. Mr. Femat obtained the weasel in 1997 while
hunting near the mouth of the Rio Santo Domingo at W Vicente

232

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Guerrero (30° 43' N, 116° 02' W). Finally, a fifth record is based
upon a sight record from San Telmo de Arriba, in the vicinity of the
Sierra San Pedro Martir, on 25 May 2002 (30° 58’ 09” N, 116° 05’
33” W) (Ruben Bustamante, pers. comm.). Based upon these
specimens, the current distribution of M frenata latirostra in
northwestern Baja California may be extended southward to the
coastal valley of the Rio Santo Domingo. Future intensive
sampling may confirm the presence of this subspecies as far as 30°
N latitude at the boundary the Californian faunal province
(Bancroft, 1926).

Acknowledgements

We thank Areli Castillo, Francisco J. Valverde and Hector Yee
for helping us with field sampling and preparation of specimens.
Also, we grateftilly acknowledge Ruben Bustamante and Ramon
Femat for providing the records for San Telmo de Arriba and
Lower Rio Santo Domingo, respectively. Likewise we thank two
anonymous reviewers for their useful comments that improved the
content and clarity of the manuscript.

Literature Cited

Bancroft, G. 1926. The faunal areas of Baja California del Norte. Condor, 28(5):
209-215.

Hall, R. E. 1981. The mammals of North America. John Wiley & sons. New York,

1181 pp.

Huey, L. M. 1964. Mammals of Baja California, Mexico. Transactions of the San
Diego Society of Natural History, 13(7):85-168.

Ralston, G. L. & W. H. Clark. 1971. Occurrence of Mustela frenata in northern Baja
California, Mexico. The Southwestern Naturalist, 16(2):209.

Sheffield, S. R. 1999. Long-Tail weasel {Mustela frenata). Pp. 169-171 in (D.E.
Wilson & S. Ruff, eds.). The Smithsonian Book of North American Mammals.
Smithsonian Institution Press. Washington, 750 pp.

Svendsen, G. E. 2003. Weasel and Black-footed Ferret {Mustela species). Pp. 650-
661 in (G.A. Feldhamer, B.C. Thompson & J.A. Chapman, eds.), Wild Mammals
of North America. The Johns Hopkins University Press, Baltimore, 1216 pp.

G. R-C at: gruiz@uabc.mx

TEXAS J. SCI. 61(3), AUGUST, 2009

233

STOMACH CONTENTS OF CALIDRIS MINUTILLA
(CHARADRIIFORMES: SCOLOPACIDAE) WINTERING AT A
FRESHWATER RESERVOIR IN WEST-CENTRAL TEXAS

Andrew C. Kasner, Randall H. Ruddick, and Terry C. Maxwell

Department of Biology, Wayland Baptist University,

1900 West 7th Street, Plainview, Texas 79072,

Department of Biology, Lamar University, P.O. Box 10037
Beaumont, Texas 77710 and
Department of Biology, Angelo State University
San Angelo, Texas 76909

Sandpipers (Charadriiformes: Scolopacidae) feed primarily on
invertebrates (Skagen & Oman 1996). Shorebird diets consist
primarily of polychaete and oligochaete worms (Schneider 1987;
Davis 1996; Tsipoura & Burger 1999), amphipods (Wilson &
Parker 1996; Shepherd & Boates 1999), horseshoe crab (Limulus
polyphemus) eggs (Tsipoura & Burger 1999), and insects,
especially dipterans (Schneider 1987; Alexander et al. 1996; Davis
1996). Few dietary studies have focused on sandpipers at inland
sites (Alexander et al. 1996; Davis 1996), and all of these focused
on migrating populations. Studies are needed to determine the
dietary preferences of sandpipers wintering at inland sites in order
to better understand why some individuals remain at more northern
latitudes over winter, when populations of invertebrate prey are
likely at their lowest levels of the year. It is of particular
importance to determine whether shorebirds at inland sites are able
to locate high quality food resources (easily captured and digested;
predictable in occurrence) or must subsist on low quality food items
(difficult to capture or digest; unpredictable occurrence).
Furthermore, habitat heterogeneity is typically lower at inland,
freshwater reservoirs (such as in this study) compared to coastal
areas, and it is likely that prey diversity and potentially abundance
is similarly poor relative to coastal regions.

The purpose of this research was to determine the diet of
wintering sandpipers at an inland freshwater reservoir. Least
Sandpipers (Calidris minutilla) and Western Sandpipers {Calidris

234

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

mauri) migrating along the Central Flyway (Smith et al. 1991; Neill
1992; Davis 1996; DeLeon 1996) winter in small numbers (<100
individuals per flock with only one or two flocks observed on any
given day) in west-central Texas (Tarter 1997; Kasner 1999). This
paper reports analysis of stomach contents from wintering
sandpipers collected at Twin Buttes Reservoir, Tom Green County,
in west-central Texas.

A total of 14 birds representing two species of Calidris, Least
Sandpiper (C. minutilla, j7=13) and Western Sandpiper (C mauri,
n=\), were collected. On 17 January 1998 and 28 February 1998,
six Least Sandpipers and one Western Sandpiper were collected
from an algal mudflat on the west shore of the south pool of Twin
Buttes Reservoir, near the mouth of the South Concho River, Tom
Green County, Texas. Birds were observed foraging on the
mudflat, wading in water approximately one to five centimeters
deep. On 19 January 2002, seven Least Sandpipers were collected
on the north shore of the south pool of Twin Buttes Reservoir.
Birds at this site were foraging in a flooded area of Bermudagrass
(Cynodon dactyl on) over a mud and gravel substrate. Avian
specimens were deposited in Angelo State Natural History
Collections, Department of Biology, Angelo State University, San
Angelo, Texas.

Bird stomachs were injected with ten percent formalin through
the mouth and esophagus with a hypodermic syringe immediately
after collection. Esophagus and stomach were removed from each
specimen in the lab later the same day. Stomach, esophagus, and
contents were stored in 10% formalin for analysis. Each stomach
and esophagus was dissected, contents were identified to the lowest
taxonomic rank possible (family or genus), and heads and other
parts were counted using a Bausch and Lomb 2X compound
dissecting microscope. Percent composition (proportion of total
stomach contents) of prey was visually determined for specimens
collected in 1998. For each specimen collected in 2002, stomach
and esophagus was dissected and flushed with formalin onto filter

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235

paper in a Buchner funnel. Additional formalin was used to
disperse aggregations of contents on the filter-fiinnel, recording the
volume of formalin used for flushing/dispersing. After being
allowed to air dry for 24 hrs, contents were identified using Merritt
& Cummings (1984) and grouped. Each subset of the contents was
massed (mg) using a Mettler balance and percent mass was
calculated for each type of item. To control for mass of formalin
contributing to stomach content mass measurements, a control trial
was conducted using filters and formalin. The two filters were
massed and then treated with 20 mL and 40 mL of formalin,
respectively (representing the minimum and maximum volumes of
formalin used for flushing/dispersing contents), allowed to air dry
for 24 hrs, and massed to determine the mass of residual formalin in
the filter (mg/mL). Mass contribution of formalin averaged 1.5
mg/mL of formalin used for flushing. Data were adjusted for the
contribution of formalin to mass measurements.

Stomach contents of all seven specimens in 1998 consisted
almost exclusively (>95%) of chironomid (Diptera: Chironomidae)
larvae, with all stomachs completely full. Mean number of
chironomid heads was 176.4 heads/stomach, with approximately
15% of chironomids in each stomach completely intact (Table 1).
The remaining contents consisted of unidentified insect parts, one
adult mesoveliid (Hemiptera: Mesoveliidae), and two seeds (Table
1). Additionally, all stomachs contained small amounts of minerals
(calcite or quartz), small pebbles, and strands of filamentous green
algae. Unidentified insect parts were suspected to be primarily
Coleopteran (many wing fragments and robust, dark colored leg
fragments and jaw fragments), however, this is speculative because
fragments were too incomplete for identification.

All but one of the seven stomachs from 2002 contained
identifiable fragments (heads, jaws, and/or legs) of Coleoptera
(Table 2), Families represented included Curculionidae
{Emphyastes, Stenopelmus, Tanysphyrus), Dytiscidae (Rhantus),
Hydrophilidae (Berosus), and Elmidae. Hemiptera was also present

236

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

Table 1. Stomach contents of Calidris minutilla {n=6) collected 17 January and 28
February 1998 at Twin Buttes Reservoir, Tom Green County, Texas.

specimen Number

No. Chironomid heads

Other prey items

1003-TCM3776

138

1 mesoveliid, misc.

1003-TCM3777

201

miscellaneous

1003-TCM3778

221

miscellaneous

1012-TCM3782

153

2 seeds, misc.

1012-TCM3783

174

miscellaneous

1012-TCM3784

205

miscellaneous

in one specimen, but only two heads were found, therefore mass
was not measurable and not included in Table 2. Minerals (calcite
and quartz) and pebbles contributed a large proportion of overall
mass in each specimen from 2002, indicative of a hard diet, such as
Coleoptera, that would require more mechanical digestion. In
specimens from 1998, the diet was primarily softer Chironimids,
and much less mineral content was observed. Alternatively,
inefficient foraging on beetles in the flooded Bermuda-grass habitat
could have led to accidental intake of minerals and pebbles. One
specimen in 2002 also contained small seeds (Table 2). The
remaining unidentified contents were insect fragments lacking
sufficient detail for identification and strands of filamentous green
algae.

These findings are consistent with other studies in the types of
prey found and proportion of prey types. Chironomids were
previously found to be an important component of the diet of
calidrine sandpipers (Schneider 1987; Alexander et al. 1996; Davis
1996; Skagen & Oman 1996), however, the highest reported
average percentage was 80.1% (Davis 1996). This study found the
percentage of chironomid larvae to be greater than 95% for all
specimens in 1998, but the small sample size and limited number of
collection dates precludes any inference that this is significantly
greater than other studies, and chironomids were noticeably absent
in 2002 specimens. The frequent occurrence of Chironomidae in
1998 specimens and Coleoptera in 2002 specimens suggests that
these groups may be important prey in winter, however, Coleop-

TEXAS J. SCL 61(3), AUGUST, 2009

237

Table 2. Stomach contents of Calidris minutilla collected 19 January 2002 (percent by
mass [mg]).

specimen No.

Coieoptera
(% mass)

Minerals
(% mass)

Seeds
(% mass)

Unidentified
(% mass)

20021

0.0

32.9

0.0

67.1

20022

5.0

40.7

0.0

54.4

20023

4.3

12.0

0.4

83.2

20024

36.6

0.0

62.2

20025

5.5

39,2

0.0

55.3

20026

3.0

28.4

0.0

68.6

20027

1.8

37.8

0.0

60.4

Mean

3.0

32.5

0.06

64.5

terans were much less abundant in 2002 specimens than
Chironomids were in 1998 specimens, Coleopteran prey are harder
and less energetically rewarding due to increased need for
mechanical digestion using minerals and extended time required for
digestion compared to soft-bodied chironomid larvae. Minerals
also take up space that could otherwise be used for additional prey
contents. Indeed, a high percentage of minerals were found in
stomachs from 2002, either due to accidental intake or potentially to
aid maceration of hard-bodied beetles. Alternatively, habitat
differences where birds were foraging in 1998 compared to 2002
could account for the differences observed in stomach contents.

Results suggest that sandpipers wintering at reservoirs in the
region are generalists in diet, with prey capture dependent on
ephemeral populations of aquatic invertebrates and terrestrial
invertebrates existing on or near the shoreline. It is likely that
hydrologic features play some role in aquatic insect populations at
such sites. For example, in 1998 when specimens contained
chironomids, foraging occurred on mudflats under conditions of
slowly receding reservoir levels with no recent runoff events.
However, in 2002, when specimens contained primarily
coleopterans, recent runoff events had caused reservoir levels to
rise, flooding shorelines and providing a source of invertebrates that
would more commonly be found in streams or terrestrial habitats

238

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3, 2009

than on reservoir shorelines (e.g., Elmidae and Curculionidae).
Rising water levels flooding flats covered with Bermudagrass and
winter temperatures in clear, clean water would allow the
occurrence of a unique aquatic insect community compared to
drought periods with turbid, receding water levels.

Colwell & Landrum (1993) found nonrandom distributions of
sandpipers in relation to fine-scale variations in prey abundance.
Sandpipers at this study site appear to use available habitats equally
(Kasner 1999), and may choose prey opportunistically as they are
encountered in ephemeral aggregations. Hockey et al. (1992)
suggest that nonbreeding wader populations are proportionate to the
carrying capacity of coastal wetlands along a latitudinal gradient. If
this is also applicable to freshwater, inland sites, the results of this
study suggest that chironomid and coleopteran populations may be
a limiting factor for sandpipers wintering in west-central Texas.
Local abundances of sandpipers in winter are low relative to the
extent of habitat that is generally available, suggesting ecological
factors other than habitat may be limiting. Further investigation of
prey populations is necessary to answer these questions. Future
research should consider the spatio-temporal dynamics of
invertebrate abundance during the winter months at the site.

Acknowledgments

We would like to thank Dr. Richard Harrel and Dr. Ned Strenth
for assistance and verification of insect identification. We thank
Dr. Kim Withers for helpful suggestions to improve this
manuscript. Collections made under permits held by T. Maxwell
(USFWS Permit MB674149-1, Texas Scientific Collecting Permit
SPR-0290-021).

Literature Cited

Alexander, S. A., K. A. Hobson, C. L. Gratto-Trevor & A. W. Diamond. 1996.
Conventional and isotopic determinations of shorebird diets at an inland stopover:
the importance of invertebrates and Potamogeton pectinatus tubers. Can. J.
Zool., 74(6):1057-1068.

TEXAS J. SCI. 61(3), AUGUST, 2009

239

Colwell, M. A. & S. L. Landrum. 1993. Nonrandom shorebird distribution and fine-
scale variation in prey abundance. Condor, 95:94-103.

Davis, C. A. 1996. Ecology of Spring and Fall Migrant Shorebirds in the Playa
Lakes Region of Texas. Unpublished Ph.D. dissertation, Texas Tech Univ.,
Lubbock, Texas, 204pp.

DeLeon, M. T. 1996. Use of habitat and behavior of migrant shorebirds in North
Dakota. Unpublished M.S. thesis, Texas Tech Univ., Lubbock, Texas, 97pp..

Hockey, P. A. R., R. A. Navarro, B. Kalejta & C. R. Velasquez. 1992. The riddle of
the sands: Why are shorebird densities so high in southern estuaries? Am. Nat.,
140(6):961-979.

Kasner, A. C. 1999. Effects of abiotic factors on inland wintering Least Sandpipers
(Calidris minutilla). Unpublished M.S. thesis, Angelo State Univ., San Angelo,
Texas, 48pp.

Merritt, R. W. & K. W. Cummings. 1984. An Introduction to the Aquatic Insects of
North America, 2"^* ed. Kendall/Hunt Publishing Co., Dubuque, Iowa, 722pp..

Neill, R. L. 1992. Recent trends in shorebird migration for North-central Texas.
Southw. Nat., 37(l):87-88.

Schneider, C. J. 1987. Comparative Ecology of Two Guilds of Shorebirds on the
South Texas Coast. Unpublished M.S. thesis, Univ. Texas, Austin, Texas, 45pp.

Shepherd, P. C. F. & J. S. Boates. 1999. Effects of a commercial baitworm harvest
on Semipalmated Sandpipers and their prey in the Bay of Fundy Hemispheric
Shorebird Reserve. Con. Biol., 13(2):347-356.

Skagen, S. K. & H. D. Oman. 1996. Dietary flexibility of shorebirds in the western
hemisphere. Can. Field-Nat., 1 10(3):419-444.

Smith, K. G., J. C. Neal & M. A. Mlodinow. 1991. Shorebird migration at artificial
fish ponds in the prairie-forest ecotone of Northwestern Arkansas. Southw. Nat.,
36(1):107-113.

Tartar, D. G. 1997. A field checklist birds of the Concho Valley Region, Texas, 3"^^^
revision.

Tsipoura, N. & J. Burger. 1999. Shorebird diet during spring migration stopover on
Delaware Bay. Condor, 101:635-644.

Wilson, W. H. Jr. & K. Parker. 1996. The life history of the amphipod, Corophium
volutator: The effects of temperature and shorebird predation. J. Exp. Mar.
Biol.,Ecol. 196:239-250.

ACK at: kasnera(gwbu.edu

240

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 3

IN RECOGNITION OF THEIR ADDITIONAL SUPPORT OF
THE TEXAS ACADEMY OF SCIENCE DURING 2009

PATRON MEMBERS

Goldberg, Stephen R.
Killebrew, Don W.
Marsh, David S.
Strenth, Ned E.

SUSTAINING MEMBERS

Davidson, David L.
Kowalski, Joseph L.
Kruger, Joseph M.

Lee, Thomas E. Jr.
Valdes, Arcadio

SUPPORTING MEMBERS

Collins, James
Harper, Donald E., Jr.
Hettinger, Deborah D.

Looney, Michael
Lundelius, Ernest L., Jr.
Mckinney, Larry
Sieben, John
Simpson, Lynn
Stevens, Fred
Weller, Milton W.

THE TEXAS ACADEMY OF SCIENCE, 2009-2010

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2008 Christopher M. Ritzi, Sul Ross State University
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2009 Ana B. Christensen, Lamar University
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SECTIONAL CHAIRPERSONS

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Volume 61
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THE TEXAS JOURNAL OF SCIENCE

Volume 6 1 , No. 4 November, 2009

CONTENTS

Gas Exchange Rates of Sun and Shade Leaves of Sophora secundlflora

(Leguminosae, Texas Mountain Laurel).

By Mitsuru Furuya and O. W. Van Auken 243

Characterization of Arsenic-Tolerant Bacterial Cultures from the
Lower Laguna Madre of South Texas.

By Gemma A. Berlanga, Michael W Persans, Thomas M. Eubanks

and Kristine L. Lowe 259

Diversity and Abundance of Unionid Mussels in Three Sanctuaries on the
Sabine River in Northeast Texas.

By Neil B. Ford, Jessica Gullett and Marsha E. May 279

Morning and Evening Densities of White-Winged and Mourning Doves in
the Lower Rio Grande Valley, Texas.

By Michael F. Small, Margaret L. Collins, John T. Baccus

and Steven J. Benn 295

General Notes

Prevalence of Hematozoan Parasites (Apicomplexa) in some common
Passerine Birds (Passeriformes) from East-Central Oklahoma.

By Michael D. Bay and Kenneth D. Andrews 3 1 1

Notes on Reproduction of the Knob-Scaled Lizard, Xenosaurus grandis
(Squamata: Xenosauridae), from Veracruz, Mexico.

By Stephen R. Goldberg 3 1 7

Population Dynamics of an Established Reproducing Population of the Invasive
Apple Snail {Pomacea insularum) in Suburban Southeast Houston, Texas.

By Colin H. Kyle, Matthew K Trawick, James P. McDonough

and Romi L. Burks 323

Recognition of Special Members 328

Index to Volume 61 (Subject, Authors & Reviewers) 329

Postal Notice

335

THE TEXAS JOURNAL OF SCIENCE
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Jim R. Goetze, Laredo Community College
Associate Editor for Botany:

Janis K. Bush, The University of Texas at San Antonio
Associate Editor for Chemistry:

John R. Villarreal, The University of Texas-Pan American
Associate Editor for Computer Science:

Nelson Passos, Midwestern State University
Associate Editor for Geology:

Ernest L. Lundelius, University of Texas at Austin
Associate Editor for Mathematics and Statistics:

E. Donice McCune, Stephen F. Austin State University

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American Association for the Advancement of Science,

TEXAS J. OF SCI. 61(4):243-258

NOVEMBER, 2009

GAS EXCHANGE RATES OF

SUN AND SHADE LEAVES OF SOPHORA SECUNDIFLORA
(LEGUMINOSAE, TEXAS MOUNTAIN LAUREL)

Mitsuru Furuya* and O. W. Van Auken

Department of Biology, University of Texas at San Antonio
San Antonio, Texas 78249
"^Current address:

7-1 1-20 Nakagawa, Tsuzuki-ku Yokohama-shi
Kanagawa-ken, 224-0001 Japan

Abstract.-Gas exchange rates of sun and shade leaves of Sophora secundiflora
(Leguminosae, Texas Mountain Laurel) were measured. Maximum photosynthetic rates
(^max), light saturation points, dark respiration rates (Rd), stomatal conductance rates and
ambient light levels for sun leaves were significantly different than shade leaves. There
were no significant differences between sun and shade leaves for the light compensation
point, transpiration rates, leaf water potential, leaf mass or leaf area. Mean A^ax rates
were 12.94 ± 0.58 (± one SE) pmolC02/mVs for sun leaves and 7.49 ± 1.35 pmolC02/
m^/s for shade leaves. Mean Rd rates were 3.48 ± 0.44 pmolC02/m^/s for sun leaves and
2.28 ± 0.21 pmolC02/m ^/s for shade leaves. Mean g^ rates were 0.24 ± 0.02 molH20/m
Vs for sun leaves and 0.12 ± 0.02 molH20/m Vs for shade leaves. A^ax rates for sun
leaves were within levels expected for sun plants, but A^ax rates for shade leaves were
fairly high as well. Sophora secundiflora sun and shade leaves also had high Rd rates
which suggest that this is not a sun or shade species. The ranges of many of the other
measurements overlapped and were not significantly different suggesting this species is
an intermediate or facultative species. This would explain why it is present in full sun
and shaded or partially shaded habitats in central Texas and other areas where it is found.

Communities are composed of a variety of species that occur
together spatially and temporally (Begon et al. 2006). Measuring the
density and basal area of plants found in these terrestrial communities
is relatively easy to do (Van Auken et al. 2005). In addition, taking
population data from these plant communities, ranking the species
from highest to lowest, and showing which ones have the greatest
basal area or density is relatively routine. Species in a community are
found together because they require or are tolerant of the conditions
present in specific areas or habitats. However, sorting out the
characteristics or factors that determine why a species is dominant is
much more challenging. It is also very difficult to ascertain why
similar species fit together or exist in communities. Some species are
restricted to open habitats, some to woodlands or forests, and others
seem to occur at the edge or in-between communities (Begon et al.

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

2006). Factors that cause these limitations might be biotic or abiotic,
but are not always easy to delineate. Light levels, soil depth, soil
moisture, nutrient levels, competition between species, biotic
characteristics, or combinations of these factors are all possibilities
(Valladares & Niinemets 2008).

In the central Texas Edwards Plateau region, savannas are
associated with Juniperus-Quercus woodlands or forests in many
areas (Van Auken et al. 1981; Van Auken & McKinley 2008; Van
Auken & Smeins 2008). A species found in many of these
communities is Sophora secundiflora (Ort.) DC. (Leguminosae, Texas
Mountain Laurel) (Van Auken et al. 1981). It has not been reported
as a dominant species, but a secondary species with lower density and
basal area (Van Auken et al. 1981). However, it forms almost mono-
specific communities in some open habitats especially on drier and
shallow soils including ridges and some south facing slopes. In
addition, isolated plants are found below the canopy in some
communities. As a Legume, it could establish in disturbances or low
nutrient soils, but no studies were found concerning its establishment
or successional status.

Physiological differences between plants native to shady habitats
compared to those found in full sun are fairly well known (Begon et
al. 2006; Valladares & Niinemets 2008). There are major differences
in most photosynthetic characteristics. Shade plants usually have
lower photosynthetic rates at high light levels, light saturate at lower
light levels (light saturation), have lower light compensation points
(photosynthetic rate equals respiration rate) and lower dark respiration
rates (Boardman 1977; Larcher 2003; Valladares & Niinemets 2008).
Sun plants characteristically have higher transpiration and stomatal
conductance rates (Young & Smith 1980). Some species display
adaptive crossover and are capable of acclimating to high or low light
environments, thus they could have a broader ecological niche
(Givnish 1988; Givnish et al. 2004). In addition, shade leaves from
plants grown in full sun have been used as surrogates for plants grown
in shade conditions to understand a species’ ecological requirements
(Hamerlynck & Knapp 1994).

FURUYA & VAN AUKEN

245

Gas exchange rates of sun and shade leaves of Sophora
secundiflora were measured. Based on most information about this
species, it was hypothesized that S. secundiflora is a sun plant and
would have a high maximum photosynthetic rate, light saturation
point, light compensation point, respiration, conductance, and
transpiration compared to shade adapted plants.

Methods

Study species. -Sophora secundiflora (Texas Mountain Laurel,
Leguminosae) is a shrub or small tree. It is a North American species
native to Texas, New Mexico, and northeastern Mexico (Correll &
Johnston 1979). Sophora secundiflora grows 5 to 8 m tall with a
maximum canopy diameter of about 3 m. It has 5 to 13 cm long,
pendulous clusters of purple or blue, fragrant flowers in spring.
Leaves are usually dark-green, glossy, evergreen, alternate, and
pinnately compound. Sophora secundiflora is reported to grow in full
sun or partial shade on well-drained soil (Enquist 1987). It seems to
tolerate hot, windy conditions, alkaline or wet, but not compacted soil.
Plants may have a deep root system although characteristics of the
rooting system are unreported.

Study area -This field study was carried out on the southern edge
of the Edwards Plateau region of central Texas just south of the
Balcones Escarpment (Correll & Johnston 1979; Van Auken et al.
1981; Van Auken & McKinley 2008). The Balcones Escarpment
consists of a rough, well-drained area, with elevations increasing from
approximately 213 m above mean sea level (AMSL) at the southern
edge to between approximately 500 and 700 m AMSL near the center,
but in most places the increase in elevation is abrupt. Most of the
subsurface of the area is Cretaceous limestone, and soils are usually
shallow, rocky or gravelly on slopes, and deep in broad valleys and
flats (Taylor et al. 1962; NRCS 2006). Soils are dark colored and
calcareous with usually neutral or slightly basic pH.

Mean annual temperature of the area is 20.0°C with monthly
means ranging from 9.6°C in January to 29.4°C in July (NOAA
2004). Mean annual precipitation is 78.7 cm and bimodal, with peaks

246

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

occurring in May and September (10 J cm and 8.7 cm, respectively),
with little summer rainfall and high evaporation (Thomthwaite 1931;
NOAA 2004).

Vegetation is Juniperus-Quercus savanna or woodland and is
representative of savannas and woodlands found throughout this
region, but higher in woody plant density than savanna communities
farther to the west (Van Auken et al. 1979; 1980; Van Auken et al.
1981; Smeins & Merrill 1988). The high density woody species are
Juniperus ashei (Ashe juniper) and Quercus virginiana {=Q.
fusiformis, Live oak) followed by Diospyros texana (Texas
persimmon) and Sophora secundiflora. Associated with these
woodlands are relatively small grasslands and sparsely vegetated
intercanopy patches or gaps (openings between canopy patches) (Van
Auken 2000). The major herbaceous species below the canopy is
Carex planostachys (Cedar sedge) (Wayne & Van Auken 2008). In
the grasslands and gaps Aristida longiseta (Red three-awn), Bouteloua
curtipendula (Side-oats grama), Bothriochloa {=Andropogon)
laguroides (Silver bluestem), B. ischaemum (KR bluestem), various
other C4 grasses, and a variety of herbaceous annuals are common
(Van Auken 2000).

Measurements exchange rates as a ftinction of light level or
photosynthetic flux density (PFD) were measured and plotted for sun
and shade leaves of Sophora secundiflora plants (Hamerlynck &
Knapp 1994). There were five replications (individual leaves)
measured for each leaf type. All plants sampled were approximately 2
m talk Sun leaves were on the outermost, southern facing canopy
branches of plants growing in the full sun and shade leaves were on
the innermost branches of these plants. Shade leaves from full sun
plants were used as surrogates for shade plants (Hamerlynck & Knapp
1994).

Measurements were made within ± 3 hr of solar noon with a LI-
COR® infrared gas analyzer (LI-6400). Irradiances were generated by
the LI-COR LED red-blue light source using a light curve program
with the LI-COR, a gas flow rate of 400 pmol/s, and a CO2

FURUYA & VAN AUKEN

247

concentration of 400 jimol/mol. One mature, undamaged, fully
expanded leaflet per replication and leaf type was used with the 2 by 3
cm chamber. The LI-COR 6400 was run at approximate ambient
summer, midday, daytime temperature (35®C) and relative humidity
(50%), and was calibrated daily. Response data were recorded after at
least two minutes when a stable total coefficient of variation was
reached (<0.3%), usually less than five minutes. Light response
curves were started at a PFD of 2000 pmol/m^/s for sun leaflets and
1800 jLimol/mVs for shade leaflets and then decreased to 1800 or 1600,
1400, 1200, 1000, 800, 600, 400, 200, 100, 75, 50, 25, 10, 5, and 0
jLimol/mVs (16 or 17 total measurements).

The measurements included net photosynthesis, stomatal
conductance, and transpiration. Repeated measure ANOVAs were
utilized to determine if significant differences occurred between
leaflet types. A one way ANOVA was used to determine if net
photosynthesis, stomatal conductance, and transpiration were
significantly different between the PFD’s tested (Sail et al. 2001).
Shapiro- Wilks tests were used to test for normal distributions and the
Bartlett’s Test was used to test for homogeneity of variances. Data
were log transformed for analyses due to unequal variances as
necessary.

The maximum photosynthesis (A^ax), PFD at A^ax, transpiration at
^max, conductance at A^ax, light saturation point, dark respiration, light
compensation point, and the quantum yield efficiency (initial slope)
were determined for each replicate, and means were calculated. The
^max was the highest net photosynthesis rate. Light saturating
photosynthesis was the PFD when the slope of the initial rate line
reached the A^ax- Dark respiration was the gas exchange rate at a PFD
of 0 pmol/m^/s (y-intercept of the line for the initial slope or rate).
The light compensation point was calculated as the PFD when the
photosynthetic rate = 0 pmol COi/m^/s (x-intercept of the line for the
initial slope or rate). The quantum yield efficiency or initial slope was
calculated using the dark value and increasing PFDs until the
regression coefficient of the slope decreased (150 pmol/m^/s PFD)
(Wayne & Van Auken 2009).

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

A pooled Mest (Sail et al. 2001) was used to detect significant
difference between leaf types for photosynthetic rates (Amax), light
saturation, dark respiration, transpiration at A^ax, conductance at A^ax,
and quantum yield efficiency. Due to unequal variances, light
compensation and PFD at A^nax were compared using a standard ^test.
Significance level for all tests was 0.05. Ambient PFD was also
measured for each sun and shade leaf with the LI-COR® integrating
quantum sensor at the time the light response curves were initiated
(LI-COR, Inc, Lincoln, NE).

Pre-dawn xylem water potential -VxQ-ddmn xylem water potential
{Px) measurements were made for leaves of each plant (Scholander et

al. 1965). Xylem water potential of sun and shade leaves was
measured with the model 1000 PMS® pressure chamber (PMS,
Instrument, Co. Corvallis, OR). Samples of each leaf type were
collected with a sharp knife and put in a zip lock plastic bag with a
wet paper towel between 4:30 and 5:00 am. The plastic bag was put
in a cooler with ice to insure that would not change.
measurements were made within 45 minutes of harvest.

Soil moisture measurements -W ohxmoXnc soil moisture measure¬
ments were made using time domain reflectometry (TDR) with a
TRIME portable TDR soil moisture meter (TRIME-FM) (MESA
System Co. Medfield, ME). TDR is a transmission line technique
used to determine soil water content by inserting two parallel metal
rods in a soil matrix to make measurements (Topp & Reynolds 1998;
Noborio 2001). Soil water content was measured in five positions
below the canopy of each plant. The five positions were the four
cardinal compass points and the site next to the bole of the plant. Soil
water content of five plants was sampled between 10:00 and 11:00

am. The site next to the bole of the plant was on the south side, and
the other locations were approximately 10 cm from the bole. Data
from five sites (north, south, east, west, and the site next to the plant)
around each plant were pooled and a mean and standard error was
determined.

FURUYA & VAN AUKEN

249

Figure L Photosynthetic light response curves for sun (♦) and shade (□) leaves of
Sophora secundijlora. There were significant differences in photosynthetic rates for
sun and shade leaves of S. secundiflora (repeated measured ANOVA, F=7.6934,
P^O.0242). Error bars represent ± one standard error of the mean. Different letters
between light levels indicate significant differences between sun leaves (upper case)
or shade leaves (lower case) and between leaf types (*).

Leaflet area and total leaflet dry mass-ArQ?i (LA) and total dry
mass (Mooney & Gulmon 1982) of five sun and five shade leaflets
were measured to determined mass per unit area (LMA) (g/cm^). The
sun and shade leaflets previously used to make gas exchange
measurements were collected to determine leaf area and total dry
mass. Leaflets were dried at 60®C to a constant mass prior to
weighing.

Results

The photosynthetic response for the two leaf types was
significantly different over the light levels measured (Repeated
Measure ANOVA; Fig. 1). At PFD’s above 400 pmol/m^/s, sun leaves
had significantly higher photosynthetic rates than shade leaves (/-test;
P < 0.05), while at PFD’s lower than 400 pmol/m^/s, shade leaves
generally had higher rates than sun leaves, but values were not
significantly different (/-test, P > 0.05). Photosynthetic rates for the
sun leaves continued to increase from 600 to 2000 pmolW/sec, but

250

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

Figure 2. Stomatal conductance curves for sun (♦) and shade (□) leaves of Sophora
secundiflora. There were significant differences in stomatal conductance rates for
sun and shade leaves of S. secundiflora (repeated measured ANOKA, F=5.6495,
P=0.0448). Error bars represent ± one standard error of the mean. Different letters
between light levels indicate significant differences between sun leaves (upper case)
or shade leaves (lower case) and between leaf types (*).

not significantly. The same was true for the shade leaves except there
were no significant differences between treatments at 200 and 1800
jLimol/mVs, the highest light treatment used for the shade leaves.
Stomatal conductance of the sun and shade leaves were significantly
different over the light levels examined (Repeated Measure ANOVA\
Fig. 2). However, the stomatal conductance rate for the sun leaves did
not change significantly over the light levels tested (One way
ANOVA) and the same was true for the shade leaves. However, the
sun leaves had higher conductance rates at most light levels tested
(Student’s Mest). Transpiration rates of the two leaf types were not
significantly different over the light levels measured (Repeated
Measure ANOVA; Fig. 3). At all light levels tested, the transpiration
rate was higher for the sun leaves, but not significantly higher
(Students Mest; P > 0.05).

The maximum photosynthetic rate (A^ax) of S. secundiflora sun
leaves was 12.9 pmol C02/m^/s and occurred at the maximum PFD

FURUYA & VAN AUKEN

251

’E

0 500 1000 1500 2000

PFD (|jmol/m2/s)

Figure 3. Transpiration curves for sun (♦) and shade (□) leaves of Sophora secundiflora.
There were no significant differences in transpiration rates for sun and shade leaves
of S. secundiflora (repeated measured ANOVA, F=L4992, P=0.2556). Error bars
represent ± one standard error of the mean. Different letters between light levels
indicate significant differences between sun leaves (upper case) or shade leaves
(lower case) and between leaf types (*).

measured (2000 jimol/m^/s) (Table 1). The rate was approximately
L7 times that of the shade leaves (7.5 pmol C02/m^/s), which
occurred at a PFD of 1000 pmol/m^/s). Light saturation for sun leaves
was 355 pmol/m^/s, which was significantly higher than the light
saturation of shade leaves (210 pmol/m^/s). The light compensation
point of sun leaves was 61 pmol/m^/sec which was 1.5 times greater
than the light compensation point of shade leaves at 41 pmol/m^/s but
not significantly different (Table 1). The dark respiration of sun
leaves was 3.5 pmol COa/m^/s and was 1.5 times greater than the dark
respiration of shade leaves at 2.3 pmol C02/m^/s and these values
were significantly different. The quantum yield efficiency or the
initial slope (slope of the line from 0-150 pmol/m^/s) was not
significantly different between the sun or shade leaves (Table 1).
Conductance at was significantly higher for sun leaves compared
to shade leaves, but there was no significant difference in transpiration
rates (Table 1).

252

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

Table 1. Means and standard error (SE) for the maximum net photosynthetic rates
(Amax), light level at the Amax, light saturation (Lsat), light compensation points
(Rd/a), dark respiration rates (Rd), initial slope or quantum yield efficiency (IS),
stomatal conductance rates, transpiration rates (E), leaflet areas (LA), leaflet
mass, leaflet mass per unit area (LMA), ambient light level, and pre-dawn xylem
water potential (T^x) of sun and shade leaves for Sophora secundiflora and soil
moisture (%).

Leaf type (Mean ± one SE)

Parameter

Sun

Shade

^max ()LimolC02/mVs)

12.9 (0.6) a*

7.5 (1.4) b

PDF at.4niax

2000 (0)

1000 (0)

^sat

355 (35) a

210 (20) b

Rd/a (pmol/mVs)

61 (13) a

41 (3) a

Rd (pmolC02/mVs)

3.5 (0.4) a

2.3 (0.2) b

IS (pmolC02/(jLimol quanta)

0.0490 (0.0022) a

0.0550 (0.0021) a

gs(molH20/mVs)at^max

0.24 (0.02) a

0.12 (0.02) b

E (mmolH20/m^/s) at

6.24 (0.78) a

4.67 (0.93) a

LA (cm“)

12.0 (1.9) a

12.8 (1.0) a

EM (g)

0.22 (0.03) a

0.23 (0.03) a

LMA (g/cm^)

0.019 (0.001) a

0.018 (0.001) a

Ambient Light (pmol/mVs)

1216 (78) a

394(95) b

'Px (Mpa)

-0.90 (0.10) a

-1.12 (0.06) a

Soil Moisture (%)

9.4 (1.6)

* Means in the same row followed by the same letter are not significantly different at the
0.05 level.

There were no significant differences in the leaflet area, leaflet
mass or specific leaflet mass (Table 1). Ambient light levels were
significantly different for the two leaf types, with the sun leaves being
exposed to 3.1 times more light (Table 1). In addition, the of the
sun and shade leaves was not significantly different, but the surface
soil was dry, having only 9.4 ±1.6 % moisture.

FURUYA & VAN AUKEN

253

Discussion

As hypothesized, sun leaves of Sophora secundiflora, which is
commonly found growing in high light environments in Central Texas
(Van Auken et al. 1981), had a high maximum photosynthetic rate
(^max)? typical of species found growing in open habitats (Begon et al.
2006). Other photosynthetic parameters, including light saturation,
light compensation, dark respiration, conductance, and transpiration,
were high for sun adapted leaves. However, shade leaves had
relatively high gas exchange parameters as well. These responses are
not consistent with findings for shade plants, but for plants that are
sun plants or are intermediate or facultative species (Boardman 1977;
Hull 2002; Larcher 2003; Givnish et al. 2004; Begon et al. 2006;
Valladares & Niinemets 2008).

Although Sophora secundiflora is a native species with a fairly
broad distribution and is used as an ornamental over much of its
range, very little is known about its photosynthetic capabilities. No
studies have been identified which evaluate the physiological
responses or growth responses of this species to light levels or other
factors. The parameters measured for both leaf types suggest that this
species is not a sun or shade species, but an intermediate or facultative
species. Therefore, it can grow in sun or shade or intermediate light
habitats.

In general, true understory species have much lower photo¬
synthetic rates than the rates reported for S. secundiflora in the current
study. Photosynthetic rates of three understory montane spruce
forests species found in central Europe all had rates 27-65 % lower
(3.4 - 5.5 pmol C02/m^/s) than S. secundiflora shade adapted leaves
(Hattenschwiler & Komer 1996). In addition to these lower ^max
rates, the European forest species reached light saturation at much
lower irradiance (~200 pmolW/s) levels than sun leaves of S.
secundiflora (355 ± 36 pmolW/s), although shade leaves of S.
secundiflora were light saturated at about the same level as the forest
understory species (210 ± 20 pmolW/s). Arnica cordifolia
(Asteraceae), an herbaceous perennial which grows in the understory
of lodgepole pine forests in southeastern Wyoming, also had

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

photosynthetic rates 44-53% lower than S. secundiflora shade adapted
leaves (3.5 - 4.2 |LimolC02/mVs), but reached light saturation at the
same level as the S. secundiflora sun leaves at 350 pmol C02/m^/s
(Young & Smith 1980). Polygonum virginianum (Polygonaceae), an
herbaceous perennial found in the forest understory and at the forest’s
edge in the eastern United States, had an A^ax of ~ 3 pmolC02/mVs at
a light saturation of ~ 500 pmolW/s (Zangerl & Bazzaz 1983).
Car ex planostachys from the central Texas Edwards Plateau
Juniperus woodland understory had an value of 4.9 ± 0.3
pmolC02/m^/s which was 65 % of the value for shade leaves of S.
secundiflora and reached light saturation at 151 ± 43 pmol/m^/s
(Wayne & Van Auken 2009). While S. secundiflora is typically
found growing in open habitats or the edge of woodlands, its high
Araax for shade adapted leaves compared to other herbaceous shade
plants would suggest it is an intermediate or facultative sun species,
and could grow in a variety of light environments including edge
habitats.

True sun plants are adapted to high light conditions and have high
rates of gas exchange. For example, Abutilon theophrasti an early
successional herbaceous perennial had rates between 15-25
pmolC02/mVs (Wieland & Bazzaz 1975; Bazzaz 1979; Munger et al.
1987a; Munger et al. 1987b; Hirose et al. 1997; Lindquist &
Mortensen 1999). Two oaks of gallery forest in tall grass prairies of
northeastern Kansas, Quercus muehlenbergii and Q. macrocarpa had
^max rates 1.40-2.02 times higher than S. secundiflora at 18-26
pmolC02/mVs for sun leaves and 11-13 pmolC02/mVs or 1.47-1.73
times higher for shade leaves (Hamerlynck & Knapp 1994).

Plants can acclimate to the variability of the light environment in
which they live, particularly early successional species or plants from
disturbed (open) communities (Bazzaz & Carlson 1982). Polygonum
pensylvanicum, a colonizing annual of open fields, had an^^ax of ~ 12
|LimolC02/mVs at ~ 1500 pmol/mVs when plants from a shaded-habitat
(200 pmol/mVs) were measured (Bazzaz & Carlson 1982; Zangerl &
Bazzaz 1983); however the rate was - 24 pmol/mVs at ~ 1800
pmol/m^/s when plants from a full sun habitat were measured (Bazzaz

FURUYA & VAN AUKEN

255

& Carlson 1982). The high light individuals we sampled in the
present study were growing in an area which received 1216 ± 78
pmol/m^/s (~ 60% full sunlight). We might expect that individuals
from higher light environments could have higher maximum
photosynthetic rates, while those from lower light environments
would be lower. Further studies would be needed to determine if S.
secundiflora does acclimate to variability in the light environment as
reported for other species.

The dark respiration of sun leaves of S. secundiflora (3.5 ± 0.4
pmolC02/m^/s) is similar to other sun-adapted plants (Hamerlynck &
Knapp 1994). This rate is 1.52 times higher than the of shade
adapted leaves of S. secundiflora. However, the for shade adapted
leaves of S. secundiflora is about 4.6 times higher than rates for other
shade adapted species (Hirose & Bazzaz 1998; Hull 2002). Dark
respiration for shade-adapted species is typically lower than sun-
adapted species, due to the lower metabolism of shade-adapted
species (Bjorkman 1968; Bazzaz & Carlson 1982). Polygonum
pensylvanicum grown at 200 pmol/mVs had a respiration rate of ~ 0.5
pmolC02/m^/s, although the rate was twice as high when plants from
full sun were measured (Bazzaz & Carlson 1982).

Other photosynthetic parameters reported in this study for S.
secundiflora are similar to those values reported in the literature.
Quantum yield efficiency reported here (0.049 and 0.055
pmolC02/pmol quanta, sun leaves and shade leaves respectively) are
within the range or similar to values (0.035 - 0.052 pmolC02/pmol
quanta) reported for other species (Hirose et al. 1997). Stomatal
conductance and transpiration reported in the current study were
similar to other studies, however many factors affect these parameters
(Wieland & Bazzaz 1975; Zangerl & Bazzaz 1984; Yun & Taylor
1986; Munger et al. 1987a; Munger et al. 1987b; Stafford 1989).

The sun and shade leaves of S. secundiflora show some distinct
and varied photosynthetic responses. These differential physiological
responses to various light levels more than likely are contributors to
the niche breadth observed for this species in the field. In general.

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

resource utilization is spatially partitioned among species along
complex environmental gradients^ such as changes in light from open
areas to woodland or forest edges (Wayne & Van Auken 2009). The
ability of S. secundiflora to reach high photosynthetic rates at lower
light level, its light saturation, and light compensation point allows it
to exist in a variety of communities. At light levels below 300
ILimolC02/m^/s, data suggests that other more shade tolerant species
would probably be able to out-compete S. secundiflora. At light
levels above 300 pmolC02/mVs, S. secundiflora could dominate, in
part because it has photosynthetic rates as high as or higher than most
co-occurring species (Grunstra 2008).

Acknowledgements

We would like to thank the Center for Water Research at The
University of Texas at San Antonio for support provided for the senior
author. Help from M. Grunstra and J. K. Bush during various stages
of the study is truly appreciated.

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oscar.vanauken@utsa.edu

TEXAS J. OF SCL 61(4):259-278

NOVEMBER, 2009

CHARACTERIZATION OF

ARSENIC-TOLERANT BACTERIAL CULTURES FROM
THE LOWER LAGUNA MADRE OF
SOUTH TEXAS

Gemma A. Berlanga, Michael W. Persans, Thomas M. Eubanks
and Kristine L. Lowe

Department of Biology, University ofTexas-Pan American
Edinburg, Texas 78539

Abstract -Two forms of arsenic are found in the environment: As(V) and
As(III), the latter being more toxic, water-soluble, and mobile. Microorganisms may
increase the mobility of arsenic by reducing As(V) to As(III); however, detoxification
and immobilization can occur via the oxidation of As(III) to As(V). The US EPA has
set a minimum contaminant level of 10 parts per billion (ppb) for arsenic in drinking
water. The research objective was to confirm the presence of arsenic-tolerant
bacteria in the Lower Laguna Madre of south Texas. Sediment samples were
collected and inoculated into growth media which contained either 2 mM As(III) or 2
mM As(V) to enrich for As(III)-tolerant and As(V)-tolerant bacteria, respectively.
Twenty six (26) As(III)-tolerant and 12 As(V)-tolerant cultures were obtained. Most
isolates were small white colonies of Gram-positive rods. Biochemical tests using
commercially-made test strips showed that As(V)-tolerant isolates displayed greater
resource usage compared to As(III)-tolerant isolates but overall, few cultures
demonstrated a wide-range of biochemical capabilities. Isolates with distinct
morphological and biochemical phenotypes were subjected to Polymerase Chain
Reaction (PCR) amplification and sequencing of the 16S rRNA genes to identify the
bacteria. Closest sequence matches were to the eubacterial genera Mycoplasma,
Salinispora, Frankia, and Pelodictyon. These results suggest that the Lower Laguna
Madre is inhabited by a diverse group of microorganisms able to tolerate toxic
concentrations of different arsenic species.

Arsenic contamination is a significant problem in the
environment because arsenic is toxic and carcinogenic. Although
arsenic may exist in several oxidation states, it is typically found in
two forms in the environment; arsenate (As(V)), which is typically
insoluble, and arsenite (As(III)), which is water soluble and more
toxic (Stumm & Morgan 1996). Insoluble As(V) often precipitates
to the bottom of bodies of water making the arsenic immobile, but
soluble As(III) is mobile and of much greater concern. The US
EPA has set a Maximum Contaminant Level (MCL) of 10 ppb for
total arsenic (As(III) + As(V)) in drinking water (USEPA 2001). A

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1995 Study showed sediment arsenic levels greater than 27,000 ppb
in the Upper Laguna Madre (Barrera et al. 1995).

The Laguna Madre is a hypersaline estuary, one of only five in
the world, with an average salt content ranging from 35 to 45 ppt;
however, salt concentrations higher than 80 ppt have been reported
(Quannen & Onuf 1993; Tunnell & Judd 2002; Whelan et al. 2005).
The high salinity of the Laguna Madre is due to the fact that there
are few freshwater inputs into the estuary and few outlets. The
Lower Laguna Madre receives fresh water discharged from the
highly-impacted Arroyo Colorado and from precipitation; however,
the estuary is shallow (average depth 1.5 m) (Tunnell & Judd
2002). Salts concentrate during periods of drought and high rates
of evaporation occur in the summer due to the warm regional
climate of South Texas, Regional pollution has resulted in toxic
chemical concentrations in water, sediment, and animals of the
Laguna Madre (Davis et al. 1995). Among the contaminants at
toxic concentrations is arsenic. Much of the arsenic detected in the
Lower Rio Grande Valley of Texas and along the US-Mexico
border is attributed to past usage of arsenical pesticides, but may
also be related to sewage treatment discharge into regional
waterways and non-point sources (Davis et al. 1995; Tunnell &
Judd 2002).

Davis et al. (1995) reported that large amounts of arsenic were
present in the Lower Laguna Madre; however, this study did not
distinguish between insoluble As(V) and toxic, soluble As(III).
Arsenic is also of concern in the Arroyo Colorado, a waterway in
South Texas that is part of the natural drainage system for the
Lower Rio Grande Valley and lies on the north of the Rio Grande
Delta. The Arroyo Colorado is part of a floodway system, receives
treated wastewater from several towns, runs through the Port of
Harlingen, and discharges into the Lower Laguna Madre. Elevated
arsenic levels have been detected in the Arroyo Colorado, most
likely due to agricultural runoff (Wells et al, 1988). Thus, arsenic
in the Laguna Madre presents a potential threat to the ecosystem

BERLANGA, ET AL.

261

and may impact organisms in the lagoon. Some microorganisms
may increase the mobility of arsenic by reducing As(V) to As(III)
(Macy et al. 2000), However, aquatic plants and microorganisms
may lessen the toxicity of arsenic by immobilizing it via oxidation,
methylation, or accumulation (Oremland et al. 2002, Bentley &
Chasteen 2002, Schmoger et al. 2000).

As(V) and As(III) can change oxidation state by chemical or
biological processes. In marine sediments, As(V) can form a
variety of insoluble mineral compounds that are structurally similar

to phosphates. These occur when arsenate anions (ASO4 ") react
with transition metals, such as iron and manganese (Smedley &
Kinniburgh 2002). Reduction of metal-arsenate minerals can occur
under low redox (reducing) conditions, thus liberating the arsenic as
As(III). As(III) typically exists as soluble arsenic acids, such as
H3ASO3, at marine pH values (Smedley & Kinniburgh 2002).

Biologically-mediated transformations of arsenic depend on the
form of arsenic involved. As(V) may be reduced to As(III) by
anaerobic sediment bacteria through dissimilatory anaerobic
respiration (Newman et al. 1998, Macy et al. 2000). Plants can
reduce As(V) to As(III) through a detoxification mechanism
(Meharg & Hartley- Whitaker 2002) or the bioaccumulation of
arsenic may take place in plant tissues (Davis et al. 1995). Arsenic
can also be detoxified via biomethylation in microorganisms, algae,
plants, and animals (Wang et al. 2004). Methylation detoxifies
arsenic by making it volatile, converting it to gaseous forms (Qin et
al. 2006). Reduction of As(V) to As(III) can potentially release
arsenic bound to sediment particles or bound in minerals, and may
impact the mobility of arsenic in the Laguna Madre ecosystem.
Increased arsenic mobility will potentially impact plants and
animals by introducing toxic, soluble As(III) into sediment pore
waters and estuary waters. Conversely, As(III) oxidation is an
important detoxification reaction since the resulting As(V) is less
toxic and less bioavailable (Oremland et al. 2002).

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

The macro-ecology of the Laguna Madre has been extensively
studied for decades. Several rare, endangered, threatened, and
migratory animal species feed or nest in the Laguna Madre.
Despite this, little is known about the microbial ecology of the
Laguna Madre and the importance of microorganisms in nutrient
and chemical cycling within the sediments of the ecosystem.
Interest in microbial ecology has increased dramatically as the
importance of microorganisms within sediments and subsurface
environments are related to biogeochemistry, bioremediation, and
biotechnology (Atlas & Bartha 1998; Nealson 1997). The interplay
between biologically-mediated As(III) oxidation and biologically-
mediated As(V) reduction is important in determining the fate of
arsenic species in the Laguna Madre. In this research, culture-
based and molecular methods were used to characterize bacterial
populations that demonstrated tolerance to As(III) or As(V).

Materials and Methods

Sample collection. samples (approximately 50 g) were
collected in March 2007 from the top 10 cm of sediment at four
sites in the Lower Laguna Madre (Figure 1). Samples were
transported on ice to the laboratory. The collection sites were:
LMT-050 (N26°08n7.4”,W97M0’4L0”), located near the South
Padre Island Wastewater Treatment Plant; South Bay (SB;
N26°02’48” W97Mr3.3”); site ABC (N26°10’09.7” W97Mr
05.3”) and LMT-051 (N26°10’09.7”,W97Mr05.3”). All sites
were south of the mouth of the Arroyo Colorado and have been
described previously (Whelan et al. 2005).

Enrichment cultures for arsenic-oxidizing and arsenic-reducing
bacteria. from each site was homogenized and 1 g was
placed into sterile 20-mL glass vials containing 10 mL of minimal
liquid medium (Lowe et al. 2000). The final concentrations and
composition of the medium was (per liter): CH3COO Na [15 mM],
(NH4)2S04 [0.9 mM], K2HPO4 » 3H2O [0.57 mM], KH2PO4 [0.33
mM], NaHCOs [0.2 mM], NazEDTA ^ 2H2O [7pM], H3BO3 [6pM],
FeS04 • 7H2O [0.6 ^M], C0CI2 • 6H2O [0.5 (iM], Ni(NH4)SOA •

BERLANGA, ET AL.

263

Figure 1. Map of the Lower Laguna Madre showing the sampling locations. (From
Whelan et al. 2005).

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

6H2O [0,5 juM], Na2Mo04 • 2H2O [0,4 |liM], Na2Se04 (anhyd) [0,15
juM], MnS04 * H2O [0,13 |aM], ZnS04 ^ 7H2O [0,1 jaM], CUSO4 *
5H2O [0.02 jLiM], casamino acids [0.01% w/v], Vitamin Bi [0.001
mg mL“^], L-arginine HCl [0.02 mg mL“^], L-glutamic acid [0,02
mg mL'^], L"glutamine [0.02 mg mF^], L-serine [0.04 mg mL"^],
MgS04 *7H20 [1 mM], CaCl2 • 2H2O [0.5 mM] and NaCl [3%
w/v]. After sterilization, the medium was amended with either
As(V) (Na2HAs04 •7H2O) or As(III) (NaAs02) to a final
concentration of 2 mM. Vials with As(V) as the terminal electron
acceptor were placed in glass canisters with an anaerobic gas
generating system (BBL GasPak, Becton Dickenson Co.,
Cockeysville, MD), Vials containing As(III) were incubated in an
aerobic environment. Cultures were performed in triplicate. Vials
were incubated at 25°C until there was visible turbidity. Once there
was visible growth in the vials, the samples were streaked onto the
appropriate medium as described above plus 1.5% Bacto Agar
(Difco-BBL, Sparks, MD) to isolate pure colonies.

Isolation of As (III) -tolerant and As (V) -tolerant bacteria-ThQ
As(III) enrichment vials that showed bacterial growth were
inoculated onto appropriate agar consisting of minimal media with
As(III), These samples were incubated aerobically for 14 d at
25°C. Resulting isolated colonies were sub-cultured on the same
medium and incubated aerobically at 25°C. Agar plates that had
been inoculated from the As(V) enrichment vials were incubated at
25 °C in glass canisters with an anaerobic gas generating system
(BBL GasPak, Becton Dickenson Co., Cockeysville MD) for 30 d.
The resulting cultures were sub-cultured on the same medium and
incubated in an anaerobic environment at 25°C.

Random As(III)-tolerant and As(V)-tolerant bacterial isolates
from the arsenic media agar plates were chosen for ftirther
investigation. Isolates were characterized by Gram-stain and by
observed colony morphology (i.e., color, size, shape). Biochemical
profiles for isolates were generated using API 20E® strips

BERLANGA, ET AL.

265

(bioMerieux Inc., Durham, NC). Isolates were stored in 25%
glycerol at -80°C for molecular studies.

API 20E® strips 20E® strips include enzymatic tests for
fermentation or oxidation of glucose, mannitol, inositol, sorbitol,
rhamnose, saccharose, melibiose, amygdalin, and arabinose, along
with nitrate reduction to nitrite and nitrate reduction to nitrogen gas.
API 20E® strips also test for the presence of P-galactosidase,
arginine dihydrolase, lysine decarboxylase, ornithine
decarboxylase, citrate utilization, H2S production, urease,
tryptophan deaminase, indole production, acetoin production
(Voges ~ Proskauer), and gelatinase. API 20E® tests were
performed according to the manufacturer’s instructions. The
number and types of positive tests were tabulated for the isolates
and used to construct biochemical profiles of the As(III)“tolerant
and As(V)-tolerant cultures. The API 20E® profiles were used to
compare biochemical phenotypes amongst the isolates.

Molecular identification of isolates -yio\QCu\^x identification of
the isolated As(III)-tolerant and As(V)-tolerant bacteria was
performed by the Polymerase Chain Reaction (PCR) amplification
and sequencing of the 16S ribosomal RNA (16S rRNA) genes
(Sambrook & Russell 2001). Genomic DNA was extracted from
the bacterial isolates using the QIAamp DNA Mini Kit (Qiagen,
Valencia, CA). Extracted genomic DNA integrity was verified on a
1% agarose electrophoresis gel. Genomic DNA concentration was
quantified by reading the absorbance in a UV-VIS spectro¬
photometer at 260 nm. Purity of the extracted DNA was
determined by the ratio of the absorbances at 260 and 280 nm.
Ratios of 260/280 absorbance measurements were between 1.7 and
2.0 for all samples; thus, the extraction yielded mostly DNA
(Sambrook & Russell 2001).

A sample (50 ng) of template genomic DNA was placed in a 0.5
mL thin-walled PCR tube with 25 pL of PCR Master Mix
(Promega, Madison, WI). PCR primers used to amplify the 1505
bp target of the 16S rRNA gene were 5'- AGA GTT TGA TCC

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

TGG CTC AG ~ 3' (forward) and 5'- ACG GCT ACC TTG TTA
CGA CTT - 3' (reverse) (Integrated DNA Technologies, Coralville,
lA). The final concentration of each primer was 10 pM. The total
PCR mixture volume was 50 pL. The PCR mixture was placed into
a MyCycler PCR thermocycler (Bio Rad, Hercules, CA) and heated
to 95°C for 6 min in order to initially denature the template DNA.
After the initial denaturation, 40 cycles were run with the following
conditions: denaturing at 95°C for 30 sec, annealing of the primers
at 52°C for 30 sec, and primer extension at 72°C for 30 sec. A final
extension at 72°C for 1 min was done and the PCR products were
held at 4°C. An aliquot (5 pL) of the PCR products was visualized
by electrophoresis on a 1% agarose gel. The remaining PCR
product mixture was purified using a Wizard PCR Clean Up Kit
(Promega, Madison, WI). Purified PCR products were used for
DNA sequencing described below.

For sequencing, a second round of PCR was done using a
commercially-available sequencing kit (Genome Lab DTCS Quick
Start Kit; Beckman Coulter, Fullerton CA) according to the
manufacturer ’ s instructions. Dye-tagged dideoxynucleotides
(ddUTP, ddGTP, ddCTP, and ddATP) were added to terminate
elongation (Sambrook & Russell 2001). The resulting PCR product
was loaded into an automated DNA sequencer (CEQ 8000 Genetic
Analysis System; Beckman Coulter, Fullerton, CA). The sequences
were compared to known bacterial sequences available in the
National Center for Biotechnology Information Basic Local
Alignment Search Tool (BLASTN) database (www.ncbi.nml.nih.
gov/BLAST) to identify the microorganisms (Altschul et al. 1997).

Results and Discussion

Pure colonies {n = 26) were successfully isolated from As(III)
enrichments [Table 1]; 12 pure colonies were isolated from As(V)
enrichments [Table 2]. Every isolate cultured in As(III) media was
100% Gram-positive [Table 1]. The As(III) Gram stain results
were unusual because marine sediments usually contain mixtures of
Gram-positive and Gram-negative bacteria (Atlas & Bartha 1998).

BERLANGA, ET AL.

267

Table 1 . Morphology of Laguna Madre bacteria isolated from As(III)-contaimng media.
As(III)"tolerant bacteria were isolated from the Laguna Madre from different sites.
Isolates were streaked onto minimal media supplemented with 2 mM As(III) and
incubated aerobically for 14 d. Cells were Gram-stained and the colony morphology
was visually observed and recorded. All isolates were Gram-positive and smooth in
appearance.

Isolate

Sediment Source

Shape

Appearance

AS3-A

South Bay

bacillus

white, small , round

AS3-B

South Bay

bacillus

white, round

AS3--C

ABC

bacillus

orange, round

AS3--D

South Bay

bacillus

white, irregular

AS3-E

South Bay

bacillus

ivory, small, irregular

AS3-F

LMT-050

bacillus

clear, small, irregular

AS3-G

LMT-050

bacillus

light yellow-orange, small, round

AS3-H

LMT-050

bacillus

light yellow, small, round

AS3-I

LMT-050

bacillus

pink, small, round

AS3-J

ABC

bacillus

peach, small, round

AS3-K

South Bay

coccus

transparent, small, round

AS3-L

South Bay

coccus

white, small, round

AS3-M

South Bay

bacillus

yellow/orange, small, round

AS3-N

LMT-050

bacillus

ivory, small, round

AS3-0

LMT-051

bacillus

white, small, round

AS3-P

ABC

bacillus

white, small, round

AS3-Q

South Bay

bacillus

white, small, round

AS3-R

South Bay

bacillus

white, small , irregular

AS3-S

South Bay

cocco-bacillus

white, small, round

AS3-T

ABC

bacillus

orange, small, round

AS3-U

ABC

bacillus

orange, small, round

AS3-V

LMT-051

bacillus

pink/orange, small, round

AS3-W

LMT-050

bacillus

ivory, small, round

AS3-X

LMT-051

bacillus

peach, small, round

AS3- Y

ABC

bacillus

ivory, tiny, round

AS3-Z

ABC

bacillus

orange, small, round

The high percentage of Gram-positive bacteria may reflect the
toxicity of arsenic, which may result in high selection pressure for
Gram-positive organisms. Once inside a cell, As(III) disrupts
protein folding and protein-DNA interactions (Norman 1998).
Gram-positive cells have a thicker cell wall compared to Gram¬
negative cells. The thicker cell wall might make it more difficult
for As(III) to enter the cell. The isolates enriched in As(III)
medium were rod-shaped or bacillus (23/26), cocci (2/26) and

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Table 2. Morphology of Laguna Madre bacteria isolated from As(V)-containmg media.
As(V)-tolerant bacteria were isolated from the Laguna Madre from different sites.
Isolates were streaked onto minimal media supplemented with 2 mM As(V) and
incubated anaerobically for 30 d. Cells were Gram-stained and the colony
morphology was visually observed and recorded.

Isolate

Sediment

Source

Gram

Stain

Shape

Colony Appearance

ASS - A

LMT-050

bacillus

transparent, small, round, smooth

AS5-B

ABC

bacillus

transparent, small, round, smooth

AS5-C

South Bay

bacillus

ivory, small, irregular, smooth

AS5-D

LMT-051

cocco-bacillus

ivory, small, irregular smooth

AS5-E

LMT-050

coccus

transparent, small, round, smooth

AS5-F

LMT-050

coccus

ivory, round with filaments, smooth

AS5-G

LMT-050

cocco-bacillus

ivory clear, small, irregular, smooth

AS5-H

LMT-051

coccus

white, small,, round, smooth

ASS -I

LMT-050

cocco-bacillus

white, small, irregular, smooth,

AS5-J

LMT-050

bacillus

transparent, small, round, smooth

ASS -K

LMT-050

coccus

ivory, small, irregular, filamentous

ASS-L

LMT-050

coccus

transparent, small, round, smooth

cocco-bacillus (1/26) [Table 1]. The typical colony morphology of
the As(III) isolates was smooth and round but varied in color and
shape. The morphologies observed ranged from small, white
punctiform colonies to irregular-shaped, colored colonies. Colony
colors included pink and orange [Table 1].

The isolates cultured in As(V) were either Gram-positive (7/12)
or Gram-negative bacteria (5/12) and were bacillus (4/12), cocci
(5/12), and cocco-bacillus (3/12) [Table 2].

The colony morphologies also varied for the As(V) isolates.
Colonies were mostly irregular and white, or almost transparent
[Table 2]. Two of the As(V) isolates, AS5-F and AS5-K, were
filamentous; both were isolated from LMT-050 [Table 2]. In
general, isolates from the As(V) enrichments displayed slower
growth rates than those isolated from As(III) media (data not
shown). However, this was most likely due to incubation
conditions - anaerobic for As(V) versus aerobic conditions for
As(III) - and not arsenic toxicity.

BERLANGA, ET AL.

269

For the As(III) isolates {n = 26) tested with API 20E® strips,
three of the isolates were positive for P-galactosidase and six were
positive for gelatinase; two were positive for mannitol and
saccharose oxidation; one was positive for melibiose utilization.
The remaining seventeen tests were negative for all As(III) isolates
[Table 3].

Among the As(V) isolates {n= 12) tested with API 20E® strips,
five were positive for p~galactosidase, two were positive for
arginine dihydrolase, and seven were positive for gelatinase. In
addition, three were positive for glucose, mannitol, rhamnose,
saccharose, amygdalin, or arabinose oxidation. The remaining
thirteen tests were negative for all As(V) isolates [Table 4].

The high number of negative tests made putative identification
of the isolates difficult; however, the isolates from the anaerobic
As(V) enrichment cultures appeared more metabolically diverse
than those cultured in the aerobic As(III) enrichment cultures. The
enzymatic flexibility that some As(V)-tolerant bacteria displayed
may be due to the lower toxicity of As(V) relative to that of As(III)
or As(III) may select for characteristics that were not included on
the test strips and not observed.

There were a low number of positive API tests from the
microorganisms isolated from As(III) enrichments [Table 3].
Among the As(III)-tolerant isolates tested, isolates AS3-J from site
ABC and isolate AS3-K from South Bay each displayed only three
positive tests which was the highest number of positive tests [Table
3], Isolates AS3-J and AS3-K are likely to be different
microorganisms because their morphology was different and their
API profiles displayed two tests in common but differed in one
[Table 3], AS3-J and AS3-K were both able to oxidize mannitol
and saccharose; however, only AS3-J was able to metabolize
melibiose while AS3-K produced p-galactosidase indicating lactose
utilization [Table 3]. Isolates AS3-G, AS3-H, AS3-T, AS3-U,
AS3-W and AS3-Z tested positive for gelatinase [Table 3];

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Table 3. Biochemical profiles of As(III)-tolerant bacteria. Twenty-six (26) As(III)-
tolerant bacteria were isolated from the Laguna Madre sediment. The isolates’
metabolic activities were tested using API 20E® strips. A plus sign (+) indicates that
the isolate was positive for the test; a negative sign (-) indicates a negative reaction

for the test. Only isolates that displayed at least one positive test are shown.

Isolate

AS3- AS3- AS3- AS3-

AS3- AS3- AS3- AS3- AS3-

AS3-

E G H J

K M T U W

ONPG

+ - - -

+ + - - -

ADH

_ _ _ _

_____

LDC

- _ - -

_____

ODC

- - - -

_____

CIT

- - - -

_____

H2S

_ _ _ _

_____

URE

_ _ _ _

_____

TDA

- - - -

_____

IND

- - - -

_____

_ _ _ _

_____

GEL

- + + -

- - + + +

GLU

_ _ _ _

_____

MAN

- - - +

+ _ _ _ _

INO

- - - -

_____

SOR

_ _ _ _

_____

RHA

_ _ _ _

_____

SAC

- - - +

+ _ _ _ _

MEL

- - - +

_____

AMY

- - - -

_____

ARA

- - - -

_____

N02

- - - -

_____

- - - -

_____

Tests: ONPG, |3-galactosidase activity; ADH, arginine dihydrolase; LDC, lysine decarboxylase;
ODC, ornithine decarboxylase; CIT, citrate utilization; H2S, hydrogen sulfide production;
URE, urease; TDA, tryptophan deaminase; IND, indole production; VP, acetoin production
(Voges-Proskaur); GEL, gelatinase; GLU, glucose; MAN, mannitol; INO, inositol; SOR,
sorbitol; RHA, rhamnose; SAC, sucrose; MEL, melibiose; AMY, amygdalin; ARA,
arabinose; N02, nitrate reduction to nitrite; N2, nitrate reduction to nitrogen gas.

moreover, AS3-T, AS3-U and AS3-Z all showed similar colony
morphologies and are potentially the same organism despite being
isolated from different locations in the Laguna Madre [Table 1].

The As(V) isolates from the sediment samples collected at LMT-
050 had the highest number of positive API tests for all isolates
tested [Table 4]. The LMT-050 population included AS5-A, ASS-

BERLANGA, ET AL.

271

Table 4. Biochemical profiles of As(V)-tolerant bacteria. Twelve (12) As(V)-tolerant
bacteria were isolated from the Laguna Madre sediment. The isolates’ metabolic
activities were tested using API 20E® strips. A plus sign (+) indicates that the isolate
was positive for the test; a negative sign (-) indicates a negative reaction for the test.

Isolate

ASS

-A

-B

-C

= D

= E

-F

-G

-H

-I

- J

-K

-L

ONPG

ADH

LDC

ODC

CIT

H2S

URE

TDA

IND

GEL

GLU

MAN

INO

SOR

RHA

SAC

MEL

AMY

ARA

N02

Tests: ONPG, P-galactosidase activity; ADH, arginine dihydrolase; LDC, lysine decarboxylase;
ODC, ornithine decarboxylase; CIT, citrate utilization; H2S, hydrogen sulfide production;
URE, urease; TDA, tryptophan deaminase; IND, indole production; VP, acetoin production
(Voges-Proskaur); GEL, gelatinase; GLU, glucose; MAN, mannitol; INO, inositol; SOR,
sorbitol; RHA, rhamnose; SAC, sucrose; MEL, melibiose; AMY, amygdalin; ARA,
arabmose; N02, nitrate reduction to nitrite; N2, nitrate reduction to nitrogen gas.

E, and AS 5-1, which were all isolated from the same sediment
sample and have similar morphologies; however, they varied
slightly in their biochemical profiles. The three isolates all tested
positive for P-galactosidase and utilization of glucose, mannitol and
rhamnose, but only AS5-A and AS5-J tested positive for arginine
dihydrolase and arabinose metabolism, and AS5-E and AS5-J tested
positive for amygdalin utilization [Table 4]. Even though AS5-A,
AS5-E, and AS5-J isolates from LMT-050 were the most
metabolically diverse, they all tested negative for gelatinase, which

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

overall had the highest number of positive reactions [Table 4]. The
highest number of gelatinase-positive isolates were obtained in the
anaerobically-grown As(V) isolates compared to the aerobic As(III)
isolates. Only 6/26 As(III) isolates were positive for gelatinase
whereas 7/12 of the As(V) isolates were gelatinase positive [Table
3 and 4], Gelatinase is a digestive enzyme necessary to break down
gelatin, a protein found in animal connective tissue, which is
sometimes degraded by bacteria during biofilm formation
(McNamara et ah 1997). The production of gelatinase is also used
to differentiate and identify anaerobes (Whaley et ah 1982). The
overall high number of positive gelatinase tests suggests that some
of these organisms may be anaerobic decomposers of organic
matter (McNamara et al. 1997).

Isolates (5/12) from the As(V) enrichments were positive for fJ-
galactosidase activity which indicates that some As(V)-tolerant
bacteria from the Laguna Madre can use lactose as a carbon source
[Table 4]. This was unexpected because lactose is not a commonly
used carbon source in marine environments as it does not easily
absorb to sediment particles (Sansone et al. 1987). Acetate is more
easily absorbed to sediment particles and is more readily available
to sediment bacteria; thus, it was used as the principle carbon
source in the isolation medium. No isolate was capable of nitrate
reduction [Table 3 and 4] which was also unexpected because many
facultative and anaerobic sediment bacteria are able reduce NO3".
NOs' is a better electron acceptor energetically compared to As(V)
(Dowdle et ah 1996).

Results from the 16S rRNA gene sequencing showed an array of
positive matches. Table 5 shows the top BLASTN result for the
As(III) and As(V) isolates that were successMly sequenced. The
top similarity match (% Match) is shown. Isolate AS3-E displayed
high similarity to Mycoplasma hyopneumoniae. Isolate AS3-K
showed good similarity to Salinispora tropica CNB-440, a marine
Gram-positive bacterium with a high percentage of G+C bases in
the DNA (Maldonado et al. 2005; Williams et al. 2005). Isolate

BERLANGA, ET AL.

273

Table 5. As(III)-tolerant and As(V)-tolerant bacteria isolated from the Laguna Madre
sediment. The 16S rRNA genes from the isolates were amplified by PCR,
sequenced, and compared to other known bacteria using the BLASTN search engine
and data-base. The isolates were matched to the highest percentage match. The top
BLASTN matches and the percent sequence match are shown.

Isolate

Sediment Source

Best Match

Percent Match

AS3-E

South Bay

Mycoplasma hyopneumoniae

100

AS3-K

South Bay

Salinispora tropica CNB-440

AS5-B

ABC

Frankia alni ACN14A

100

AS5-E

LMT-050

Pelodictyon luteolum

AS5-B had high similarity with Frankia alni strain ACN14A which
is a Gram-positive, nitrogen-fixing bacterium that can live
symbiotically with some non-legume plants (Atlas & Bartha 1998;
Normand et al. 2007.). Isolate AS5-E showed similarity with
Pelodictyon luteolum, a photosynthetic, green-sulfur bacterium
(Overmann & Tuschak 1997).

Although the 16S rRNA gene sequencing resulted in a variety of
potential matches, the identity of the organisms is yet to be
conclusively determined. This is because the sequencing results do
not agree with the morphological and physiological data for the
isolates. For example, isolate AS3-E was observed to be a Gram¬
positive, rod-shaped bacterium [Table 3] and showed high identity
with M. hyopneumoniae. Mycoplasma hyopneumoniae is related to
low G+C Gram-positive organisms, it lacks a cell wall and is
associated with a mild, chronic form of pneumonia that affects pigs
(Atlas & Bartha 1998; Minion et al. 2004). However, mycoplasma
cells are typically small, coccus-shaped cells with a convex dense
region in the center sometimes referred to a “fried egg” appearance
(see Madigan & Martinko 2005). Thus, the observed morphology
of this isolate does not agree with what would be expected for a
mycoplasma cell and it is unclear why a swine pathogen would be
present in Laguna Madre sediments. Recent research has involved
inserting arsenic resistance genes as genetic markers for the

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

development of genetically-modified vaccines against Mycoplasma
hyopneumoniae in infected swine (Matic 2008). What role this may
play in arsenic tolerance is not yet known, especially for
environmental strains of the bacterium.

Isolates AS5-B and AS3-K showed high similarity to F. alni
and S. tropica, respectively. Organisms are Gram-positive, aerobic
bacteria, which does agree with these observations [Table 3 & 4]
yet these bacteria are typically filamentous and produce
hyphae/mycelia when grown in laboratory media (Benson &
Silvester 1993; Maldonado et ah 2005). This morphology was not
observed for either isolate. The lack of agreement between the
morphology and sequencing data may be due to PCR amplification
of extraneous sequences or limited sequence information in the
BLASTN database that did not allow for an ideal match. Thus,
As(III)“tolerant and As(V)-tolerant isolates from the Laguna Madre
appear to be diverse groups of yet unidentified microorganisms.

The results of this investigation provide insights into the
potential for arsenic mobilization in the Lower Laguna Madre.
Arsenic is carcinogenic and toxic, especially As(III); thus it is
advantageous to have information into possible arsenic cycling in
the Lower Laguna Madre due to recreational use of the lagoon and
fisheries that occupy regions adjacent to the lagoon. The Laguna
Madre is a rare hypersaline estuarine ecosystem, and it is important
to ascertain whether arsenic and other contaminants are potentially
harmful to the ecosystem. Arsenic was used as a pesticide for
several decades and cannot be degraded like an organic pollutant; it
can only be converted to different forms. Thus, once in the
ecosystem, it will stay there in some manner. It is likely that the
shallow water levels, low water flow, high evaporation rates, high
salinity and pH of the Laguna Madre affect the concentration and
speciation of the arsenic. To what extent is not yet known.

These results suggest that there is the potential for arsenic
mobilization (i.e., reduction of As(V) to As(III)) in the Laguna

BERLANGA, ET AL.

275

Madre due to the presence of several different arsenic-tolerant
bacterial types isolated under arsenic-reducing (As(V)) conditions.
These organisms displayed different metabolic abilities and
different biochemical profiles, suggesting that they are not the same
species. Results of the 16S rRNA gene sequencing experiments
support this. Furthermore, these organisms were isolated from
several locations in the Lower Laguna Madre with varying
environmental conditions, which suggests that they might be
widespread in the ecosystem.

If there is the potential for As(V) reduction to As(III), it is
important to consider how this may affect the arsenic toxicity in the
Laguna Madre and how this in turn will affect the biota in the
ecosystem. One would think that bacterial cells reducing As(V) to
As(III) would be making a toxic environment for themselves and be
affected by their own metabolic products. Why this does not
appear to affect the bacterial cells is not known but it necessitates
the involvement of some type of arsenic tolerance. It is possible
that the produced As(III) may re-oxidize by chemical or biological
means, or that the cells may have some mechanism of resisting the
produced As(III). Some bacterial cells have arsenic resistance
mechanisms such as efflux pumps that keep arsenic out of the cell
(Newman et al. 1998). Such mechanisms were not tested for in the
current study.

The presence of As(III)-tolerant bacteria in the lagoon implies a
possible mechanism for counteracting the mobilization of arsenic if
these As(III)-tolerant organisms can also oxidize As(III) to As(V).
Future studies will include the determination of the relative
abundance and density of As(III)-oxidizing and As(V)-reducing
bacteria in the Laguna Madre, comparing arsenic oxidation and
reduction at different sites, conducting seasonal studies on arsenic
transformations in the Laguna Madre, and the detection of genes
associated with arsenic oxidation and arsenic reduction (Saltikov &
Newman 2003; Murphy & Saltikov 2007). Additionally arsenic

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

reduction rates can be conducted in the future to conclusively
demonstrate arsenic mobilization in the Laguna Madre.

Acknowledgements

We wish to thank Hudson DeYoe, Thomas Whelan III and the
UTPA Coastal Studies Lab for assistance in sample collection.
Funding was supplied by the Howard Hughes Medical Institute
Undergraduate Science Education Program Grant (#520006321),
the NSF-Collaborative Research at Undergraduate Institutions
(CRUI) Program, the Guerra Honors Program, and the Faculty
Research Council at UTPA.

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KLL at: klowe@utpa.edu

TEXAS J. OF SCI. 61(4):279-294

NOVEMBER, 2009

DIVERSITY AND ABUNDANCE OF UNIONID MUSSELS
IN THREE SANCTUARIES ON THE SABINE RIVER
IN NORTHEAST TEXAS

NeD B. Ford, Jessica Gullett and Marsha E. May*

Department of Biology, University of Texas at Tyler
Tyler, Texas 75799 and

^Wildlife Diversity Branch, Texas Parks and Wildlife Department
4200 Smith School Road, Austin, Texas 78744

Abstract.-Populations of freshwater mussels (Bivalvia: Unionidae) are declining
for reasons that are primarily anthropogenic. The Texas Administrative Code lists 18
freshwater mussel sanctuaries (“no-take” areas) within Texas stream segments and
reservoirs with three being on the Sabine River in northeast Texas. Visits to each
Sabine River sanctuary were made multiple times between April and September 2007
with two goals: to establish species richness by locating rarer species not found in
earlier surveys and to collect unionid data that could be used to evaluate abundances
among the sanctuaries. Using timed and density surveys (0.25 meter square quadrats)
1596 individuals of 18 unionid species were recorded. Densities ranged from means
of over 21 per meter square in one sanctuary to 3.6 per meter square in the sanctuary
nearest the dam at Lake Tawakoni. Because a range of sizes were found for several
species at the two downstream sanctuaries, recruitment evidently occurs. One of the
healthiest unionid populations in these areas was Fusconaia askewi, which is a
species of concern in the Texas Wildlife Action Plan. The mussel beds were found
only in small, isolated patches in any sanctuary and silting over of beds with sand
from bankfalls was evident throughout the river. Whether these sanctuaries will
sustain all species within the upper Sabine River is questionable and it will be
important to continue to monitor them.

It is increasingly evident that freshwater mussels (Bivalvia:
Unionidae) are important components of riverine ecosystems
(Christian & Berg 2000; Vaughn & Hakenkamp 2001; Howard &
Cuffey 2006; Vaughn & Spooner 2006). Unionids have historically
dominated lotic environments of the southeastern United States in
terms of benthic biomass (Parmalee & Bogan 1998) and in
undisturbed rivers may exceed other assemblages by an order of
magnitude (Stray er et al. 1994). With the greatest diversity in the
world, the continental United States supported nearly 300 species of
unionid mussels (Neves 1993; Turgeon et al. 1998). However, their
sedentary, slow-growing and long-lived (many > 25 years) life
histories plus early parasitic phase usually requiring a host fish (Kat

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

1984; Watters 1994; Vaughn & Taylor 2000) has made them highly
susceptible to human impacts such as wetland drainage, channeli¬
zation, sedimentation, dredging, pollution, invasive species and
impoundments (Vaughn & Taylor 1999; Howells et al. 2000;
Lydeard et al. 2004). The decline of North American unionid
populations has been occurring for over a century (Neves et al.
1997; Vaughn 1997) with the extinction of at least 35 species and
up to 65% imperiled to some degree (Turgeon et al. 1998). For
many states, including Texas, the extent of freshwater mussel
decline is simply not known (Bogan 1993; Layzer et al. 1993;
Neves 1993).

At least 52 species of unionids occur in Texas and yet our
understanding of their conservation status is quite limited (Howells
et al. 1996; 1997). Although specific data are not available, it
seems likely that East Texas unionid populations have declined at
least equivalent to unionids in the other regions (Neck 1986;
Howells 1997; Bordelon & Harrel 2004; Ford & Nicholson 2006).
The human population of the region has been growing rapidly with
dramatically increasing demands on its water resources, as
illustrated by the 31 large reservoirs on its rivers (Ford & Nicholson
2006). Most of the rivers of eastern Texas are isolated from each
other and many drain independently into the Gulf of Mexico. For
example, the Sabine River begins in North-Central Texas in Hunt,
Rains and Van Zandt counties and flows southeasterly first to a
large reservoir on the border with Louisiana, Toledo Bend Lake,
then ends in Sabine Lake, an estuary of the Gulf of Mexico.
Additionally, one reservoir was built at the headwaters of the river.
Lake Tawakoni, and a second, Lake Fork, is located in Wood,
Rains and Hopkins counties and contributes much initial flow to the
river through Lake Fork Creek. These two reservoirs have likely
changed the river downstream both in flow patterns and
geomorphology (Ford & Nicholson 2007). The only recent
published surveys of mussels from the Sabine River drainage are
for Lake Tawakoni (Neck 1986), a study on the Old Sabine Bottom
Wildlife Management Area, which has the Sabine River as its

FORD, GULLETT & MAY

281

northern border (Ford & Nicholson 2006) and a number of
unpublished Texas Parks and Wildlife Department (TPWD) surveys

(summarized in Howells 1997; 2006).

Mussel harvesting in Texas has occurred for over one hundred
years, however, the intense overharvesting that occurred in the
Mississippi Valley apparently did not occur in Texas (Howells et al.
1996). Although harvesting permits were required, little effort to
monitor the mussel-harvesting was implemented until the
increasing demand from the cultured pearl industry for American
mussel shell begin in the late 1970s. In 1992, the Texas
Administrative Code listed 28 freshwater mussel sanctuaries within
Texas stream segments and reservoirs, but in 2006, Rule 57.157
reduced the number to 18 (Fig. 1). Harvesting is not permitted in
these "no-take" areas with the intention that they will provide adult
unionids producing glochidia for dispersal by fish hosts to non¬
protected areas.

Three of the sanctuaries occur on the Sabine River in Northeast
Texas. Texas Parks and Wildlife Department conducted some
limited surveys at the bridge crossings of these sanctuaries in 1993
(Howells 1995) and 1994 (Howells 1995; 1996a; 1996b) and again
in 2005 and 2006 (Howells 2006). The goal of this study was to
survey unionid mussels throughout the full extent of each sanctuary
to establish total species richness for each and to collect data that
could be used to evaluate densities of mussels within each
sanctuary.

Materials and Methods

Study areas first sanctuary directly below Lake Tawakoni
(hereafter called Lake Tawakoni Sanctuary) begins at the dam at
Lake Tawakoni and ends downstream at State Highway 19 in Rains
and Van Zandt counties. The riverbed from the dam to Highway 19
was obviously heavily impacted by scouring that occurred during
high water releases. Daily discharges in this section of the river
ranged from lows of 5 cfs (cubic feet per second) to high releases of

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

Fig. 1. Texas Mussel Sanctuaries (Texas Administrative Code Title 31, part 2, Ch. 57,
subch. B, rule 57.157); A. Big Cypress Creek in Camp County; Bl. Sabine River in
Rains and Van Zandt counties; B2. Sabine River in Smith, Upshur and Wood
counties; B3. Sabine River in Harrison and Panola counties; C. Angelina River in
Angelina, Cherokee, Jasper, Nacogdoches, Rusk, San Augustine, and Tyler counties;
D. Neches River in Hardin, Jasper, Orange and Tyler counties; E. Trinity River in
Houston, Leon Madison, Trinity and Walker counties; F. Live Oak Creek in Gillespie
County; G. Brazos River in Palo Pinto and Parker counties; H. Guadalupe River in
Kerr County; 1. Concho River in Concho County; J. San Saba River in Menard
County; K. Guadalupe River in Gonzales County; L. San Marcos River in Hays,
Guadalupe and Gonzales counties; M. Pine Creek in Lamar and Red River counties;
N. Sanders Creek in Fannin and Lamar counties; O. Elm Creek in Runnels and
Taylor counties; P. Rio Grande in Webb County.

FORD, GULLET! & MAY

283

over 7000 cfs in just one day (United States Geological Survey
[USGS] 2007). The initial first km is channelized and deep. The
rest consists of mud and silty substratum with large amounts of
detritus and nonorganic trash (plastic and styrofoam).

The second sanctuary below the bridge at Highway 14 (hereafter
called Highway 14 Sanctuary) is located from Farm to Market Road
14 to State Highway 155 in Smith, Upshur and Wood counties. In
this section, the river was relatively wide (20-30 m at low water)
and so a number of shallow sites with mussels and shells were
evident. Some exposure of rocky outcrops of Cretaceous origin
occurred with areas of small cobble. However, a large percentage
of the sanctuary had severe erosion of the steep riverbanks,
including numerous bankfalls. Daily discharges in this section of
the river ranged from a low of 45 cfs to a high of 18,500 cfs in the
year of the study (USGS 2007). This part of the river normally
experiences flooding several times in the winter but during the year
of the survey the high flows occurred during midsummer.

The third sanctuary below the bridge at Highway 43 (hereafter
Highway 43 Sanctuary) is located from State Highway 43 down¬
stream to U. S. Highway 59 in Harrison and Panola counties. The
river in this section was also relatively shallow and wide. However,
daily discharges in this section of the river were more dramatic and
ranged from lows of 1 1 cfs to high releases of 20,400 cfs (USGS
2007). It also had some areas of bedrock and extremely large
boulders. Few reaches of any length with smaller rocks and cobble
were evident.

Sampling techniques sanctuary was surveyed multiple
times between April and September 2007 using two methods. The
total extent of each sanctuary was initially explored by kayaking
during low water with reconnaissance for shells and stream
characteristics appropriate for mussels such as current and the
presence of cobble (Vaughn et al. 1997; Strayer et al. 1997; Strayer
& Smith 2003). Five to seven sites spaced throughout the length of

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

each sanctuary were sampled. At selected sites, timed surveys were
conducted and in areas with adequate mussel numbers, density
surveys were performed. Both methods are necessary as timed
surveys are useM for locating rare species but cannot be used for
statistical comparison between areas (Strayer & Smith 2003).

Timed surveys -VimQ searches were conducted by surveying a
100m stretch of the river visually and tactilely for live and recently
dead mussels in shallow areas and along the banks. Each site was
sampled for a total of one-person hour. All live unionids and shells
that were complete with both valves were collected, identified and
counted. Live specimens were returned to the river. One voucher
of each species was retained in the University of Texas at Tyler
collection and any questionable specimens were collected and sent
to Robert Howells of TPWD for identification.

Density surveys -In timed survey sites where unionids were
abundant (at least 12 per 1 person hour), nearby areas were sampled
using 0.25 meter square quadrats to estimate density (expressed as
mussels per square meter). A random plot design was used
with three starting points (k) and a sample total of 10 quadrats (n)
(Strayer & Smith 2003). An approximate width of river of 20 m
(W) and 100 m for the reach sampled (L) was used. This produced
a distance of three meters between samples. Two surveyors
searched the substratum by hand and excavated all mussels to a
depth of 15 cm until no more specimens were found. Both live and
recently dead (complete with both valves) were identified and
counted. Measurements of length, width and height were taken
only on living unionids.

Data analysis -Ml individuals counted in the timed surveys
were used to calculate a Shannon-Wiener species diversity (H* base
e) and evenness (J') indices. Rank abundance was determined for
unionids for both methods. A Jaccard’s Coefficient of Community
was used to compare species similarity between sanctuaries for the
timed surveys (Brower et al. 1997). Richness and densities were

FORD, GULLET! & MAY

285

(f)

Quadrula verrucosa
Fusconaia askewi
Quadrula apiculata
Lampsilis teres
Truncilla truncata
Potamilus purpuratus
Quadrula mortoni
Obliquaria reflexa
Plectomerus dombeyanus
Leptodea fragilis
Megalonaias nervosa
Pyganodon grandis
Potamilus amphichaenus
Arcidens confragosus
Amblema plicata
Lampsilis hydiana
Lampsilis satura
Anodonta suborbiculata

■

□ Density

■ Timed

0 100 200 300 400 500 600

Number

Figure 2. Comparison of the number of unionid mussels collected at the three sanctuaries
of the Sabine River by sampling technique. The species are ranked by their
abundances in the timed surveys.

compared for the density surveys using a single classification
nested ANOVA with the sanctuary and sites nested with sanctuary
as effects to be tested (SYSTAT® 1 1 2004).

Results

Eighteen unionid species totaling 1596 individuals were found in
the survey of 19 sites in the three sanctuaries on the Sabine River
(Figure 2). Only one species, Anodonta suborbiculata, was found
in this study that was not recorded in Howells' surveys of these
sanctuaries (Table 1). In the timed survey all 18 unionid species
were found (Table 2) whereas in the density survey only 15 were
recorded (Table 3). Four species were abundant in both timed and
density surveys (Fig. 2). These species were Quadrula verrucosa,
Fusconaia askewi, Q. apiculata and Truncilla truncata. Several
other species were abundant in the time surveys, but less so in the

286

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

Table 1. Totals of unionids collected by various methods including visual examination,
wading and snorkeling with hand collection recorded by the Texas Parks and
Wildlife Department at three sanctuaries in the Sabine River. Collections occurred in
1993 and 1994 at the Lake Tawakoni sanctuary, in 1994 also at the Highway 14
sanctuary and in 1994, 1995 and twice in 2005 at the highway 43 sanctuary
(Howells 1995; 1996a; 1996b; 2006).

Lake

Tawakoni

Highway

Total

Fusconaia askewi

Quadrula verrucosa

Lamps il is teres

Leptodea fragilis

Quadrula mortoni

Obliquaria reflexa

Quadrula apiculata

Potamilus purpuratus

Potamilus amphichaenus

Amblema plicata

Plectomerus dombeyanus

Truncilla truncata

Lamps ills satura

Pyganodon grandis

Lamps ills hydiana

Megalonaias nervosa

Utterbackia imbecillis

Arcidens confragosus

Toxolasma texasiensis

Total Number

417

486

Species Richness

density surveys. Timed surveys are generally more successful at
locating rare species but tend to record more of the large species
(Stray er et al. 1997; Vaughn et ah 1997). Seven species were
relatively rare in both methods (Table 2 & 3). Measurements of
live specimens found during the density survey are shown in Table
4. A few species, including Fusconaia askewi, exhibited a wide
range of sizes in the Highway 14 and 43 Sanctuaries.

Jaccard's Coefficient of Community index indicated fewer
species in common between the Lake Tawakoni Sanctuary and the
other sanctuaries (CCj 14 vs. Tawakoni = 0.44%; CCj 43 vs.

FORD, GULLETT & MAY

287

Table 2. Totals of the species collected by the timed method at the three sanctuaries in
the Sabine River.

Lake

Tawakoni

Hwy

Total

Quadrula verrucosa

397

129

526

Fusconaia askewi

169

154

323

Quadrula apiculata

117

182

Lamp sills teres

101

Truncilla truncata

Potamilus purpuratus

Quadrula mortoni

Obliquaria reflexa

Plectomerus dombeyanus

Leptodea fragilis

Megalonaias nervosa

Pyganodon grandis

Potamilus amphichaenus

Arcidens confragosus

Amblema plicata

Lampsilis hydiana

Lampsilis satura

Anodonta suborbiculata

Total number

921

619

1596

Species richness

Shannon diversity

1.63

1.82

2.27

2.13

Evenness

0.74

0.69

0.78

0.73

Tawakoni = 0.50%; CCj 14 vs. 43 = 0.78%). Lake Tawakoni
Sanctuary had the fewest individuals and lowest species richness
(Table 2). The highway 14 had the greatest number of individuals
but the highway 43 sanctuary had the greatest richness (Table 2).

Densities of unionids were significantly different among all
three sanctuaries (F = 7.93; df = 7,102; P < 0.0001). The density
for Lake Tawakoni Sanctuary was the lowest, Highway 43
sanctuary a little higher and the Highway 14 sanctuary was the
highest (Table 3).

Discussion

The Sabine River historically supported approximately 33
unionid species (Howells et al. 1996). Recent surveys have

288

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

Table 3. Totals of the species collected by quadrate sampling method (for density
measurement) at all three sanctuaries in the Sabine River.

Lake

Tawakoni

Hwy

Total

Truncilla truncata

Quadrula apiculata

Quadrula verrucosa

Fusconaia askewi

Obliquaria reflexa

Quadrula mortoni

Lampsilis teres

Potamilus purpuratus

Leptodea fragilis

Plectomerus dombeyanus

Pyganodon grandis

Arcidens confragosus

Megalonaias nervosa

Potamilus amphichaenus

Amblema plicata

Number

268

360

Mean per square meter

3.60

21.44

7.60

Standard error

1.64

3.56

1.08

Species richness

recorded only a portion of those (Neck 1986; Ford & Nicholson
2006; Howells 1997; 2006). Ford & Nicholson's (2006) study on
the Old Sabine Bottom Wildlife Management Area (OSBWMA)
used timed searches and found 13 unionid species in that limited
section of the Sabine River. The major substratum was sand and
clay, neither of which are stable habitats for unionids and may
explain the lower richness at the OSBWMA. The TPWD surveys
involved visual, tactile and some snorkeling searches at the bridges
bordering these sanctuaries in 1993 (Howells 1995) and 1994
(Howells 1995; 1996a; 1996b) and again in 2005 and 2006
(Howells 2006). In those surveys, TPWD found 486 live and
recently dead individuals of 19 species (Table 1). Nearly all these
specimens were from the sanctuary furthest downstream at the
bridges on Highways 43 and 59. The addition to the current survey
of the two species (Utterbackia imbecillis and Toxolasma
texasiensis) that TPWD recorded in their surveys means that the

FORD, GULLETT & MAY

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289

Table 4. Measurements on live unionids collected during density surveys in the three sanctuaries in the Sabine River. Mean Length and
Standard Deviation given with the Range in parenthesis.

LakeTawakoni Hwy 14 Hw>'43

290

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

mussel fauna of the sanctuaries of the Sabine River currently
exhibits 69.7% of the total species that historically occurred in the
Sabine River.

It appears that, as in other east Texas rivers, anthropogenic
impacts are likely reducing unionid diversity and abundance in the
upper Sabine River. Near the dam at Lake Tawakoni the scouring
impact of high water releases on the river substrate was very
evident. The substrate was silt and sand, with only one very small
site (30 m) of cobble where a few unionids were found. Further
downstream, the riverbanks were relatively low and erosion was not
evident but debris from the reservoir was abundant. The most
apparent factor that could be impacting the mussels in other two
sanctuaries was erosion. The Highway 14 Sanctuary had a number
of shallow reaches with cobble, which produced riffles where live
unionids were abundant. This sanctuary also had steep banks and
surrounding agricultural land often came adjacent to the river. A
number of recent bankfalls, which released large amounts of sand
downstream were evident. Highway 43 Sanctuary was also shallow
and wide, but had areas with extremely large boulders and little
cobble. Mussels were not found in reaches with solid bedrock. The
greatest densities were found just downstream of the bridge at
Highway 43, which had stable geomorphology but where smaller
rocks and cobble were present. During the time of this survey,
construction was occurring near the bridge that released sand into
the river. Then a rare summer flood occurred, which shifted the
sand downstream covering much of one of the study sites.

Timed surveys can be used to examine species richness and
ranked species abundances and the unionids in the sanctuaries were
comparable to recent surveys in other east Texas rivers (Howells et
al 2000; Bordelon & Harrel 2004). As is typical of unionid
diversity studies, the rank abundance curve exhibited a few very
abundant species with several intermediately abundant species and
a large number of rare species (Fig. 2). The highest density of
unionids of over 21 per square meter was found in the Highway 14

FORD, GULLETT & MAY

291

Sanctuary (Table 2). This density compares favorably to those in
Little River (17 unionids per square meter) and the Kiamichi river
(20 unionids per square meter) in southeastern Oklahoma (Vaughn
& Spooner 2004). However, it is important to point out that these
sites in the Sabine River were chosen particularly due to the
presence of abundant unionids. Observations made during this
study were that such optimal sites were relatively limited in each
sanctuary. Indeed, there was significant variation among sanctuary
mussel density. The Highway 43 Sanctuary had a mean density of
only 7.6 unionids per square meter and the Lake Tawakoni
Sanctuary had a much lower mean density of 3.6 unionids per
square meter.

It does appear that recruitment of young is occurring in both
Highway 14 and Highway 43 Sanctuaries since a range of sizes
were found for several species (Table 4). One of the healthiest
populations was Fusconaia askewi, which is a species of concern in
the Texas Wildlife Action Plan (TPWD 2006). This population had
very large individuals as well as very small specimens.

Unionid beds were found only in the sanctuaries below
Highway 14 and 43 with only one very small area in the Lake
Tawakoni Sanctuary with enough mussels to do a density survey.
Although some of these beds in the downstream sanctuaries
appeared to have significant numbers of unionids, it is evident that
the beds do not extend for any length but rather are very sporadic.
From limited observations made elsewhere and the literature (Neck
1986; Ford & Nicholson 2006; Howells 1997; 2006), it is likely
that this pattern is true throughout the extent of the upper Sabine
River. To understand the species composition as is exists today in
the sanctuaries would require a landscape level approach detailing
the various habitats that support the different species of unionids.

The impact of high water releases on erosion of the banks and
covering of beds with sand was obviously a problem for these
unionid populations. This was most evident by the lowest density

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THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

of mussel occurring in the sanctuary near the dam. Even though
this area had less erosion of banks (probably because bank-fiill
level was relatively low and water likely spread over the
surrounding wetlands), there were no mussels in most of the
samples. The scouring effect of high water releases is known to
impact mussels near reservoirs. Recruitment of mussels from this
sanctuary is probably limited and it is unlikely that the small areas
of dense mussels in the other sanctuaries will sustain all species
within the upper Sabine River. It will be important to monitor these
sanctuaries in the future.

Acknowledgements

We thank TPWD for the State Wildlife Grant supporting this
research. We thank David Kimberly and Daymon Hail for field
assistance. We also thank Robert G. Howells (TPWD - currently
BioStudies, Kerrville, Texas) and Lyubov E. Burlakova for
confirming species identifications, and Robert Howells and Matt
Troia for reviewing a draft of this paper. Voucher specimens have
been deposited at the University of Texas at Tyler invertebrate
collection.

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Howells, R. G. 1996a. Distributional surveys of freshwater bivalves in Texas:
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NBF at: Neil_Ford@uttyler.edu

TEXAS J. OF SCI. 61(4):295-310

NOVEMBER, 2009

MORNING AND EVENING DENSITIES OF
WHITE- WINGED AND MOURNING DOVES IN
THE LOWER RIO GRANDE VALLEY, TEXAS

Michael F. Small, Margaret L. Collins, John T. Baccus
and Steven J. Benn*

Department of Biology, Wildlife Ecology Program
Texas State University-San Marcos, San Marcos, Texas 78666 and
^Texas Parks and Wildlife Department,

Weslaco, Texas 78596

Abstract.-This study evaluated post-nesting, pre-hunting season densities of eastern
white-winged (Zenaida asiatica asiatica) and mourning doves (Z macrourd) based on
morning versus evening distance sampling in south Texas. Probability of detection
curves did not differ by diel period within species. White-winged dove density estimates
were significantly greater in the evening compared to the morning (0.97 doves/ha, 95%
Cl: 0.65-1.44 morning and 2.09 doves/ha, 95% Cl: 1.53-2.86 evening). Morning dove
densities did not differ by diel period. Analysis of the raw data suggests the disparity in
white-winged dove density estimates is most likely attributable to pre-sunrise activity by
white-winged doves causing lower morning counts.

Historically, the lower Rio Grande Valley (LRGV) of Texas was
the only area in Texas where eastern white- winged doves {Zenaida
asiatica asiatica) and mourning doves (Z. macroura) exhibited
sympatry. Although mourning doves are generally cosmopolitan in
Texas (Haskett & Sayre 1993), the breeding range of eastern white¬
winged doves was originally restricted to the LRGV at the southern
tip of the state (Schwertner et al. 2002; Small et al. 2006).
However, beginning about 1920, industrialized farming and
municipal development led to an estimated overall loss of white¬
winged dove breeding habitat exceeding 90% (Purdy & Tomlinson
1991; Tremblay et al. 2005).

Following this habitat loss, and possibly associated with it,
white-winged doves began expanding northward, forming urban
populations. This range expansion continued, peaking sometime in
the late 1970s to present (Small et al. 2006). As of 1990, more
breeding white-winged doves occurred outside the LRGV than
within it (George et al. 1990). During this same period, mourning

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dove distribution remained essentially unchanged in Texas (Otis et
al 2008).

Because both dove species are migratory game birds, they are
monitored annually. Mourning doves are monitored as part of
national call-count and breeding bird surveys. The Texas Parks and
Wildlife Department (TPWD) has monitored white- winged doves
in the LRGV since about 1949 using an auditory-count index. This
auditory-count index involves counting calling white-winged doves
and using a conversion table to estimate the breeding-pair density
(Rappole & Waggerman 1986). However, how and when the
conversion table was developed is unknown; rendering the results
spurious, except to establish trends (Sepulveda et al. 2006).

To remedy this, and following pilot studies in 2004 and 2005,
TPWD implemented distance sampling to estimate the size of urban
white-winged dove populations statewide. However, the remaining
traditional rural, nesting habitat in south Texas is still occupied by
white-winged doves, but monitoring of these populations has been
suspended by TPWD. Distance sampling of urban white-winged
dove populations in Texas is conducted only in the early morning,
when it is believed detection rates are highest. Yet Small (2006)
found evenings were just as effective as mornings in sampling
white-winged dove populations in Mason, (Mason County) Texas.

Monitoring of urban white- winged dove populations in Texas is
conducted annually between 15 May and 15 June. Therefore, these
densities effectively estimate potentially breeding adults. To
estimate individuals available for harvest during the hunting season,
some measure of productivity is needed in addition to the number
of potentially breeding adults.

The objectives of this study were to determine: (1) if using point
transects in distance sampling are effective for simultaneously
estimating white-winged and mourning dove densities in native
brush of the LRGV during the post-breeding/pre-hunting season;
and (2) if morning and evening distance sampling yielded the same

SMALL ET AL.

297

density estimates for either white-winged or mourning doves or
both.

The reasoning for these objectives was to evaluate whether using
the covariate “species” in the multiple covariate distance sampling
(MCDS) engine of Program DISTANCE (Thomas et al. 2006)
would provide similar probability of detection curves from a single
model for each diel sampling period by species. Also, if the
probability of detection curves proved to be similar, as was
anticipated, would density estimates derived for each diel period be
similar; thus, indicating populations sampled in mornings versus
evenings were sufficiently closed to produce reliable population
estimates.

Methods

Study area -This study was conducted at the Anacua Unit of the
Las Palomas Wildlife Management Area (Lat 26°03T7.2", Long
97°50'46.15") between Santa Maria, Texas and the U.S. -Mexico
border (Cameron County). The Anacua Unit consists of 92.1 ha of
riparian and upland native brush habitat interspersed with three
fallow fields from north to south. Vegetation is dominated by
Texas ebony (Pithecellobium ebano), anacua (Ehretia anacua),
retama {Parkinsonia aculeate), and mesquite {Prosopis glandulosa)
(Jahrsdoerfer & Leslie 1988). It is bordered by agricultural fields
separated by dirt roads and bisected from east to west by a flood
control levee.

Assumptions of distance sampling-Voini transect distance
sampling requires that three primary assumptions be met to obtain
unbiased density estimates (Buckland et al. 2001). These
assumptions are: (1) objects on the point are detected with certainty
(i.e., g(0) =1); (2) objects are detected at their initial location; and
(3) measurements of distances to objects are exact. Protocol for
this study emphasized visually scanning each point upon approach
and then scanning outward from the point. Consequently,
individuals on the point did not go undetected. In instances when a
dove was observed on or near the point but moved in response to

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the approach of the observer, the distance from the point to the
location of the dove prior to movement was recorded. All points
were clearly marked and visible to the observer from a distance
sufficient to determine the position of doves prior to movement in
response to observer approach. Thus, assumption one was not
violated.

For assumption two, movements prior to or during the
observation period were random with respect to the observer and
assumption two was considered met (Tumock & Quinn 1991;
Buckland & Tumock 1992; Buckland et al. 2001). Lastly,
assumption three, requiring accurate measurement of distances to
detected doves, was not violated. All observers used laser range
finders which recorded distances to the nearest meter. No guesses
of distance were made for any observations.

Sampling protocol -A satellite photograph of the study area was
imported into ArcGIS version 9.2 (Environmental Systems
Research Institute, Inc., Redlands, CA, USA). A polyline layer was
drawn along dirt roads surrounding the Anacua Unit, the outside
edge of fallow fields, and along the levee traversing the area. Five
sets of 20 points (100 total points) were randomly generated along
the polyline. A random number generator was then used to shuffle
each of the five sets of 20 points.

Two observers trained in distance sampling of doves conducted
two sets of point transects from 31 My-4 August and 10- 14 August
2008. Each sampling period consisted of the 100 randomly chosen
points. Because an entire sampling period of 100 points could not
be completely sampled within a morning or evening period (2 h),
each transect was conducted on five consecutive days (20
points/day). Previous experience indicated that 20 points could be
sampled within < 2 h. Because of heavy precipitation during the
second sampling period on the evening of 13 August, only 4 points
could be accessed per observer. As a result, the first survey
consisted of 100 points and the second survey consisted of 88
points.

SMALL ET AL.

299

On each day, the 20-point subsets were divided between the two
observers, with each observer surveying the same points in the
morning and evening. One observer surveyed the first 10 points,
while the second observer simultaneously surveyed the remaining
10 points. The sequence of the points was randomized each day to
avoid points in close proximity being sampled consecutively. The
same 100 points were used for both 5-day sampling periods but
observers switched points between transects; i.e., observer one for
transect one became observer two for transect two and vice versa.
Consequently, neither observer visited the same points while
conducting the second sampling period that they surveyed during
the first sampling period.

Observers conducted surveys concurrently beginning about 15
min after official sunrise and completing surveys no later than 2 h
post-sunrise. Each day, transect surveys were duplicated in the
evenings beginning no earlier than 2 h pre-sunset and completed no
later than 15 min pre-sunset. Transect points were visited in the
same sequence in the evening as in the morning. Survey protocol
followed Schwertner & Johnson (2005). Each point was visited for
a 2-min period and distances to all white-winged and mourning
doves observed were recorded to the nearest meter using a
Bushnell™ Yardage Pro Legend laser range-finder (Bushnell, Inc.,
Overland Park, KS, USA). For this study, only visual detections
were used; auditory detections were not recorded.

Data analysis were analyzed in program DISTANCE
(Thomas et al. 2006) using the MCDS engine because two
covariates were of interest; time of day (morning and evening) at
the stratum level and species (white-winged dove and mourning
dove) as an added covariate. The two trials were pooled to ensure
adequate numbers of observations were available for analysis at the
covariate levels (Marquez & Buckland 2004; Marquez et al. 2007).
Data were combined for both observers because Program
DISTANCE uses detection models that are pooling robust and
because the covariates of interest were time of day (morning versus
evening) and species (white- winged dove versus mourning dove).

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 4, 2009

The two sample periods were pooled into a single point transect to
maximize numbers of observations available for analysis at the
covariate levels (Marquez & Buckland 2004; Marquez et al. 2007).
Although the conventional distance sampling (CDS) engine allows
for a single covariate (at the stratum level), separate runs would
have to be made for the additional covariate (Buckland et al. 2001).
Probability of detection fiinction curves within species was
compared using a Kolmogorov-Smimov test.

The advantage of the MCDS engine over the CDS engine for
this study was that a single truncation point and a single detection
model could be used. This affords greater precision in density
estimates via smaller 95% confidence intervals (CIs), In addition,
use of the MCDS engine provides output data that allows
calculation of a scaling parameter (5) which is then used to plot the
probability of detection curves at the co variate level. Evaluation of
these curves in relation to each other allows assessment of the
validity of the covariates (Marquez et al. 2007).

Initially, five candidate models from the MCDS engine were run
using all data (no truncation) with no covariates and restricted to no
more than two adjustment terms. The Akaike Information Criterion
(AIC) was used to select the most parsimonious model for
estimating population density (Burnham & Anderson 2003), Using
the most parsimonious model, a likely truncation point was chosen
based on diagnostic output from Program DISTANCE. Once
identified, data were run again with a single covariate (species) and
using various truncation points around the original choice until the
data satisfactorily fit the probability of detection curve both visually
and statistically; using the Komolgorov-Smimov test P-value
calculated by DISTANCE.

Output from Program DISTANCE, using the most parsimonious
model (that with the lowest AIC value), was then used to determine
scale parameters (s) which were used to model probability of
detection curves for each species by morning and evening.
Program DISTANCE was then run to calculate population density

SMALL ET AL.

301

estimates for each species by time of day with corresponding 95%
confidence intervals.

As a further evaluation of morning versus evening sampling, raw
data obtained from each sampling period were examined. Paired t-
tests were used to compare daily differences in number of
individuals, number of observations (clusters), and mean cluster
size between morning and evening sampling for each species.
Also, scatter plots of number of doves seen each minute post¬
sunrise and pre-sunset were generated for each species. If any plot
showed a pattern of increasing values at either end of the plot, these
data were grouped into 10 min intervals and regressed against
number of individuals seen.

Results

From the pool of 100 randomly placed points, 188 point surveys
were conducted. A total of 105 and 108 individual mourning doves
were recorded across 78 and 84 observations made during morning
and evening sampling, respectively; this compared to 815 and 1,346
individual white- winged doves across 207 and 357 observations,
respectively.

The most parsimonious model selected by Program DISTANCE
was a half-normal with a cosine key function and one expansion
term of order two {D = 0.03, P = 0.74) with data truncated at 160
m. Probability of detection functions constructed from these data
were extremely similar between species and nearly identical within
species for morning and evening sample periods (D = 0.12, P =
0.16 for both species. Fig. 1). However, the density estimate was
significantly higher for evening sampling than for morning
sampling for white-winged doves (0.97 doves/ha, 95% Cl: 0.65-
1.44 morning and 2.09 doves/ha, 95% Cl: 1.53-2.86 evening. Fig.
2), Estimates did not differ for mourning doves (0.38 doves/ha,
95% Cl: 0.24-0.59 morning and 0.53 doves/ha, 95% Cl: 0.35-0.80
evening. Fig. 2).

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 4, 2009

Distance (m)

-WWD-M

-WWD-E

-MODaM

-MODO-E

Figure 1. Probability of detection curves for (a) white- winged doves sampled in the
morning (WWD-M) and the evening (WWD-E) and for (b) mourning doves sampled
in the morning (MODO-M) and the evening (MODO-E).

For white- winged doves, total number of doves observed and
total number of observations (clusters) were higher during evening
sampling = -2.62, P = 0.03 and = -5.36, P < 0.01, respective¬
ly). Mean white- winged dove cluster size per day did not differ
between morning and evening sampling (fg = 0.04, P = 0.97). For
mourning doves, total number of doves observed, total number of

SMALL ET AL.

303

3.0

2.5

2.0 ■

p 1.5 -
o
Q

1.0 -

0.5 -

0.0

Sample

Figure 2. Density estimates and 95% confidence intervals for white-winged and
mourning doves sampled in mornings and evenings.

observations (clusters), and mean cluster size per day did not differ
between morning and evening sampling (tg = -0.23, P = 0.82, /g = -
0.72, P = 0.49, and tg = 0.24, P = 0.81, respectively). Evaluation of
scatter plots showed only white-winged doves sampled in the
morning had substantially more individuals seen closer to sunrise.
Regression of number of white- winged doves over 10 min intervals
(the last interval was only 8 min) post-sunrise showed a strongly

significant negative relationship (r = 0.81, P < 0.01, Fig. 3).

These results are consistent with counts of daily data. White¬
winged doves had higher counts during evenings than mornings for
number of individuals (7 of 10 days) and number of clusters
observed (9 of 10 days, 1 equal). Mean daily cluster size differ-

304 THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 4, 2009

Minutes post-siinrise

Figure 3. (a) Scatter plot of white- winged doves seen by minute post-surrise and (b)
regression of white-winged doves seen over 10 min intervals.

ences were equal, each diel period having 5 days with higher
values. For mourning doves, number of individuals and number of

SMALL ET AL.

305

clusters was similar in the mornings and evenings with evenings
having higher values on 4 of 10 days, 2 equal and 5 of 10 days,
respectively. Evening counts of mean cluster size were higher on 6
of 10 days (Fig. 4).

Discussion

This study had a narrow time window for completion. Because
doves commonly nest more than once (often late in the breeding
season), initiate fall feeding flights, and exhibit some variability in
migration, there was only about a four-week, post-nesting, pre¬
hunting season period of time to complete this study. Also, on 23
July, category III Hurricane Dolly hit the Texas coast about 90 km
east of the study site. Doves may have scattered, or left the area
entirely, as a result of effects of Hurricane Dolly, resulting in lower
than normal density estimates

Significantly different density estimates by diel period for white¬
winged doves, specifically, a higher evening density estimate
compared to morning, suggest morning and evening populations
were not comprised of the same individuals; the population was not
closed. Although some of the individuals present may have
remained constant, the proportion is unclear. Further, these results
do not preclude the use of morning and evening distance sampling
for determining white-winged dove density during other times of
the year.

The most plausible explanation for this discrepancy is that
white-winged doves began leaving roosts prior to or at sunrise and
were consequently unavailable for counting because of lack of
available daylight. This conclusion is supported by the number of
white-winged doves seen relative to time (Fig. 3). White-winged
doves are considered diurnal (Schwertner et al. 2002); thus,
individuals moving between roosts at night seems highly
improbable. However, they do exhibit some degree of crepuscular
activity (Cottam & Trefethen 1968). It is assumed that, under this
scenario, white-winged doves in the morning do not leave roosts at

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THE TEXAS JOURNAL OF SCIENCE, VOL. 61, NO. 4, 2009

. ♦

vwvnw

vwvnE

-■ A-

- MODaW

- ■ MODaE

VW\^M

vwvnE

» MODaM

• ■ - JK-

• - MODaE

Figure 4. Daily values for (a) number of individuals, (b) number of observations, and (c)
mean cluster size for white-winged doves sampled in die morning (WWD-Ld) and the
evening (WWD-E) and for mourning doves sampled in the morning (MODO-M) and
the evening (MODO-E).

the same time, but do so in a streaming fashion, beginning earlier
than 15 min post-sunrise. If so, this count bias would account for

SMALL ET AL.

307

the differences in the number of white-winged doves seen in
morning and evening samples. For evenings, white- winged doves
returning to roosts may be more time-structured (i.e., occurs within
a shorter time-frame and more consistently on a daily basis). A
study of morning distance sampling with a starting time of 15 min
pre-sunrise may alleviate differences found in this study between
diel periods provided visibility is sufficient to count doves.

White-winged doves monitored by radio-telemetry in urban
areas exhibited high site fidelity and structured home ranges
centered on their nest (Small et al. 2005), Because this study was
conducted during the post-breeding period and pre-hunting season,
it is conceivable that white- winged doves do not retain the same site
fidelity associated with nesting. Generally, post-nesting white¬
winged doves become gregarious in roosting, aggregate into large
feeding flocks, and forage together to build energy reserves for the
fall migration (Cottam & Trefethen 1968; Schwertner et al. 2002).
Roosting in proximity to changing available food and water
resources may contribute to a lack of site fidelity in white-winged
doves and may also, at least partially, explain the high variation in
daily morning and evening counts (Fig. 3).

Mourning doves, conversely, exhibited nearly identical density
estimates for morning and evening surveys (Fig. 2) and very low
daily variation in number of individuals, number of observations,
and mean cluster size (Fig. 3). While this result does not
definitively prove that the mourning dove population being sampled
was closed, it provides strong support for this hypothesis. It also
suggests that, pending further research, mourning doves can be
sampled as reliably in the evening as in the morning.

This study indicates distinctly different behavior in post-nesting
white-winged and mourning doves. Should fiirther study indicate
these differences to be consistent over time, it may be beneficial to
monitor the two species differently. Mourning doves are currently
monitored annually as part of the federal mourning dove call-count

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survey conducted in late May and early June (Dolton 1993). The
purpose of the call-count survey is to monitor relative abundance
and population trends which are used to determine hunting seasons.
However, this method is retroactive as opposed to adaptive.
Analysis of previous trends elicits a response in future regulations
as opposed to regulations influencing trends by using adaptive
management models. Also, productivity, which is highly variable
in mourning doves (Hayslette et al. 1996; 2000), is not measured.
Only estimates of breeding adults are quantified, which may not be
reflective of numbers of individuals available for harvest in a given
year.

Based on this study, and pending future research, several
recommendations can be made. First, the monitoring of urban
white-winged doves using distance sampling should continue to
occur at the onset of the breeding season in mid-May. However,
white-winged dove monitoring should be expanded to include rural
nesting populations and a nesting productivity monitoring initiative
should be implemented to more accurately estimate doves available
for harvest.

Secondly, further evaluation of post-nesting distance sampling
of mourning doves should be implemented. If late summer
sampling of mourning doves proves to provide accurate estimates
of individuals available for harvest, consideration should be given
to shifting the monitoring of mourning doves from late spring to
late fall. Both of these suggestions have the potential for providing
more precision in estimating population trends for both these game
species while improving efficiency of human resources required for
conducting these surveys.

Acknowledgments

Thanks to J. Veech for reviewing earlier drafts of this
manuscript. J. Timmons was instrumental in data collection. This
study was funded by the Texas Parks and Wildlife Department
white-winged dove stamp fund.

SMALL ET AL.

309

Literature Cited

Baskett, T. S. & M. W. Sayre. 1993. Characteristics and importance. Pp. 1-6, in
Ecology and Management of the Mourning Dove (T. S. Baskett, M. W. Sayre, R. E.
Tomlinson, & R. E. Mirarchi, eds.), Stackpole Books, Harrisburg, 608 pp.

Buckland, S. T., D. R. Anderson, K. P. Burnham, J. L. Laake, D. L. Borchers & L.
Thomas. 2001. Introduction to Distance Sampling: Estimating Abundance of
Animal Populations. Oxford University Press, New York, 448 pp.

Buckland, S. T. & Tumock, B. J. 1992. A robust line transect method. Biometrics
48(3):901-909.

Burnham, K. P. & D. Anderson. 2003. Model Selection and Multi-model Inference: A
Practical Information-theoretic Approach. Springer, 488 pp.

Cottam, C. & J. B. Trefethen. 1968. Whitewings: The Life History, Status, and
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Dolton, D. D. 1993. The call-count survey: historic development and current
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TEXAS J. SCI. 61(4), NOVEMBER, 2009

311

GENERAL NOTES

PREVALENCE OF HEMATOZOAN PARASITES (APICOMPLEXA)
IN SOME COMMON PASSERINE BIRDS (PASSERIFORMES)
FROM EAST-CENTRAL OKLAHOMA

Michael D. Bay and Kenneth D. Andrews

Department of Biology, East Central University
Ada, Oklahoma 74820

Blood parasites (hematozoa) occur in the plasma {Trypanosoma)
and within the blood cells {Plasmodium^ Leucocytozoon and
Haemoproteus) of a number of avian species. Most of these
infections are likely benign in wild birds (Bennett et al 1988;
Weatherhead & Bennett 1991; 1992), and occasionally pathogenic
during stressful times (Atkinson & Van Riper 1991; Weatherhead &
Bennett 1991). Furthermore, parasites may have some influence on
sexual selection (Hamilton & Zuk 1982; Zuk 1991); however, this
has yet to be resolved (Weatherhead 1 990; Weatherhead et al 1991;
Seutin 1994; Scheuerlein & Ricklefs 2004).

The distribution of blood parasites has been studied in many
regions (e.g., Bernard & Bair 1 986 in Vermont; Rodriguez & Matta
2001 in Colombia), while other areas have been less well studied or
not at all. Parasitism of Oklahoma birds has been investigated in
only one previous instance (Janovy 1963), and few data exist
concerning the levels of infection or types of parasites involved. In
this study, the incidence of infection is detailed for four common
passeriform species, and also the types of blood parasites involved
and their intensities.

Methods and Materials

This study was conducted in Pontotoc, Murray and Okfiiskee
counties of Oklahoma from May 2002 through August 2005. Since
most hemosporidian infections are patent by early to mid-summer.

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this time frame is optimal in sampling infected hosts (Valkiunas
2001). All birds were captured using 30- and 38- mm mesh mist
nets, and banded with U.S. Bird Banding Laboratory aluminum
bands. All captured individuals were sexed and aged where
possible, as well as measured and weighed. Blood samples were
obtained from toenail clippings and smeared on slides. These
samples were air dried, fixed with 100% methanol and then stained
with Giemsa (Bennett 1970). All slides were examined at 400-
lOOOX. Parasites were quantified by counting the number per
10,000 blood cells to obtain an estimate of parasite intensity
(Godfrey et al. 1987). All blood smears were submitted as voucher
specimens to the U.S. National Parasite Collection (accession
numbers 100814- 100835), Beltsville Agricultural Research Center,
10300 Baltimore Avenue, Beltsville, Maryland 20705.

Results and Discussion

A total of 175 birds representing four species (49 Carolina
Chickadees, Parus carolinensis, 55 House Sparrows, Passer
domesticus, 37 Northern Cardinals, Cardinalis cardinalis and 34
House Finches, Carpodacus mexicanus) were examined. No
significant difference (x, df= 3, P>0.05) between years in the
prevalence of infection, was noted, permitting data to be pooled
across years for each species. Overall prevalence of blood parasite
infections was 12.5% for the total sample. Four blood parasites
(identified to genus) were found ranging from a prevalence of 1.8 to
13.5% for the host species (Table 1), with Haemoproteus sp. being
the most common (54.5% of those infected). Though the
prevalence of parasites was low in the species sampled, they were
significantly different (x^=17.3, df=3, P<0.001), possibly indicating
some degree of susceptibility to vectors and/or differences in host
resistance to infection. Although prevalence tends to vary from
region to region, and from species to species, most studies report a
low percentage of infection in passerine birds. For instance,
Williams & Bennett (1978) found that 10.5% of Northern Cardinals

TEXAS J. SCI. 61(4), NOVEMBER, 2009

313

Table L Prevalence (number infected / number examined, %) of blood parasites in four
common bird species from east-central Oklahoma. HP= Haemoproteus; PL=
Plasmodium', HG= Haemogregarina', LK= Leukocytozoon.

Parasite

Host

Cardinalis cardinalis (Northern Cardinal)

13.5%

5.4%

2.7%

Passer domesticus (House Sparrow)

9.1%

1.8%

Carpodacus mexicanus (House Finch)

2.9%

Parus caroUnensis (Carolina Chickadee)

2.0%

(«=180) and 15,2% of House Sparrows {n=\ll), in New Jersey were
infected, compared to 21% of cardinals and 12.7% of House Sparrows
in this study. In contrast, Greiner et al (1975) found a higher
percentage of cardinals were infected (42.9%), though prevalence
findings for the House Finch and Carolina Chickadee were similar to
this study (14.8% and 6.8% respectively). Studies by Bernard & Bair
( 1 986), Stabler & Kitzmiller (1970) and Al-Dabagh ( 1 964), also report
a low infection prevalence in the House Sparrow, comparable to this
study. The only other study conducted in Oklahoma (Janovy 1963)
reported that only 5.8% of passerine birds («=102) were infected,
although only the House Sparrow was common to this study.

Haemoproteus was the most frequent occurring parasite in this
study as well as in the studies of passeriform birds in other regions
(e.g., Williams & Bennett 1978 in New Jersey and Maryland; Bennett
et al. 1991 in Mexico; Scheuerlein & Ricklefs 2004 in Europe), while
Leucocytozoon was most prevalent in the studies of Stabler &
Kitzmiller (1970) in Colorado, Bernard & Bair (1986) in Vermont, and
Merino et al. (1997) in Spain. Janovy’s (1963) study in Oklahoma
reports Haemoproteus was the most frequent parasite encountered,
though most were from a non-passeriform bird, the Mourning Dove
{Zenaidura macroura).

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The level of intensity (parasites per 10,000 blood cells) was low for
all infected individuals, so it was not possible to determine if any
pattern existed for each host species relative to the sampling period.
Of the four parasites, Plasmodium spp. had the highest mean intensity
in the Northern Cardinal (9.9%) and the House Sparrow (1.2%),
followed by Haemoproteus spp (1.5%) and Leukocytozoon spp.
(0.84%) in the House Finch. Haemogregarina spp. was detected in
three of the four bird species sampled (except the Northern Cardinal)
but had the lowest intensity (0.03% to 0.28).

The interpretation of parasitemia levels can be difficult, since they
tend to vary considerably both temporally and spatially and may often
be associated with considerable error in comparison to prevalence
(McCurdy et al. 1998). Although the intensity levels of this study
were low and too infrequent for analysis, it is important to report them
here for comparison to future studies. Also, since parasites might be
metabolically costly to birds, the body weights of infected individuals
were compared to those that were not infected in three of the four host
species (excluding Carolina Chickadee because of low infection).
Since there are no gender difference in the body weights of males and
females, the weights were combined in the analysis. For the Northern
Cardinal and House Finch there was no significant difference in the
mean body weights of those infected compared to those that were not
(t=0.07, P>0.05 and t=1.47, P>0.05, respectively), however, in the
House Sparrow, infected individuals had a higher mean body weight
(t=2.71,P<0.01) (Table 2).

Assessing the effects of blood parasites on wild hosts is a difficult
process and there is even some debate as to whether these parasites are
pathogenic or not (Atkinson & Van Riper 1991). It is possible that
some species are more resistant than others to a buildup of high levels
of parasitemia. Some studies show that high parasite levels could
affect working adults (feeding nestlings) (Merino et al. 2000), or
possibly early aged individuals (hatchlings) or first year adults
(Weatherhead & Bennett 1 992). In this study, the level of parasitemia
was relatively low, and any pathogenic effects might be minimal. This

TEXAS J. SCI. 61(4), NOVEMBER, 2009

315

Table 2. Body weights (X ± SE) of bird species, comparing infected individuals to those
not infected from east-central, Oklahoma 2002-2005.

Host Species

Infected

Uninfected

Northern Cardinal

40.9 ± 6.4%

41.4 ± 1.4%

.07

>0.05

House Sparrow

30.4 ± 6.9%

27.6 ± 0.72%

2.71

<0.01

House Finch

20.3 ± 5.8%

20.7 ± 0.68%

1.47

>0.05

may explain why there was no difference in body weights between
infected and uninfected individuals in three of the four species,
although it is interesting that infected House Sparrows had a higher
mean body weight in comparison to uninfected individuals. While
some studies report little if any difference in body condition of
infected and uninfected birds (e.g., Weatherhead & Bennett 1992),
some do report a higher mean body weight in infected individuals as
Bennett et al. (1988) found in the Black and White Warbler {Mniotilta
varia) and Fox Sparrow {Passerella iliaca). Bennett et al. (1988)
suggests that it is possible that blood parasites simply do not cause a
loss of body mass, even in individuals with a high parasitemia. Also,
given the small body mass of passerine species, changes due to
parasite infection may be too small to distinguish from other natural
occurring effects on body weight, such as foraging and breeding
activities (Bennett et al. 1988).

Acknowledgments

Many thanks to Rick Davis for helping mist net birds and Tiffany
Howell for inspecting slides and counting infected blood cells. This
research was aided by a Federal Bird Banding Permit (U.S.
Department of the Interior, Bird Banding Laboratory, Laurel,
Maryland) issued to the senior author.

Literature Cited

Al-Dabagh, M. A. 1964. The incidence of blood parasites in wild and domestic birds of
Columbus, Ohio. Am. Midi. Nat., 72:148-150.

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Atkinson, C.T. & C. Van Riper IIL 1991. Pathogenicity and epizootiology of avian
haematozoa: Plasmodium, Leucocytozoon, and Haemoproteus. Pp 19-48 in Bird-
parasite interactions (J.E. Loye and M. Zuk, eds), Oxford Univ. Press, 406pp.

Bennett, G. F. 1970. Simple techniques for making avian blood smears. Can. J. Zook,
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Bennett, G. F., J. R. Caines & M. A. Bishop. 1988. Influence of blood parasites on the
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northeastern Mexico. J. ParasitoL, 77:38-41.

Bernard, W. H. & R. D. Bair 1986. Prevalence of avian hematozoa in central Vermont.
J. Wildl. Dis., 22:365-374.

Godfrey, R. D., Fedynich, A. M. & D. B. Pence. 1987. Quantification of hematozoa in
blood smears. J. Wildl. Dis., 23:558-565.

Greiner, E. C., Bennett, G. F. E. M. White & R. F. Coombs. 1975. Distribution of the
avian hematozoa of North America. Can J. Zook, 53:1762-1787.

Hamilton, W. D. & M. Zuk. 1982. Heritable true fitness and bright birds: a role for
parasites. Science, 218:384-387.

Janovy, J. 1 963 . A preliminary survey of blood parasites of Oklahoma birds. Proc. Okla.
Acad. Sck, 43:59-61.

McCurdy, D.G., D.Shtler, A. Mullie & M.R. Forbes. 1998. Sex- biased parasitsm of
avian hosts: relations to blood parasite taxon and mating system. Oikos, 82:303-3 12.

Merino, S., Potti, J. & J. A. Fargallo. 1997. Blood parasites of passerine birds from
central Spain. J. Wildl. Dis., 33:638-641.

Merino, S., Morreno, J., J. J. Sanz & E. Arriero. 2000. Are avian blood parasites
pathogenic in the wild? A medication experiment in blue tits {Parus caeruleus).
Proc. R. Soc. Lond., 267:2507-2510.

Rodriguez, O. A. & N. E. Matta. 2001 . Blood parasites in some birds from eastern plains
of Colombia. Mem. Inst. Oswaldo Cruz, 96:1 173-1 176.

Scheuerlein, A. & R. E. Ricklefs. 2004. Prevalence of blood parasites in European
passeriform birds. Proc. R. Soc. Lond., B 271.

Seutin, G. 1994. Plumage redness in redpoll finches does not affect hemoparasitic
infection. Oikos, 70:280-286.

Stabler, R. M. & N. Kitzmiller. 1970. Hematozoa from Colorado birds. III.
Passeriformes. J. Parasit., 56:12-16.

Valkiunas, G. 2001. Blood parasites of birds: some obstacles in their use in ecological
and evolutionary biology studies. Avian Ecok Behav., 7:87-100.

Weatherhead, P. J. 1 990. Secondary sexual traits, parasites and polygyny in Red winged
Blackbirds. Behav. Ecok, 1:125-130.

Weatherhead, P. J. & G. F. Bennett. 1991. Ecology of Red winged Blackbird parasitism
by haematozoa. Can. J. Zook, 69:2352-2359.

Weatherhead, P. J. Bennett, G. F., & D. Shutler. 1991 . Sexual selection and parasites in
wood warblers. Auk, 108:147-152.

Weatherhead, P. J. & G. F. Bennett. 1992. Ecology of parasitism of Brown headed
Cowbirds by haematozoa. Can J. Zook, 70:1-7.

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Williams, N. A. & G. F. Bennett. 1978. Hematozoa of some birds of New Jersey and
Maryland. Can. J. Zool., 56:596-603.

Zuk, M. 1991. Parasites and bright birds: new data and a new prediction. Pp 3 1 7-327,
in Bird-parasite interactions (J. E. Loye and M. Zuk, eds), Oxford Univ. Press,
406pp.

MDB at: mbay@ecok.edu

* * * * *

NOTES ON REPRODUCTION OF THE KNOB-SCALED LIZARD,
XENOSAURUS GRANDIS (SQUAMATA: XENOSAURIDAE),
FROM VERACRUZ, MEXICO

Stephen R. Goldberg

Department of Biology, Whittier College, PO Box 634
Whittier, California 90608

The knob-scaled lizard, Xenosaurus grandis is distributed as
disjunct populations from west central Veracruz southward to
Guerrero, Oaxaca and Chiapas, Mexico to Alta Verapaz, Guatemala
(Ballinger et al. 2000a). It is viviparous (Ballinger et al. 2000b).
Information on the reproduction of A. grandis is in Fritts (1966);
Alvarez del Toro (1982); Ballinger et al. (2000b); Smith et al.
(2000). The purpose of this note is to add information on the
reproductive biology of X. grandis from the first histological
examination of gonadal material from this species.

Eleven males (mean snout-vent length, SVL = 104.5 mm ± 12.5
SD, range = 83-120 mm); 27 females (mean SVL = 108.7 mm ±
12.3 SD, range = 87-133 mm); 3 juveniles (mean SVL 69.3 mm ±
8.1 SD, range = 62-78 mm) and one neonate (SVL = 42 mm) X.
grandis collected from 1969 to 1983 at Cuautlapan (18°52’12”N,
97°r48”W), Veracruz, Mexico were examined from the
herpetology collection of the Natural History Museum of Los

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Angeles County (LACM), Los Angeles, California.

The left testis, epididymis and left ovary were removed from
males and females, respectively. Histology slides were prepared by
conventional methods and stained by Harris hematoxylin and eosin
(Presnell & Schreibman 1997). Oviductal eggs or enlarged ovarian
follicles (> 4 mm length) were counted. Histology slides are
deposited in LACM. An unpaired ^test was used to compare male
and female mean body sizes, and linear regression analyses
examined the relationship between female body size and clutch
number (Instat vers. 3.0b, Graphpad Software, San Diego, CA).

The following Xenosaurus grandis specimens from Veracruz,
Mexico were examined: LACM 5927, 10981, 75420, 75688, 75690,
75691,75694, 104942, 120049-120057, 120059, 120060, 120064,
120065, 120070, 120071, 120075-120077, 120080-120084,
120086, 120087, 121537, 135539, 135541, 135544, 135545,
135547, 136334-136336.

There was no significant size difference between mean male and
female SVLs (unpaired t test, P = 0.34). Monthly stages in the
testicular cycle are in Table 1. Three stages were present: (1)
Regression: germinal epithelium of the seminiferous tubules is 1-2
layers thick and is composed mainly of spermatogonia with
occasional Sertoli cells; (2) Recrudescence: there is an increase of
cellularity in the seminiferous tubules due to a proliferation of germ
cells; primary spermatocytes predominate. In late stage
recrudescence, secondary spermatocytes may be abundant and
occasional spermatids are noted; (3) Spermiogenesis: seminiferous
tubules are lined by clusters of spermatozoa and groups of
metamorphosing spermatids are present.

TEXAS J. SCL 61(4), NOVEMBER, 2009

319

Table 1. Monthly stages in testicular cycle of 11 Xenosaurus grandis from Veracruz,
Mexico.

Month

Regression

Recrudescence

Spermiogenesis

January

May

August

September

Regressed testes were present in January (Table 1).
Recrudescence (= renewal) was in progress in the single male X.
grandis from May. Testes from August and September were
undergoing spermiogenesis (= sperm formation). The smallest
reproductively active males (spermiogenesis in progress) both
measured 83 mm SVL and were from August (LACM 120080,
121537). One apparently sub-adult male from March with a
regressed testis (LACM 120065) measured 78 mm SVL. Two other
subadults, (LACM 120054) SVL = 68 mm from January and
(LACM 120087) SVL= 62 mm from July were not sexed.

Smith et al. (2000) reported that grandis males from Veracruz,
Mexico mate in fall at which time testes are enlarged. Findings
from this study confirm a late-summer fall period of sperm
production for X. grandis, consistent with other fall/winter active
viviparous lizards (Goldberg 1971; 2002; Smith et al. 2000).

Monthly stages in the ovarian cycle are in Table 2. Five stages
were noted: (1) Quiescent: no yolk deposition; (2) Yolk deposition:
early vitellogenesis in progress with basophilic yolk granules; (3)
Enlarged follicles > 5 mm; yolk filled follicles are approaching
ovulation; (4) Oviductal eggs: ovulation has occurred; (5) Embryos
are developing within the oviducts.

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Table 2. Monthly stages in the ovarian cycle of 27 Xenosaurus grandis from Veracruz,
Mexico.

Month

Quiescent

Yolk

deposition

Enlarged
follicles >5inni

Oviductal

eggs

Embryos

January

April

May

June

July

August

October

* Reproductive organs were damaged making it impossible to count the number of
embryos.

Embryos were observed in January, June and July suggesting
parturition occurs in late summer. One neonate that measured 42
mm SVL was ‘‘prematurely” bom dead in captivity during May
(LACM 135547). The presence of reproductively quiescent females
during all months sampled indicates X. grandis do not produce
young each year. Histological examination revealed none of these
quiescent ovaries exhibited any trace of yolk deposition. Moreover,
it is doubtful if two May A grandis females with enlarged follicles
> 5 mm that had not yet ovulated would have produced young that
summer, suggesting it may take some females more than two years
to complete gestation. Ballinger et al. (2000b) reported X grandis
females reproduce every other year. The smallest reproductively
active X. grandis female (developing embryos) measured 90 mm
SVL (LACM 10981). The correlation between female body size
and litter size was not significant {P = 0.77). Mean clutch size for
10 gravid X. grandis was 5.7 ± 0.95 SD, range: 4-7. This mean
litter size is larger than the 3.2 ± 0.2 SE reported for X. grandis

TEXAS J. SCI. 61(4), NOVEMBER, 2009

321

agrenon from Oaxaca, Mexico (Lemos-Espinal et al. 2003) where
the two smallest reproductively active females (embryos present)
measured 97 mm SVL. It is however close to the mean litter size of
5.1 ± 0.2 SE, range 2-8 forX. grandis from Veracruz (Ballinger et
al. 2000b). This suggests geographic variation occurs inX. grandis
litter sizes. Fritts (1966) reported young ofX. grandis were bom in
July which is consistent with Ballinger et al. (2000b) who reported
parturition from June through August.

The factors responsible for only part of a female lizard
population reproducing in a given year are unknown. It is typical
for adult females of most species to be in various stages of the
ovarian activity during the reproductive cycle in the temperate zone
(see for example Goldberg 1973; 1975) or to exhibit some
reproductive activity (often associated with moisture) throughout
the year at low elevations in the tropics (Fitch 1982). Reports of
female lizards with biennial production of young are more typical of
those living in very harsh environments (Cree & Guillette 1995;
Boretto & Ibarguengoytia 2006; Ibargiiengoytia & Casalins 2007).
In addition to X. grandis, another lizard in which only parts of the
adult female population reproduces each year is the xantusiid lizard,
Xantusia riversiana (cf. Goldberg & Bezy 1974). In contrast toX.
grandis, females of the viviparous congeners X. newmanorum and
X. platyceps reproduce each year (Ballinger et al. 2000b).
Additional study is needed to elucidate the factors responsible for
only part of the X grandis female population reproducing in a given
year.

Acknowledgments

I thank Christine Thacker (LACM) for permission to examine
specimens.

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Literature Cited

Alvarez del Toro, M. 1982. Los Reptiles de Chiapas. Tercera Edic. Publicacion del
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Ballinger, R. E., J. A. Lemos-Espinal & G. R. Smith. 2000a. Xenosaurus grandis
(Gray). Knob-scaled lizard. Catalog. Amer. Amphib. ReptiL, 713:1-4.

Ballinger, R. E., J. A. Lemos-Espinal & G.R. Smith. 2000b. Reproduction in females of
three species of crevice-dwelling lizards (genus Xenosaurus) from Mexico. Stud.
Neotrop. Fauna Environ., 35:179-183.

Boretto, J. M. & N. R. Ibarguengoytia. 2006. Asynchronous spermatogenesis and
biennial female cycle of the viviparous lizard Phymaturus antofagastensis
(Liolaemidae): reproductive responses to high altitudes and temperate climate of
Catamarca, Argentina. Amphib.-Reptil., 27:25-36.

Cree, A. & L. L. Guillette, Jr. 1995. Biennial reproduction with a fourteen-month
pregnancy in the gecko Hoplodactylus maculatus from Southern New Zealand. J.
Herpetol., 29:163-173.

Fitch, H. S. 1982. Reproductive cycles in tropical reptiles. Occas. Pap. Mus. Nat. Hist.
Univ. Kansas, 96:1-53.

Fritts, T. H. 1966. Notes on the reproduction of Xenosaurus grandis (Squamata:
Xenosauridae). Copeia, 1966:598.

Goldberg, S. R. 1971. Reproductive cycle of the ovoviviparous iguanid lizard
Sceloporus jarrovi Cope. Herpetologica, 27:123-131.

Goldberg, S. R. 1973. Ovarian cycle of the western fence lizard, Sceloporus
occidentalis. Herpetologica, 29:284-289.

Goldberg, S. R. 1975. Reproduction in the sagebrush lizard, Sceloporus graciosus.
Amer. Midi. Nat., 93:177-187.

Goldberg, S. R. 2002. Eumeces brevirostris (Short-nosed Skink). Reproduction.
Herpetol. Rev., 33:134.

Goldberg, S. R. & R. L. Bezy. 1974. Reproduction in the island night lizard, Xantusia
riversiana. Herpetologica, 30:350-360.

Ibarguengoytia, N. R. & L. M. Casalins. 2007. Reproductive biology of the
southernmost gecko Homonota darwini: convergent life-history patterns among
southern hemisphere reptiles living in harsh environments. J. Herpetol., 41 :72-80.

Lemos-Espinal, J, A., G. R. Smith & R. E. Ballinger. 2003. Ecology of Xenosaurus
grandis agrenon, a knob-scaled lizard from Oaxaca, Mexico. J. Herpetol., 37:192-
196.

Presnell, J. K. & M. P. Schreibman. 1997. Humason’s Animal Tissue Techniques, 5*^
Edit., The Johns Hopkins University Press, Baltimore, MD. 572 pp.

Smith, G. R., R. E. Ballinger & J. A. Lemos-Espinal. 2000. Male reproductive cycle of
the knob-scaled lizard, Xenosaurus grandis. Southwest. Nat., 45:356-357.

SRG at: sgoldberg@whittier.edu

TEXAS J. SCI. 61(4), NOVEMBER, 2009

323

POPULATION DYNAMICS OF
AN ESTABLISHED REPRODUCING POPULATION OF
THE INVASIVE APPLE SNAIL (POMACEA INSULARUM)
IN SUBURBAN SOUTHEAST HOUSTON, TEXAS

Colin H. Kyle, Matthew K. Trawick, James P. McDonough
and Romi L, Burks

Department of Biology, 1001 East University Avenue
Southwestern University, Georgetown, Texas 78626

Over the past 15 years in the United States, a rise in introductions
of non-native gastropod species has prompted major concern from
both the U.S. Department of Agriculture and aquatic ecologists
(Robinson 1999; Levine & Antonio 2003). Due to their large size
(i.e., mass and operculum width up to approximately 150 g and 55
mm, respectively; Youens & Burks 2008) and high rate of
reproduction (i.e., up to 4000 eggs per clutch; Barnes et al. 2008), the
freshwater gastropod family Ampullariidae contains a number of
destructive invasive species (Rawlings et al. 2007) that alter ecosystem
function and threaten native biodiversity (Carlsson et al. 2004;
Carlsson & Bronmark 2006; Boland et al. 2008; Connors et al. 2008).
Invading habitats worldwide, apple snails of the genus Pomacea now
represent an increasing environmental problem in the U.S. Multiple
introductions (Rawlings et al. 2007) of numerous species from
multiple origins (Hayes et al. 2008) complicate this problem.

Native to temperate South America, the apple snail species now
found in Texas (Karatayev et al. 2009), Pomacea insularum, possesses
a round shell with a characteristic deep groove on the whorl (i.e.,
channeled) (Howells et al. 2006). Unlike most snails, P. insularum,
and closely related P. canaliculata (another channeled species),
consume macroscopic plants rather than algae (Carlsson &
Lacoursiere 2005; Burlakova et al. 2008). Because molecular
geneticists only recently identified P. insularum as a distinct species
from its close relative, the mode of introduction, spread, and current
distribution of this newly introduced species requires more attention

324

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

from ecologists (Rawlings et al. 2007; Karatayev et al. 2009).

To address whether or not newly established populations of P.
insularum now persist, this report analyzed size structure of P.
insularum populations from 2006-2008. This study examined changes
in proportions of juveniles and adults within sample populations. An
increase in the proportion of juveniles between samples taken from
2006 to 2008 indicates changes in the size distributions of introduced
populations of P. insularum.

Live snails were collected during five trips: May 2006 (^=100),
May 2007 {n=2\\ September 2007 (n=ll\ May 2008 (/?=82) and
August 2008 {n=\16). Each trip involved two sample sites in the
Houston area: Horsepen Bay located within Armand Bayou and a
drainage ditch in nearby Clear Lake. All snails taken from both
sampling sites were pooled into one sampling event based on date.
The same water system connects both sample sites, thus creating five
population samples. To obtain a comprehensive population sample,
teams explored the banks of sampling areas and pulled any observable
snails using nets. This frequently involved traveling via canoe to areas
inaccessible by foot. Bank sampling included careful exploration of
emergent macrophyte stands and within dense patches of freely
floating plants. For the Armand Bayou site, sampling days varied
between 8-10 hours on the water, with a consistent traveling distance
of approximately 2 km upstream. Researchers separated snails into
two broad groups based upon their operculum widths: juveniles (< 40
mm) and adults (> 40 mm). Operculum width serves as a good
indicator of snail size (Youens & Burks 2008) and suggestions for size
at maturity for P. insularum currently do not exist in the literature.
Cazzaniga ( 1 990) noted that an operculum size of 40 mm for larger P.
insularum may be similar to the 25 mm threshold in operculum width
that delineates smaller adult P. canaliculata. To test differences in
proportions of juveniles over different sample populations, all five
sample populations were compared against each other using a z-test of
significance for two proportions (Baldi & Moore 1996).

Data analysis revealed significant differences between proportions

TEXAS J. SCI. 61(4), NOVEMBER, 2009

325

of juveniles across sample populations, with all samples occurring in a
non-random distribution (all /^-values <0.05). According to pair-wise
z-tests, the sample from August of 2008 exhibited a significantly
higher proportion of juveniles than three of the other four samples,
with the September 2007 sample not statistically different than any of
the samples (Table 1). The largest difference between proportions
existed between the earliest sample, May 2006, and the most recent,
August 2008. The August 2008 sample showed a change in the size
structure distribution, which indicates a larger proportion of juveniles.

The increase in the overall proportion of juveniles collected from
2006 to 2008 (Table 1) suggests that the total number of juvenile P.
insularum present in the study sites increased over this period.
Juvenile snails (<40 mm) were not observed in samples taken from the
earliest sampling events. The use of consistent and comprehensive
sampling methods suggests that research teams did not simply miss
juveniles in earlier sampling events. The continued presence of egg
clutches observed during field studies coupled with the change in
proportion of juveniles indicates that invasive populations of P.
insularum in the study sites are increasing in overall size distribution.
An increasing number of juveniles may imply that exotic P. insularum
populations are growing in the aquatic ecosystems of southeast
Houston. For P. canaliculata, Carlsson & Bronmark (2006)
demonstrated that smaller snails exhibited higher feeding rates.
Therefore, an increase in the population size of non-native P.
insularum could produce more damage to aquatic vegetation due to
their higher rate of consumption (Boland et al. 2008; Burlakova et al.
2008). However, invasive ecologists need future research on
population size per unit area and snail consumption rates of
macroscopic plants to support these predictions.

For the first time, this study documents the presence of the growing
size distributions of populations of invasive P. insularum in southeast
Texas. However, scientists still know little about the ability of this
species to damage local ecosystems. Due to their recent presence,
only limited estimates on the reproductive ability of P. insularum exist

326

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

Table 1: Individual percentages for each size class present with the total number of snails collected
from each sampling event. Samples occur across six operculum uddth size classes with a
threshold at 40 mm for adults. Different small letters denote statistical significance between
sampling events in the total proportion of juveniles (10-40 mm) according to a z-test of
significance for proportions.

Sampling

Event

Juveniles

Adults

10-20

20-30

30-40

Significance

40-50

50-60

>60

May-06

100

73%

22%

May-07

4.8%

66.6%

28.6%

Sep-07

1.3%

abc

14.8%

72.7%

10.4%

May-08

1.2%

6.1%

8.5%

43.9%

40.3%

Aug-08

176

16.5%

18.8%

2.8%

11.9%

40.9%

9.1%

(Barnes et al. 2008). However, the potential population growth made
possible by females routinely laying large egg clutches (each
containing approximately 2000 eggs; Barnes et al. 2008) warrants
serious concern. Without further investigation of the population size
structure, invasive ecologists cannot accurately predict effects of P.
insularum on aquatic Texas ecosystems. Ecologists must conduct
future research, specifically density estimations and consumption
rates, to understand fully the overall effect P. insularum will have in
southeast Texas and possibly along the entire Gulf coast.

Acknowledgements

We thank the Texas Academy of Sciences, Southwestern
University and H-E-B for providing the funds necessary for this study.
Our gratitude also goes to Mark Kramer, George Regmond, Ann
Brinly and the entire staff at Armand Bayou Nature Center for helping
us with our data collection. We would also like to thank Rebecca
Marfurt, Sarah Hensley, Matt Barnes, Abby Youens and Brandon
Boland for providing initial life history data for P. insularum.

Literature Cited

Baldi, B. & D. S. Moore. 1996. The practice of statistics in the life sciences. W. H.
Freeman and Company, New York, 516pp.

Barnes, M. A., R. K. Marfurt, J. J. Hand & R. L. Burks. 2008. Fecundity of the exotic
applesnail, Pomacea insularum. Journal of the North American Benthological

TEXAS J. SCI. 61(4), NOVEMBER, 2009

327

Society, 27(3):738-745.

Boland, B., M. Meerhoff, C. Fosalba, N. Mazzeo, M. Barnes & R. Burks. 2008,
Juvenile snails, adult appetites: Contrasting resource consumption between two
species of applesnails (Pomacea). Journal of Molluscan Studies, 74(l):47-54.

Burlakova, L. E., A. Y. Karatayev, D. P. Padilla, L. D. Cartwright & D. N. Hollas. 2008.
Wetland restoration and invasive species: Apple snail {Pomacea insularum) feeding
on native and invasive aquatic plants. Restoration Ecology, 17(3): 433-440.

Cazzinga, N. J. 1990. Sexual dimorphism in Pomacea canaliculata. The Veliger,
33(4):384-388.

Carlsson, N. O. L., C. Bronmark & L. A. Hansson. 2004. Invading herbivory: The
golden apple snail alters ecosystem tlmctioning in Asian wetlands. Ecology,
85(6):1575-1580.

Carlsson, N. O. L. & C. Bronmark. 2006. Size-dependent effects of an invasive
herbivorous snail {Pomacea canaliculata) on macrophyte and periphyton in Asian
wetlands. Freshwater Biology, 51:695-704.

Carlsson, N. O. L. & J. O. Lacoursiere. 2005. Herbivory on aquatic vascular plants by
the introduced golden apple snail {Pomacea canaliculata) in Lao PDR. Biological
Invasions, 7:233-241.

Conner, S. L,, C. M. Pomory & P. C, Darby. 2008. Density effects of native and exotic
snails on growth in juvenile apple snails Pomacea paliidosa (Gastiopoda:
Ampullariidae): A laboratory experiment. Jounial of Molluscan Studies, 74:355-362.

Hayes, K. A., R. C. Joshi, S. C. Thiengo & R. H. Cowie. 2008. Out of South America:
Multiple origins of non-native apple snails in Asia. Diversity and Distributions,
14(4):701-712.

Howells, R. G., L. E. Burlakova, A. Y. Karatayev, R. K. Marfiirt & R. L. Burks. 2006.
Native and introduced Ampullariidae in North America: History, status, and ecology,
pp. 73-112, in Global advancements in ecology and management of golden apple
snails (R. Joshi, & L. Sebastian eds). Philippine Rice Research Institute, 588pp.

Karatayev, A. Y., L. E. Burlakova, V. A. Karatayev & D. K. Padilla. 2009.
Introduction, distribution, spread, and impacts of exotic freshwater gastropods in
Texas. Hydrobiologia, 619:181-194.

Levine, J. M. & C. M. Antonio. 2003. Forecasting biological invasions with increaseing
international trade. Conservation Biology, 17:322-326.

Rawlings, T. A., K. A. Hayes, R. H. Cowie & T. M. Collins. 2007. The identity,
distribution, and impacts of non-native apple snails in the continental United States.
BMC Evolutionary Biology, 7:97-1 1 1.

Robinson, D. G. 1999. Alien invasions: The effects of the global economy on
nonmarine gastropod introductions into the United States. Malacologia, 41(2):413-
438.

Youens, A. K. & R. B. Burks. 2008. Comparing applesnails with oranges: The need to
standardize measuring techniques when studying Pomacea. Aquatic Ecology,
42:679-684.

burksr@southwestem.edu

328

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

IN RECOGNITION OF THEIR ADDITIONAL SUPPORT OF
THE TEXAS ACADEMY OF SCIENCE DURING 2009

PATRON MEMBERS

Goldberg, Stephen R.
Killebrew, Don W.
Marsh, David S.
Strenth, Ned E.

SUSTAINING MEMBERS

Davidson, David L.
Kowalski, Joseph L.
Kruger, Joseph M.

Lee, Thomas E. Jr.
Valdes, Arcadio

SUPPORTING MEMBERS

Collins, James
Harper, Donald E., Jr.
Hettinger, Deborah D.

Looney, Michael
Lundelius, Ernest L., Jr.
McKinney, Larry
Sieben, John
Simpson, Lynn
Stevens, Fred
Weller, Milton W.

TEXAS J. SCI. 61(4), NOVEMBER, 2009

329

INDEX TO VOLUME 61 (2009)
THE TEXAS JOURNAL OF SCIENCE

Rigel K. Rilling

Department of Biology, Angelo State University
San Angelo, Texas 76909

This index has separate subject and author sections. Words,
phrases, locations, proper names and the scientific names of
organisms are followed by the initial page number of the articles in
which they appeared. The author index includes the names of all
authors followed by the initial page number of their respective
article(s).

SUBJECT INDEX

Ambystomajeffersonianum 61
Ammotragus lervia 1 5
amphibians 3, 61
Andropogon glomeratus 83
Anguillidae 3 1
Anisoptera 157
anthropogenic effects 3, 279
aoudad 1 5
Apicomplexa 311
Araneidae 203
arboreal nesting 163
Arkansas 1 5 1

central and southern 3 1
endemic fauna of 203
Arkansas Counties
Columbia County 3 1
arsenic-tolerant cultures 259
Atherinopsidae 3 1
avian taxonomy 195

bare ground 119,219
Bivalvia 203,279

brackish wetlands 83
Bromus japonicus 119

Caddell formation 181
Calidris minutilla 233
Catostomidae 3 1
Centrarchidae 3 1
Cephenemiya jellisoni 1 87
Cephenemiya phobifer 187
Cephenem iya pratti 187
Charadriiformes 233
chronic wasting disease 1 87
clutch size 1 3 1
Clypeasteroida 181
Coleoptera 203
Collembola 203
Costa Rica 147
cover board arrays 3
Cymbovula acicularis 67
Cyprinidae 3 1

330

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

detoxification, arsenic 259
density 295
diel 295

Dipodomys elator 119
Diplopoda 151, 203
Diptera 187,203
distance sampling 295
diversity 3, 83, 279
domestic goat 1 5
domestic sheep 1 5

ectoparasites 131, 187
endemic species 203
Environmental Protection Agency
(EPA) 259
Eocene, upper 1 8 1
Ephemeroptera 203
Erizathon dorsatum 65
escape terrain 1 5
Etheostoma asprigene 3 1
Etheostoma fusiforme 3 1
Eubacteria 259

facultative species 243
fatty acid profiles 45
fire 219

floodplain ecosystem 3
freshwater
wetlands 83
reservoir 233
frugivory 97
Fuirena simplex 83
Fundulidae 3 1
Fungi 203

Fusconaia askewi 279

gas chromatography 45
gas exchange 243
Gastropoda 67, 203, 323
Gekko smithii 225
Geographic Information System
(GIS) 15

global climate change 1 3 1
granivory 97
grazing intensity 1 19
grazing regime 119
ground coverage 119,219
Gulf Coast 67, 181

hatching success 131
Hematozoa 3 1 1
herpetofauna 3
Hiodon tergisus 3 1
Hiodontidae 3 1
Hirundo rustica erythrogaster
131

Hymenoptera 203

Icturalidae 3 1
Illinois 61
index fossil 181
infanticide 131
invasive species 323

Julida 151
K

karyotype 195

Laguna Madre 259
leaf litter 219

TEXAS J. SCI. 61(4), NOVEMBER, 2009

331

leaves 243
Leguminosae 243
Lertha extensa 45
limiting resource 163
lipid composition 45

Mabuya unimarginata 147
mass spectrometry 45
Mexico

Baja California 229
Coahuila 1 5

Sierra Maderas del Carmen 15
Sierra San Marcos y del Pino
15

Tamaulipas 67
Veracruz 317

minimum contaminant level 259
Mississippi 181
Muridae 97
Mussels 279
Mustela frenata 229
Mustelidae 229
Mycoplasma 259
Myxidium serotinum 61
Myxozoa 61

nasopharyngeal bots 187
Nemopteridae 45
nesting success 131
nestling success 131
Neuroptera 45
Notorus phaeus 3 1

Ochrotomys nuttalli 163
Odocoileus hemionus 1 87
Odocoileus virginianus 1 87
Oestridae 187

Oklahoma, east-central 311

Old Sabine Bottom Wildlife
Management Area 3
omnivory 97
organic substrate 2 1 9
overwintering 233
Ovis canadensis 1 5
Ovulidae 67
oxidation 259

Parajulidae 151
parasites 131,187,311
Passeriformes 311
Pelodictyon 259
Percidae 3 1
Percina sp. 31
Periarch us lyel li 181
Peromyscus pectoral is 97
phospholipid 45
photosynthetic rate 243
pine slough 219
polymerase chain reaction (PCR)
259

Pomacea insularum 323
population dynamics 323
Program R 195
Pseudoscorpionida 203
Pteronotropis hubbsi 3 1

radio-tracking 163
recolonization 219
recruitment 279
reproductive cycle 147, 225, 317
reptiles 3, 147,219,225,317
Rio Grande Valley, lower 295

332

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

Sabine River 279
Salinispora 259
sanctuaries, mussel 279
Schoenoplectus pungens 83
Scincella lateral is 219
Scincidae 147,219
Scutelliformes 1 8 1
seasonal flooding 1 63
seasonal leaf fall 219
seasonal trophic ecology 97
sediment samples 259
Sophora secundiflora 243
Spar tin a patens 83
species richness 3, 83
Squamata 147,219,225,317
Stephen F. Austin State Univ.

Experimental F orest 163
stomatal conductance 243
suburban habitat 323

Texas

central 97,157,243
eastern 3, 163
Edwards Plateau 65
northeast 131,279
south 259, 295
South Padre Island 83
west central Texas 233
Texas Cities
Houston 323

Texas Counties

Angelina County 163
Jones County 157
Houston County 163, 323
Nacogdoches County 163
Smith County 3
San Augustine County 163
Taylor County 157
Witchita County 119
Texas Wildlife Action Plan 279
transpiration 243
transplant and restoration 15
triacylglycerol 45
Trichoptera 203
trophic opportunism 97

Unionidae 279
V

vegetation height 119

water potential, leaf 243
wetland habitat 3, 83

Xenosaurus grandis 3 1 7
Z

Zenaia asiatica asiatica 295
Zenaia macroura 295

TEXAS J. SCI. 61(4), NOVEMBER, 2009

333

AUTHORS

Alaniz-Garcia, J. 229
Andrews, K. D. 311

Baccus, J. T. 97, 195, 295
Baird, A. B. 65
Bashan, M. 45
Bay,M.D. 311
Benn, S. J. 295
Berlanga, G. A. 259
Bradstreet, A. P. 163
Burks, R.L. 323

Cakmak, O. 45
Collins, M.L. 295
Contreras-Balderas, A. J. 15
Correa-Sandoval, A. 67
Crawford, J. A. 61

Edwards, C.W. 163
Espinosa-T., A. 15
Eubanks, T. M. 259

Ford,N.B. 3,279
Forstner, M. R. J. 195
Furuya, M. 243

Garcia- A., M. A. 15
Goetze, J. R. 119
Goldberg, S. R. 147,225,317
Gonzalez-Guzman, S. 229
Gullett,J. 279

Hall,D. W. 65
Hardwick, J, M. 97
Herriman, K. 3
Huffman, D. G. 97
Hunkapiller, T. R. 3

Johnson, B. W. 157
Judd,F. W. 83
Kainer, M. A. 97
Kasner, A. C. 233

Kelley, S.W. 187
Kopachena, J. G. 131
Kuhns, A. R. 61
Kyle,C.H. 323

LaDuc, T. J. 65
Lee,T.E. 157
Lonard, R. I. 83
Lowe,K.L. 259

Martinez-Gallardo, R. 229

Maxwell, T. C. 233

May, M. E. 279

McAllister, C. T. 31, 61, 151, 203

McDonough, J. P. 323

Nelson, A. D. 119
Nelson, M. 119

Patel, A. J. 157
Pauly, G. B. 65
Persans, M. W. 259

Robison, H.W, 31,151,203
Ruddick,R.H. 233
Ruiz-Campos, G. 229

Sandoval, A. V. 15
Satar, A. 45
Slay,M.E. 203
Small, M.F. 195,295
Strenth, N. E. 67

Trawick, M. K. 323
Tumlinson, R. 31
Turner, K.T. 131

Van Auken, O. W. 243
Vogtsberger, R. C. 157

Watson, C. M. 219
Watson, E. 119

Zachos, L. G. 181

334

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

REVIEWERS

The Editorial staff wishes to acknowledge the following individuals
for serving as reviewers for those manuscripts considered for publication
in Volume 61. Without your assistance it would not be possible to
maintain the quality of research results published in this volume of the
Texas Journal of Science.

Ammerman, Loren

Kakolesha, Nick

Quigg, Antonietta

Anderson, Todd

Keith, Don

Rincon-Zachary, Magaly

Baccus, John

Lee, Thomas

Ritzi, Chris

Barnes, Jeffrey

Lehman, Roy

Rylander, Ken

Branch, William

Lipscomb, Barney

Scales, John

Brant, Joel

Lonard, Bob

Shipley, Michael

Broussard, Greg

Longley, Glenn

Small, Michael

Bush, Janis

Mahrdt, Clark

Smith, Wayne

Choate, Larry

Masuoka, James

Stangl, Fred

Ciampaglio, Charles

Matthews, Bill

Starnes, Wayne

Cobb, George

McAllister, Chris

Stewart, Betty

Collins, Joseph

McDermott, Susanne

Stewart, Timothy

Cook, Jerry

McFarland, Anne

Strenth, Ned

Cook, Tamara

McMahon, Robert

Sudman, Phil

Fedynich, Alan

Miller, Tom

Thompson, Carol

Gagen, Charlie

Mills, Dana

Thompson, Cody

Goetze, Jim

Mitchell, Joseph

Tumlison, Renn

Goldberg, Stephen

Morehead, Sally

VanAuken, Bill

Harmel, Daren

Murray, Phil

Wang, Xixi

Hibbitts, Toby

Nelson, Allan

Williams, Hans

Higgins, Chris

Parmley, Dennis

Yancey, Thomas

Hoagland, Bruce

Purtlebaugh, Caleb

Zimmerman, Earl

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}a Has Not Changed During Preceding 12 Months

O Has Changed During Preceding 12 Months (Publisher must submit explanation of change with this statement)

PS Form 3526, September 2007 (Page 1 of 3 (Instructions Page 3)) PSN 7530-01 -000-9931 PRIVACY NOTICE: See our privacy policy on www.usps.com

THE TEXAS JOURNAL OF SCIENCE-VOL. 61, NO. 4, 2009

13. Publication Title

14. Issue Date for Circulation Data Below

The

Texas Journal of Science

November 2009

15. Extent and Nature of Circulation

Average No. Copies Each Issue
During Preceding 12 Months

No. Copies of Single issue
Published Nearest to

Filing Date

a. Total Number of Copies (Net press run)

900

<1)

Mailed Outside-County Paid Subscriptions Stated on

PS Form 3541 (Include paid distribution above nomi¬
nal rale, advertiser's proof copies, and exchange
copies)

658

b. Paid
Circulation
(By Mail

(2)

Mailed In-County Paid Subscriptions Slated on PS

Fonri 3541 (Include paid distribution above nominal
rate, advertiser's proof copies, and exchange copies)

Outside
the Mail)

<3)

Paid Distribution Outside the Mails Inciuding Sates

Through Dealers and Carriers, Street Vendors, Counter

Sales, and Other Paid Distribution Outside USPS®

157

(4)

Paid Distribution by Other Classes of Mall Through
the USPS (e.g, First-Class Mail®)

c. Total Paid Distribution (Sum of 1Sb (1), (2). (3), and (4))

830

(1)

Free or Nominal Rate Outside-County

Copies included on PS Form 3541

d. Free or
Nominal
Rate

Distribution

(2)

Free or Nominal Rate In-County Copies Included
on PS Form 3541

(By Mail
and

Outside
the Mail)

(3)

Free or Nominal Rale Copies Mailed at Other

Classes Through the USPS (e g. First-Class Mall)

(4)

Free or Nominal Rate Distribution Outside the Mail
(Carriers or other means)

e Total Free or Nominal Rate Distribution (Sum of 15d(1), (2), (3) and (4))

. 0 .

f. Total Distribution (Sum of f5c and ISe) ^

830

g Copies not Distributed (See fosfrucf/ons to PuW/sfters #4 (page #3j) ^

h. Total (Sum of 1 51 and g) ^

900

I Percent Paid w

(IScdivided by 15f times 100) *

100%

16. Pubtication of Statement of Ownership

If the publication is a qeneral publication, publication of this statement is required. Will be printed Q Publication not required,

in the _ _ issue of this publication

17. Signature and TiUe of Editor, Publisher. Business Manager, or Owner

Date

E'd: k.

25 Sept. 2009

I certify that all information furnished on this form is true and campiste. I understand that anyone who furnishes false or misleading information on this
form or who omits material or information requested on the form may be subject to criminal sanctions (including fines and imprisonment) and/or civil
sanctions (including civil penalties).

PS Form 3526, September 2007 (Page 2 of 3)

THE TEXAS ACADEMY OF SCIENCE, 2009-2010

OFFICERS

President
President Elect:
Vice-President:

Immediate Past President:
Executive Secretary:
Corresponding Secretary:
Managing Editor:

Manuscript Editor:

Treasurer:

AAAS Council Representative:
International Coordinator:

William J. Quinn, St. Edward’s University

Benjamin A. Pierce, Southwestern University

Romi L. Burks, Southwestern University

Raymond C. Mathews, Jr., Texas Water Dev. Board

Fred Stevens, Schreiner University

Diane B. Hyatt, Texas Water Development Board

Ned E. Strenth, Angelo State University

Frederick B. Stangl, Jr., Midwestern State University

John A. Ward, Brooke Army Medical Center

James W. Westgate, Lamar University

Armando J. Contreras, Universidad Autonoma de N.L.

DIRECTORS

2007 Renard L. Thomas, Texas Southern University
Bob Murphy, Texas Parks and Wildlife Department

2008 Christopher M. Ritzi, Sul Ross State University
Andrew C. Kasner, Audubon Texas

2009 Ana B. Christensen, Lamar University

Thomas L. Arsuffi, Texas Tech at Junction

SECTIONAL CHAIRPERSONS

Anthropology: Raymond Mauldin, University of Texas at San Antonio
Biomedical: G. Scott Weston, University of the Incarnate Word
Botany: David Lemke, Texas State University

Cell and Molecular Biology: Magaly Rincon-Zachary, Midwestern State University

Chemistry and Biochemistry: J. D. Lewis, St. Edward’s University

Computer Science: James McGuffee, St. Edward’s University

Conservation Ecology: Wendi Moran, Hardin-Simmons University

Environmental Science: Kenneth R. Summy, University of Texas-Pan American

Freshwater Sciences: Matt Chumchal, Texas Christion University

Geosciences: Chris Barken, Stephen F. Austin State University

Marine Sciences: Larry D. McKinney, Harte Research Institute

Mathematics: Elsie M. Campbell, Angelo State University

Physics: David L. Bixler, Angelo State University

Science Education: Patricia Ritschel-Trifilo, Harden-Simmons University

Systematics and Evolutionary Biology: Tara Maginnis, St. Edward’s University

Terrestrial Ecology and Management: Richard Patrock, St. Edward’s University

COUNSELORS

Collegiate Academy: David S. Marsh, Angelo State University
Junior Academy: Vince Schielack, Texas A&M University

PERIODICALS

THE TEXAS JOURNAL OF SCIENCE
Texas Academy of Science
CMB 629

Wayland Baptist University
Plainview, Texas 79072

SMITHSONIAN INSTITUTION LIBRARIES

3 9088 01558 5730

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