MARINE AND FISHERY SCIENCES 36 (2): 149-163 (2023)

https://doi.org/10.47193/mafis.3622023010508

ABSTRACT. Coastal estuarine ecosystems serve as nursery habitats for many commercially and

recreationally important fishes. Biodiversity is a structural indicator and has been used as a metric

for conservation and management. In the hypersaline Lower Laguna Madre of Texas, a variety of

organisms makes their living in and around the dominant seagrass vegetation. This study provides

a general assessment of forage fishes biodiversity collected seasonally with bag seines in two sites:

Holly Beach (HB) and South Bay (SB) within the most southern Texas bay system as part of a

broader study on fish biology. A total of 15,880 fishes representing 32 species were collected during

four quarterly samplings through a year (11,795 from HB and 4,085 from SB). Both sites are inter-

connected as no fishes similarities difference were found, nonetheless, the sites’ variable character-

istics (i.e. basin area, seagrasses coverage, connection to the Gulf of Mexico) resulted in significant

greater species richness, relative abundances, and diversity in HB than SB for most of the year, sug-

gesting differences in habitat quality or at the very least variation in the availability of habitat types,

which are known to contribute to differences in fish diversity attributes.

Key words: Fishing gear, estuaries, coastal fishes, baitfish, nursery habitat.

Biodiversidad de peces de forraje en la Laguna Madre Inferior, en el extremo sur de Texas

RESUMEN. Los ecosistemas de estuarios costeros sirven como hábitats de crianza para muchos

peces de importancia comercial y recreativa. La biodiversidad es un indicador estructural y se ha

utilizado como métrica para la conservación y la gestión. En la hipersalina Laguna Madre Inferior

de Texas, una variedad de organismos vive en y alrededor de la vegetación de pastos marinos domi-

nante. Este estudio proporciona una evaluación general de la biodiversidad de peces de forraje reco-

lectados estacionalmente con redes de cerco en dos sitios: Holly Beach (HB) y South Bay (SB) den-

tro del sistema de bahías al sur de Texas, como parte de un estudio más amplio sobre la biología de

peces. Durante cuatro muestreos trimestrales a lo largo de un año, se recolectaron un total de 15.880

peces (11.795 de HB y 4.085 de SB) que representan 32 especies. Ambos sitios están interconecta-

dos, ya que no se encontraron diferencias en las similitudes de peces; sin embargo, las características

variables de los sitios (es decir, área de la cuenca, cobertura de pastos marinos, conexión con el

Golfo de México) dieron como resultado una riqueza de especies, abundancias relativas y diversidad

significativamente mayores en HB que en SB durante la mayor parte del año, lo que sugiere dife-

rencias en la calidad del hábitat o, al menos, variación en la disponibilidad de tipos de hábitat, que

se sabe que contribuyen a las diferencias en los atributos de diversidad de peces.

Palabras clave: Artes de pesca, estuarios, peces costeros, carnada, hábitat de cría.

149

*Correspondence:

carlos.cintra@utrgv.edu

Received: 3 February 2023

Accepted: 28 March 2023

ISSN 2683-7595 (print)

ISSN 2683-7951 (online)

https://ojs.inidep.edu.ar

Journal of the Instituto Nacional de

Investigación y Desarrollo Pesquero

(INIDEP)

This work is licensed under a Creative

Commons Attribution-

NonCommercial-ShareAlike 4.0

International License

Marine and

Fishery Sciences

MAFIS

ORIGINAL RESEARCH

Biodiversity of forage fishes in the Lower Laguna Madre, southernmost

Texas

DAVID CAMARILLO JR., ELIZABETH MOGUS GARCIA and CARLOS E. CINTRA-BUENROSTRO*

Ocean, Coastal Environmental and Ecological Assessment Lab., School of Earth, Environmental, and Marine Sciences, University of Texas

Rio Grande Valley (UTRGV), One West University Blvd., 78520 - Brownsville, USA. ORCID Elizabeth Mogus Garcia

https://orcid.org/0009-0003-0268-7686, Carlos E. Cintra-Buenrostro https://orcid.org/0000-0002-5870-9624

INTRODUCTION

Coastal estuaries usually consist of a variety of

structured and productive ecosystems such as

marshes, mangroves, seagrasses, and reefs (Beck

et al. 2001) that act as a habitat for a variety of

organisms and create a nursery system for many

commercially important fish species all over the

coastal world (Hampel et al. 2005; Whitfield

2016; Breaux et al. 2019), many spawning off-

shore and using these areas as larvae and juve-

niles (Heck et al. 1997). Economically, these

ecosystems are incredibly important locally,

regionally, and nationally. In the United States of

America (USA), the Texas coast alone brings in

about USD 2 billion for recreational fishing, USD

5.4 million from tourism, and USD 250 million

for seafood production (Rosen 2013).

Smaller and lower trophic level schooling fish-

es, known as forage fish, are usually found to be

prey items to larger predatory fishes within the

estuary. Forage fishes populate these ecosystems

playing an important role within the community

and trophic web as they make up a large part of

the diet of many higher-level predators that use

the estuary to feed and grow (Pikitch et al. 2012;

Faletti et al. 2019). Many of these forage fishes

will stay within the estuary their entire life,

whereas others will travel offshore to spawn

(Murphy and Taylor 1989; Wilson and Nieland

1994; Brown-Peterson et al. 2002; Faletti et al.

2019).

On the Gulf coast of Texas, which stretches

about 644 km, there are about 10,522 km2of

estuary habitat and seven major bay systems

(Rosen 2013). Along the Texas coast, many of

the estuaries have a high presence of seagrass

that allows for sediment stabilization, nutrient

cycling, protection, and detrital production and

export (Heck et al. 2003). The most southern bay

system of the Texas Gulf coast is the Laguna

Madre. The most southern portion of the Laguna

Madre, the Lower Laguna Madre (LLM), has

been considered as hypersaline due to little fresh-

water inputs, few inlets to the Gulf of Mexico

(GOM), and high evaporation (Tunnell and Judd

2002; Rosen 2013; Kowalski et al. 2018). The

LLM has a high presence of seagrasses mostly

including Turtle Grass (Thalassia testudinum K.

D. Koenig, 1805) and Shoal Grass (Halodule

wrightii (Ascherson, 1868)), but other species

such as Manatee Grass (Syringodium filiforme

Kützing, 1841), Widgeon Grass (Ruppia mariti-

ma Linnaeus, 1753), and Clover Grass (Halophi-

la engelmannii Ascherson, 1875) can be found

(Sheridan and Minello 2003). It is expected to

find a higher number of fishes residing in areas

of the estuary that have higher densities of sea-

grasses and other plant structures, which was

shown in the LLM by Sheridan and Minello

(2003). Though with changing climatic condi-

tions, anthropogenic disturbances, and the

increase of cold fronts and storm surges, many

organisms are seeing declines, mass mortalities,

or displacement (Sheridan and Minello 2003;

Kowalski et al. 2018).

Due to the high productivity within these

ecosystems, a high diversity of species (fishes

and invertebrates) is usually found (Beck et al.

2001) which leads to interactions among differ-

ent species using the same resources (Whitfield

2016). This creates biotic interactions that may

affect local species distributions and abundances.

An ecosystem’s productivity is largely attributed

to the area’s biodiversity, making biodiversity a

crucial metric for conservation and management

(Pawluk et al. 2021). Fish diversity within the

marine environment is constantly changing and

will continue to fluctuate with changing environ-

mental conditions. Habitat heterogeneity, basin

area, physicochemistry of the water, primary

productivity, resources availability and historical

factors are known drivers of fishes diversity

(Tonn 1990; Ricklefs and Schluter 1993; Gel-

wick et al. 2001; Ricklefs 2004; Auber et al.

2017; Thompson et al. 2020). As Pawluk et al.

150 MARINE AND FISHERY SCIENCES 36 (2): 149-163 (2023)

(2021) pointed out, it is important to assess the

patterns in fish abundances and species richness

within marine systems in order to better under-

stand community dynamics given changing tem-

peratures and environmental conditions and

point out what potential vulnerabilities there are.

Given the importance of biodiversity and their

connection with productivity and conservation, it

is crucial to characterize these low-studied areas

(as is the case in the southernmost Texas coast)

and understand the dynamics of the forage fish

community that can have a major impact on the

local and state economies.

This study is derived from a broader one (to

be published elsewhere) focusing on the age,

growth, diet, and trophic web reconstruction of

Sciaenops ocellatus and looks at forage fishes’

diversity to better understand if there are any

changes due to shifting environmental factors

that are continually increased by climate

change. Objectives of this study were: (1) com-

pare forage fishes’ biodiversity between two

sites within the LLM, a bay and a lagoon, and

(2) identify variations within a single year as a

driver of any changes in forage fishes biodiver-

sity by using netting to capture and record

species along the shoreline of both sites.

Because of differences in basin area (the lagoon

site being smaller as detailed in methods), con-

nection to the GOM, and seagrass coverage, it is

hypothesized that the bay system will have

greater fishes diversity than the lagoon system.

Furthermore, fishes biodiversity is expected to

fluctuate throughout the year due to a combina-

tion of factors including fishes migration cycles

(e.g. Livingston et al. 1976; Timmerman et al.

2021), spawning and recruitment, juveniles sur-

vivorship (Livingston et al. 1976; Morin et al.

1985; Meffe and Berra 1988; Yoklavich et al.

1991), movement and dispersal of fishes that

might be affected by home ranges, and degree of

connectivity as well as physical barriers (Liv-

ingston et al. 1976; Yoklavich et al. 1991; Gel-

wick et al. 2001).

MATERIALS AND METHODS

Study sites

Study sites are South Bay (SB) and Holly

Beach (HB), both located in Cameron County,

southern Texas, USA. This area has several

knowledge gaps including baseline studies as

compared to other geographic locations in the

country. Both sites are part of the LLM, which is

one of the six largest hypersaline estuarine sys-

tems in the world (Tunnell and Judd 2002; Mar-

quez et al. 2017). The climate is categorized as

semiarid and subtropical (Tunnell and Judd 2002;

Marquez et al. 2017). Estuaries of the LLM are

connected to the GOM at the Brazos-Santiago

Pass which was created in the 1930’s (Tunnell

and Judd 2002; Marquez et al. 2017).

South Bay (26° 01'20.6"N-97° 11'03.8"W)

is classified as a bay system by name only, as it is

an enclosed lagoon which only connects to the

Brownsville Ship Channel through a narrow

opening (Figure 1). It is located south of the Bra-

zos-Santiago Pass. It is a shallow body of water

and connects to the Rio Grande River on the

south end. It has an average depth of 0.85 ±0.15

m with an area of about 14.2 km2(Marquez et al.

2017). Holly Beach (26° 07'30.5"N-97° 17'

48.4"W) is classified as a lagoon system and is

located north of SB and the Brazos Santiago Pass

(Figure 1). It lies between the Laguna Atascosa

National Refuge and the LLM with the Laguna

Vista Cove on the south end (Murphy et al. 2021).

It has an average depth of about 1 m and has a

great seagrass bed presence. As HB is part of the

LLM aquatic system, for the purposes of this

study, an area of ~42 km2was estimated using

Google Earth (2022). Both study sites are popular

bodies of water for fishing and birding as they

provide a variety of ecosystem services and a

habitat for many fishes of commercial and recre-

ational importance.

151

CAMARILLO JR. ET AL.: FORAGE FISHES IN SOUTHERNMOST TEXAS

Sampling

Sampling procedure for this study followed the

protocol from the Texas Parks and Wildlife

Department (TPWD), which uses bag seines as

part of their survey efforts. Bag seines utilized

were made to replicate those from TPWD, which

are 18.3 m long with 1.3 cm stretched nylon #5

multifilament mesh in the bag, and 1.9 cm

stretched nylon #5 multifilament mesh in the

wings and were used along the shore of both sites.

Sampling effort goal was five bag seine replicates

per site for a total of 20 replicates per quarter (Q),

given four overall visits to each site. Due to

restricted access to SB by SpaceX during rockets

testing, unpredictable weather, and a set limit of

300 individual S. ocellatus each Q under the Uni-

versity of Texas Rio Grande Valley, Institutional

Animal Care and Use Committee (IACUC, AUP-

19-40). Bearing in mind that, in order to comply

with the approved protocol under AUP-19-40, no

further sampling occurred once the set number of

S. ocellatus was achieved on each Q (i.e. 300

between both sites); hence, a balanced sampling

effort was not achievable. This resulted in vari-

able numbers in replicates per site visit, as well as

sample area covered. Nonetheless, the sampling

effort goal was exceeded every Q as follows: Q1

21 samples (n), Q2 n =35, Q3 n =36, and Q4 n

=30. For future comparison purposes to the

TPWD database, sampling was diurnal and

occurred during high tides. A total area of about

720 m2was covered at both sites. Fishes were

counted and identified for each bag seine repli-

cate. All fishes (other than S. ocellatus) and their

count were noted and then released back into the

water. For identification purposes, when needed

two voucher specimens of each species were kept

and taken back to the laboratory.

Statistical analyses

Fishes’ data were analyzed in Primer v7,

before any routine a log (abundance +1) was nec-

essary as determined by a shade plot as per

Clarke et al. (2014), which down-weighted con-

tributions by highly abundant species and allow

low-and-mid-range species to also influence

assemblage similarities calculations (Clarke and

Warwick 2006). The following Primer routines

152 MARINE AND FISHERY SCIENCES 36 (2): 149-163 (2023)

Figure 1. Gulf of Mexico map and study sites Holly Beach and South Bay located within the Lower Laguna Madre, Texas, United

States of America. Map modified from Google Earth.

Holly Beach

Gulf of Mexico

South Bay

Gulf of Mexico

United States of America

Mexico

Cuba

and tests allowed comparisons between fish

assemblages from both sites throughout the year

on a quarterly basis. Natural groups of fishes at

each site for every Q were identified with group-

averaged cluster analysis based on Bray-Curtis

similarities and non-metric multidimensional

scaling (nMDS), only the latter was presented

graphically (Figure 2), followed by a similarity

profile (SIMPROF) test to identify genuine

groups. A similarity percentage (SIMPER) test

was performed to determine fishes that con-

tributed to assemblages’ dissimilarities among Qs

and sites. Because there are only two sites, a one-

way ANOSIM was performed on Qs. The

DIVERSE routine was utilized to obtain species

richness, Shannon-Wiener diversity (H’ here-

after), and Jaccard evenness (J’ hereafter). These

metrics as well as the overall abundance (abun-

dance hereafter) per Q at each site were compared

with a model 1 two-way analysis of variance

(ANOVA) unless otherwise noticed (see below).

Fishes’ dominance ratios (DRs) were estimated

by locality for each Q and were compared graph-

ically using descriptive statistics (mean ±stan-

dard deviations), as no further statistical analyses

were deemed appropriate because of the effect of

highly abundant taxa in a given Q, which skewed

the fishes’ dominance ratio distributions.

Prior to performing any ANOVA, fish species

richness, abundance, H’, and J’s at each site were

subjected to Q-Q plots and Kolmogorov-Smirnov

tests to verify normality, while homoscedasticity

was evaluated with Levene’s test (Zar 1996).

Fishes’ abundance violated both assumptions and

were subjected to a log10 +1 transformation (Zar

1996). Species richness was not normally distrib-

uted, but the ANOVA was deemed robust enough

for such violation (Underwood 1997) and was

performed on non-transformed values. Diversity

was heteroscedastic and the log10 +1 transforma-

tion made data even less homoscedastic but as the

variances difference was <3 times, the ANOVA

was deemed robust for this violation and was per-

formed on non-transformed values. Evenness was

not complaint to either assumption (i.e. normality

or homoscedasticity) and log10 +1 transformation

made it worst. In this case, the ANOVA could

have been still performed as ANOVA is based on

153

CAMARILLO JR. ET AL.: FORAGE FISHES IN SOUTHERNMOST TEXAS

Figure 2. Non-metric multidimensional scaling plot of fishes’ assemblages similarities. Ovals indicate 50% resemblance levels.

Holly Beach =HB, South Bay =SB, quarter =Q. Q1 =October-December 2020; Q2 =January-March 2021; Q3 =April-

June 2021; Q4 =July-September 2021.

Transform: Log(X+1)

Resemblance: S17 Bray-Curtis similarity

SB1

HB1

SB2

HB2

SB3

HB3

SB4

HB4

2D Stress: 0.03

SiteQuarter

SBQ1

HBQ1

SBQ2

HBQ2

SBQ3

HBQ3

SBQ4

HBQ4

means which conforms to the Central Limit Theo-

rem making the assumption of normality not too

critical, and the same reasoning used for H’ might

have been applied to the J’ lack of homoscedastic-

ity compliance. Hence, the non-parametric option

(i.e. Kruskal-Wallis) was performed as an aca-

demic exercise only, because it is sensitive to

departures from homoscedasticity (Underwood

1997). Noteworthy, Kruskal-Wallis runs only as a

one-way therefore two tests were performed one

for Qs and the second one for sites. A Tukey Hon-

est Significant Difference (Tukey hereafter) test

was performed when ANOVA indicated signifi-

cant differences to identify the Qs responsible for

them (Zar 1996). Note as there are only two sites

the software does not perform this test for that

variable and issues a warning, but the difference

indicated by the ANOVA remains and therefore

can be determined from the mean values. All para-

metric statistics were performed with SPSS v27.

RESULTS

A total of 32 fishes (Table 1) yielded 15,880

individuals collected during the year of sampling,

74.3% of them captured at HB. Anchoa mitchilli,

Cyprinodon variegatus, Lagodon rhomboides,

and Micropogonias undulatus were present year-

round at both sites (Table 1). Fishes’ richness in

HB was 29 with ten species only present there,

while SB had 22 species, three occurring only at

SB (Table 1). Species richness varied also by Q

ranging from 12-16 fishes at HB with 12 species

occurring in a single Q, while the range for SB

was 8-14 fishes with also 12 species occurring in

a single Q (Table 1).

Similarity of fishes yielded two major clusters

separating at 40.2%, one cluster included Q2 from

both sites as well as Q3 at HB, all other similari-

ties occurred in the second cluster (figure not

shown); however, both clusters were not signifi-

cantly different from one another. Quarters were

not significantly different from one another as per

ANOSIM (R =0.417, p =0.114) supporting the

SIMPROF test. Separation of Q2 was also depict-

ed by the nMDS, with Q3 at HB in between the

remaining groups (Figure 2). Similarity among

samples ranged from 24.4 to 77.4%, with Qs 1 and

4 at SB being the more similar to one another; and

Qs 2 and 4 at SB being more dissimilar from each

other. Overall, seven or 12 fishes were needed to

explain >90% of the similarities: L. rhomboides,

A. mitchilli, M. undulatus, S. ocellatus, Eucinosto-

mus gula, Lutjanus griseus, and Brevoortia

patronus explained the similarities between Qs 1

and 4 at SB, while L. rhomboides, A. mitchilli, C.

variegatus, B. patronus, Hippocampus sp., Mugil

cephalus, Lutjanus griseus, Bairdiella chrysoura,

Opsanus beta, Synodus foetens, Hemiramphus

brasiliensis, and Fundulus grandis explained sim-

ilarities between Qs 2 and 4 at SB.

Mean ±standard deviation fishes’ richness at

HB ranged from 3.27 ±1.94 to 6.75 ±1.81, while

at SB the range was from 2.75 ±1.04 to 3.47 ±

1.46. There were significant differences between

sites (F0.05 (1,121) =60.66, p <0.001) with HB hav-

ing greater richness than SB, and among Qs

(F0.05 (3,121) =6.47, p <0.001) (Figure 3 A). The

Tukey test indicated Q4 at HB was significantly

lower than the other 3 Qs, which were not differ-

ent from one another; while at SB Q4 was signif-

icantly larger than all other Qs, which were not

significantly different among (Figure 3 A). How-

ever, but as expected, the interaction effect was

also significant (F0.05 (3,121) =8.86, p <0.001).

Fishes’ abundances (mean ±standard devia-

tion) ranged from 84.73 ±95.24 to 268.00 ±

248.33 at HB, and from 29.00 ±25.80 to 93.80 ±

38.26 at SB (Figure 3 B). Holly Beach had a sig-

nificantly larger number of individuals (log10

transformed) than SB (F0.05 (1,121) =19.25, p <

0.001), significant differences also occurred

among Qs (F0.05 (3,121) =5.79, p =0.001), and the

interaction between sites and Qs (F0.05 (3,121) =

14.00, p <0.001). At both sites, Qs 1, 2, and 4

were not significantly different from one another,

154 MARINE AND FISHERY SCIENCES 36 (2): 149-163 (2023)

155

CAMARILLO JR. ET AL.: FORAGE FISHES IN SOUTHERNMOST TEXAS

Table 1. Fishes’ richness by site and quarter (Q) presented in alphabetical order. Holly Beach =HB; South Bay =SB; Q1 =

October-December 2020; Q2 =January-March 2021; Q3 =April-June 2021; Q4 =July-September 2021; Freshw. =

freshwater.

Species HB SB Environment1

Anchoa mitchilli (Valenciennes, 1848) Q1, Q2, Q3, Q4 Q1, Q2, Q3, Q4 Euryhaline

Archosargus probatocephalus (Walbaum, 1792) Q3, Q4 Marine/brackish

Bairdiella chrysoura (Lacepède, 1802) Q1 Marine/brackish

Brevoortia patronus Goode, 1878 Q3 Q4 Euryhaline

Chaetodipterus faber (Broussonet, 1782) Q3 Marine/brackish

Cynoscion nebulosus (Cuvier, 1830) Q1, Q4 Marine/brackish

Cyprinodon variegatus Lacepède, 1803 Q1, Q2, Q3, Q4 Q2, Q3, Q4 Euryhaline

Elops saurus Linnaeus, 1766 Q2 Marine/brackish

Eucinostomus gula (Quoy and Gaimard, 1824) Q1, Q4 Q1, Q4 Euryhaline

Fundulus grandis Baird and Girard, 1853 Q1, Q2, Q3, Q4 Q2, Q4 Freshw./brackish

Fundulus majalis (Walbaum, 1792) Q2, Q3 Q2 Marine/brackish

Hemiramphus brasiliensis (Linnaeus, 1758) Q1 Q4 Marine

Hippocampus sp. Q1 Q1, Q2

Kathetostoma albigutta Bean, 1892 Q3 Q4 Marine

Lagodon rhomboides (Linnaeus, 1766) Q1, Q2, Q3, Q4 Q1, Q2, Q3, Q4 Euryhaline

Lucania parva (Baird and Girard, 1855) Q3 Marine/brackish

Lutjanus griseus (Linnaeus, 1758) Q1, Q4 Q1, Q4 Euryhaline*

Micropogonias undulatus (Linnaeus, 1766) Q1, Q2, Q3, Q4 Q1, Q2, Q3, Q4 Marine/brackish

Mugil cephalus Linnaeus, 1758 Q1, Q2, Q3, Q4 Q2, Q3, Q4 Euryhaline*

Mugil curema Valenciennes, 1836 Q3 Q4 Euryhaline*

Oligoplites saurus (Bloch and Schneider, 1801) Q1 Euryhaline*

Opsanus beta (Goode and Bean, 1880) Q1, Q2, Q3, Q4 Q1, Q3 Marine

Orthopristis chrysoptera (Linnaeus, 1766) Q4 Marine/brackish

Paralichthys lethostigma Jordan and Gilbert, 1884 Q1, Q2, Q3 Q3 Euryhaline*

Pogonias cromis (Linnaeus, 1766) Q3, Q4 Marine/brackish

Sciaenops ocellatus (Linnaeus, 1766) Q2, Q3 Q2 Marine/brackish

Scomberomorus maculatus (Mitchill, 1815) Q4 Marine

Strongylura marina (Walbaum, 1792) Q1, Q3, Q4 Euryhaline

Syngnathus louisianae Günther, 1870 Q2, Q3 Marine

Syngnathus sp. Q1

Synodus foetens (Linnaeus, 1766) Q3 Q1 Euryhaline

Trachurus trachurus (Linnaeus, 1758) Q4 Q4 Marine

1From Froese and Pauly (2022).

*Reported as occasional in freshwater or penetrating rivers, and thus considered euryhaline.

but Q3 was significantly different than Qs 2 and

4 (Figure 3 B) as per Tukey’s test.

Fishes’ H’ (mean ±standard deviation) at HB

ranged from 0.88 ±0.62 to 1.70 ±0.27, while in

SB ranged from 0.68 ±0.36 to 1.09 ±0.35 (Fig-

ure 3 C). As for the previous two metrics, there

were significant differences in H’ between sites

(F0.05 (1,121) =46.55, p <0.001), among Qs

(F0.05 (3,121) =8.32, p <0.001), and their interac-

tion (F0.05 (3,121) =7.40, p <0.001) (Figure 3 C).

Holly Beach once more had higher values than

SB for most Qs, while the Tukey test indicated

Q4 was significantly different from the other Qs,

which were not different from one another at both

study sites (Figure 3 C).

Mean ±standard deviation fishes’ J’ range at

HB was 0.83 ±0.16 to 0.90 ±0.03, and at SB 0.73

±0.12 to 0.94 ±0.04 (Figure 3 D). Fishes’ J’ was

significantly larger at HB than at SB (H0.05 (1) =

4.08, p =0.043), and among Qs (H0.05 (3) =24.28,

p <0.001) (Figure 3 D). Regarding J’ Q2 was sig-

nificantly different than the others, which were

not different from one another at both sites (Fig-

ure 3 D) as per the pairwise comparisons.

Lagodon rhomboides were overwhelmingly

the more abundant fish in SB during Qs 1, 3, and

4 DR =0.93, 0.90, and 0.93, respectively); while

Q2 in SB was dominated by M. undulatus (DR =

0.37). In HB, the same two species were also

dominant during the same Qs with L. rhomboides

156 MARINE AND FISHERY SCIENCES 36 (2): 149-163 (2023)

Figure 3. Mean ±standard deviation (Std. Dev., bars) metrics for each quarter (Q) in the study sites. A) Species richness, B) abun-

dance, C) diversity, and D) evenness. Holly Beach =HB, South Bay =SB. Sample size (n) as follows: HBQ1 n =13,

SBQ1 n =8, HBQ2 n =16, SBQ2 n =19; HBQ3 n =21; SBQ3 n =15; HBQ4 n =15; SBQ4 n =15. Q1 =October-

December 2020; Q2 =January-March 2021; Q3 =April-June 2021; Q4 =July-September 2021.

Holly Beach South Bay Holly Beach South Bay

Mean species richness ± Std. Dev.

Mean Std. Dev.abundance ±

500

400

300

200

100

Q2 Q3 Q4

Mean Std. Dev.diversity ±

Quarter

2.5

2.0

1.5

1.0

0.5

0.0 Q1

Mean Std. Dev.evenness ±

1.2

1.0

0.8

0.6

0.4

0.2

0.0

Q2 Q3 Q4

Quarter

Q2 Q3 Q4

Quarter

Q1 Q2 Q3 Q4

Quarter

bdca

DR =0.69, 0.76, and 0.85 in Qs 1, 3, and 4,

respectively; the DR in Q2 for M. undulatus was

0.39. The second highest DR (0.03) in SB for Q1

was for E. gula followed by A. mitchilli (DR =

0.02); for Q2 A. mitchilli had the 2nd highest DR

(0.32), and S. ocellatus occupied the 3rd position

with a DR of 0.16. Anchoa mitchilli had the 2nd

highest DR (0.06) during Q3 in SB followed by

C. variegatus (DR =0.02), while the 2nd and 3rd

place in the last Q were occupied by E. gula and

C. variegatus, respectively. Eucinostomus gula

also occupied the 2nd highest position with a DR

of 0.14 in HB during Q1 followed by F. grandis

(DR =0.08); 2nd and 3rd place during Q2 were

occupied by F. grandis (DR =0.29) and L. rhom-

boides (DR =0.13). Dominance was 2nd highest

for Mugil cephalus (0.13) in HB for Q3 while A.

mitchilli occupied 3rd place (DR =0.007), and

for the last Q E. gula had the 2nd highest DR

(0.06) and M. cephalus the 3rd one (DR =0.05).

Quarter 2 was the more evenly divided in terms

of fishes’ dominance as the highest abundances of

L. rhomboides resulted in high mean dominance

ratio values for the other three Qs, particularly in

SB (Figure 4).

DISCUSSION

In general, HB showed greater species rich-

ness, relative abundances, and H’ than SB for Qs

1-3; while in Q4 SB had greater values than HB.

This opposite result is likely one cause for the

significant interaction effect for relative abun-

dances and H’. As SB and HB have different

habitat characteristics (e.g. basin area, connection

to the GOM, seagrasses coverage, among others),

the observed significant differences between sites

are not surprising as HB was expected to host a

greater number of fishes than SB.

Another cause for the interaction effect

between sites and Qs is the expected changes in

fishes’ presences throughout the year, which also

helps explain the significant differences observed

in time. Twelve species were present only during

one Q at each site. Such differences are explained

by a combination of factors, e.g. fishes migration

cycles (Livingston et al. 1976; Timmerman et al.

2021), spawning, recruitment, and juvenile sur-

vivorship (Livingston et al. 1976; Morin et al.

157

CAMARILLO JR. ET AL.: FORAGE FISHES IN SOUTHERNMOST TEXAS

Figure 4. Mean ±standard deviation (Std. Dev., bars) fishes dominance ratio for each quarter (Q) in the study sites. Abbreviations

and sample sizes as in Figure 3.

Q1 Q2 Q3 Q4

Mean ominance atio Std. Dev.dr±

1.2

1.0

0.8

0.6

0.4

0.2

0.0

Quarter

Holly Beach South Bay

1985; Meffe and Berra 1988; Yoklavich et al.

1991), movement and dispersal of fishes which

might be affected by physical barriers, home

ranges, and degree of connectivity (Livingston et

al. 1976; Yoklavich et al. 1991; Gelwick et al.

2001).

As HB is not only more open to the GOM but

also has a larger area than SB, it is able to support

a more diverse ichthyofauna with the following

species only occurring there: Archosargus proba-

tocephalus, Chaetodipterus faber, Cynoscion

nebulosus, Elops saurus, Oligoplites saurus,

Pogonias cromis, Scomberomorus maculatus,

Strongylura marina, Syngnathus louisianae, and

Syngnathus sp. Out of these ten fishes, five were

present during a single Q. Three of them were

reef-associated species: Chaetodipterus faber

occurred only during the spring (i.e. Q3) and it is

known to be present in estuaries, particularly as

juveniles (Froese and Pauly 2022); while E.

saurus present in winter (i.e. Q2) are known to

occur in shallow onshore areas with common

occurrence of juveniles in lagoons (Cervigón et al.

1992); in fall (i.e. Q1) O. saurus was found, and

are known to prefer turbid waters when entering

estuaries (Fischer et al. 1995). Syngnathus sp. was

also present in Q1. Organisms in this genus are

associated with aquatic vegetation including sea-

grasses, noteworthy its congeneric (S. louisianae)

occurred in the two subsequent Qs and only at

HB. For summer (i.e. Q4) the presence of S. mac-

ulatus was recorded. They are known to be mov-

ing along the Mexican coast (which is very nearby

the study sites) between August and November

(Froese and Pauly 2022).

The three fishes that only occurred at SB were

present in a single Q: Bairdiella chrysoura (Q1),

Lucania parva (Q3), and Orthopristis

chrysoptera (Q4). The former species likely was

using the enclosed system as a nursery and/or

feeding area, as they are known to move into

estuaries during the summer months (Froese and

Pauly 2022); while L. parva benefitted from the

presence of seagrasses, which are denser than at

158 MARINE AND FISHERY SCIENCES 36 (2): 149-163 (2023)

HB but were not quantified for the purposes of

this study. Both species might have been present

but not captured in other Qs. This possibility is

less likely for O. chrysoptera which are mainly

nocturnal and apparently avoid low water temper-

atures as they seasonally migrate to deeper waters

during winter (Darcy 1983). While for B.

chrysoura, the possibility is very feasible as it is

known not only to feed but also to mature and

reproduce within estuaries, and hence considered

an estuarine resident (Grammer et al. 2009).

Lucania parva is also categorized as an estuarine

species, known to prefer areas with high seagrass-

es canopies that may provide protection from

predators and perhaps a larger amount of epiben-

thic prey items that attach to the seagrasses sur-

face areas (Tomoleoni 2007). Many studies have

shown that vegetated habitats tend to rear higher

densities of forage fishes and other organisms

than those that are non-vegetated sandy or muddy

bottoms (Summerson and Peterson 1984; Connol-

ly 1994; Jenkins et al. 1997; Rozas and Minello

1998; Sheridan and Minello 2003), but as the

goals of the present study were not to contrast

vegetated versus non-vegetated sites neither to

estimate seagrasses densities at the studied sites,

the subject is not further discussed.

Aside from the aforementioned factors associ-

ated with fishes movements (migration, disper-

sal), ontogenetic (reproduction, growth), survivor-

ship, or diel habits, one other cause for not captur-

ing the eight fishes (five in HB and three in SB) in

other Qs could be related to the fishing gear (i.e.

bag seine) used. Rozas and Minello (1997) sug-

gested that enclosure devices have a greater catch

efficiency of small nekton than towed nets in shal-

low estuarine habitats. However, see Layman and

Smith (2001) for a different perspective. Note-

worthy, gear was selected as part of the larger

study focused on the age, growth, diet, and trophic

web reconstruction of S. ocellatus and adopted

from the TPWD sampling protocol, which uses

bag seines as part of their survey efforts. As the

present results will serve as a baseline study to

compare outcomes to TPWD presented else-

where, it was necessary to sample with this gear.

Nonetheless, even with the inherent bag seine

sampling bias and limitations, four species (A.

mitchilli, C. variegatus, L. rhomboides, and M.

undulatus) use both studied sites as their perma-

nent home by being present year-round. Three of

these fishes are generally euryhaline.

In terms of fishes’ relative abundances, 11,795

and 4,085 individuals were captured at HB and

SB, respectively. Temporally, the greater abun-

dances occurred in Q3 at both sites, then changed

depending on location with the 2nd largest abun-

dances at HB during Q2 and Q4 at SB. Thus, the

spring (i.e. Q3) seems to be the more beneficial

season at both sites but as this is a single-year

study such a statement may not hold when multi-

ple years are accounted for. Nonetheless, for the

purposes of the results here presented, 47.7% and

34.4% of the fishes’ abundances occurred at HB

and SB respectively during the spring. Lagodon

rhomboides was mainly responsible for this pat-

tern with 4,271 individuals at HB and 1,267 at

SB. Noteworthy, the species was least abundant

during Q2 (~winter), and as mentioned before

occurred year-round at both sites. Hence, L.

rhomboides had higher abundances during

spring-summer, which was also reported for the

Mad Island estuary in Texas (Akin et al. 2003),

and in Tampa Bay, Florida (Chacin et al. 2016).

Lagodon rhomboides post-larvae arrive in the

GOM and southeastern USA estuaries during

winter as they reproduce offshore (Darcy 1985),

hence the higher abundances during spring to fall

(i.e. ca. Qs 3, 4, and 1) of one of the most widely

distributed and common fishes in the USA

(Hoese and Moore 1998), which follows such

ontogenetic cycle. This also explains its lesser

abundances during winter (Q2). Furthermore,

Stoner (1980) indicated L. rhomboides as impor-

tant mesograzers in the Laguna Madre seagrass

beds being abundant from early spring to fall,

while Hoss (1974) observed a minimal overwin-

tering of the L. rhomboides population in North

Carolina’s estuarine waters as the majority of the

fish migrated offshore upon the onset of colder

water temperatures.

Fish assemblages similarities formed two clus-

ters (not shown), but there was no significant dif-

ference between them. Nonetheless, the group

formed in Q2 for both sites with Q3 at HB in the

nMDS is likely the result of greater variability in

the abundances of taxa, with the two higher abun-

dances at HB and the least abundance at SB. This

exchange of abundance values among seasons is

complementary to the more abundant fishes

which might have contributed to the lack of sig-

nificant difference found and helps to also

explain the interaction effect in abundances and

diversity (discussed above) resulting in the gener-

ally observed equitability. However, winter con-

ditions should not be disregarded and likely

allowed some degree of grouping of both sites

during Q2.

Although DRs were variable throughout the

year and between sites, L. rhomboides was over-

whelmingly more abundant from spring to fall at

both sites, while M. undulatus was more domi-

nant in winter (Q2) with a DR of 0.37 at SB and

0.39 at HB. As mentioned in the results, the 2nd

and 3rd positions based on DRs varied among

species and might have resulted from resource

partition throughout the year. Likely migration

offshore for reproduction purposes, decreasing

temperatures, or both factors combined, L. rhom-

boides (see above) allowed the rise of another

year-round resident to become dominant in Q2.

Akin et al. (2003) also found seasonal changes in

estuarine fishes assemblages in Mad Island

Marsh, Matagorda Bay, Texas with M. undulatus

being more abundant from December-April when

temperatures were low, while L. rhomboides was

abundant during spring and summer. Resource

partitioning could also be exemplified by A.

mitchilli and L. rhomboides, regarding the DRs of

the former species tended to be the 2nd most

abundant in half of the Qs at SB. Although habitat

zonation was not the focus of this study, Gelwick

159

CAMARILLO JR. ET AL.: FORAGE FISHES IN SOUTHERNMOST TEXAS

et al. (2001) identified zones based on depth,

salinity, and dissolved oxygen gradients also in

Matagorda Bay and indicated A. mitchilli occu-

pied a different zone than L. rhomboides.

Lastly, six of the eight more dominant species

are important for fisheries (A. mitchilli, C. varie-

gatus, L. rhomboides, M. undulatus, M. cephalus,

and S. ocellatus) and were present year-round

highlighting the importance of both study sites as

important in their life cycle, although M.

cephalus, and S. ocellatus were not always cap-

tured by the bag seines. According to Froese and

Pauly (2022) these species as fishery resources

have the following uses: 1) major relevance as

bait A. mitchilli, C. variegatus and L. rhom-

boides; 2) considered game fishes L. rhomboides,

M. undulatus, M. cephalus and S. ocellatus; 3)

commercial M. undulatus and M. cephalus, while

L. rhomboides and S. ocellatus are used in minor

extent; 4) aquarium fishes’ exploitation C. varie-

gatus and S. ocellatus; and 5) used in aquaculture

M. undulatus, M. cephalus, and S. ocellatus.

Although the present study only encompassed

one-year, it was evident that both sites are inter-

connected as no fishes similarities difference was

found, nonetheless their multiple differences (i.e.

basin area, seagrasses coverage, connection to the

GOM, etc.) resulted in significant differences in

fishes richness, abundances, H’, and J’ suggesting

differences in habitat quality, or at the very least

variation in the availability of habitat types,

which are known to contribute to differences in

fish diversity attributes (Meffe and Berra 1988).

It also highlights the importance of HB and SB as

a permanent residency for several species that

were present year-round, including six fishes rel-

evant to fisheries. This calls for further studies in

this relatively abandoned area (compared to other

USA GOM ones), particularly as global warming

and increasing sea-level are expected to impact

the coastal areas likely impacting not only

resources useful to feed the increasing human

population but also affecting forage fishes which

should be managed at the very least as a group

because of their relevance as an energy source

within any marine food web.

ACKNOWLEDGEMENTS

The authors thank all the members of the

Ocean, Coastal Environmental and Ecological

Assessment Laboratory at UTRGV that partici-

pated in the field surveys, as well as boat captain

Skye Zufelt for skillful navigation to sampling

sites. Thanks also to the two anonymous review-

ers for their feedback allowing improvement of

this manuscript. All fishes were collected under

the Texas Parks and Wildlife Department Scien-

tific Permit (SPR-0808-314), in compliance with

UTRGV’s Institutional Animal Care and Use

Committee (IACUC) approved protocol (AUP-

19-40). Funding for the broader study from which

data presented and analyzed here is greatly

acknowledged and was provided by the National

Oceanic and Atmospheric Administration

(NOAA), Office of Education Educational Part-

nership Program (EPP) through the Center for

Coastal and Marine Ecosystems award

(NA16SEC4810009). However, publication con-

tents are solely the responsibility of the authors

and award recipient and do not necessarily repre-

sent the official views of the U.S. Department of

Commerce, NOAA.

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