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Larval Duration, Settlement, and Larval Growth Rates of the
Endangered Tidewater Goby (Eucyclogobius newberryi) and the Arrow
Goby (Clevelandia ios) (Pisces, Teleostei)
Author(s): Brenton T. Spies, Berenice C. Tarango, and Mark A. Steele
Source: Bulletin, Southern California Academy of Sciences, 113(3):165-175.
Published By: Southern California Academy of Sciences
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Larval Duration, Settlement, and Larval Growth Rates of the
Endangered Tidewater Goby (Eucyclogobius newberryi) and the
Arrow Goby (Clevelandia ios) (Pisces, Teleostei)
Brenton T. Spies, Berenice C. Tarango, and Mark A. Steele
Department of Biology, California State University, Northridge, CA 91330-8303
Abstract.—The early life history of the federally endangered Tidewater Goby
(Eucyclogobius newberryi) and its sister species the Arrow Goby (Clevelandia ios) has
been poorly documented to date. Both are endemic to estuarine habitats throughout
the California coast, however, habitat use differs between these two species. The
Arrow Goby is commonly found in fully marine tidal bays and mudflats. The
Tidewater Goby, however, prefers lagoons with some degree of seasonal isolation
from the sea. Here, we used otoliths to examine the larval duration, size at
settlement, and growth rates of newly settled gobies collected from 18 estuaries in
California. The Tidewater Goby had a larval duration that was ,2 days shorter than
the Arrow Goby (23.95 vs. 26.11 days, respectively), but a larger size at settlement
based on back-calculated size (12.38 vs. 10.00 mm SL) due to a faster larval growth
rate (2.86 vs. 2.60 mm/day
). There are several reasons that could explain these
differences in larval traits, such as differences in temperature or food resources
between the two estuary types, or the faster, annual life cycle of the Tidewater Goby
relative to the Arrow Goby.
Many fish species inhabit estuarine habitats during their early life history. These
habitats are quite variable, with some remaining open to the ocean year round and others
typically closing seasonally. Species that inhabit seasonally closing estuaries may exhibit
different larval traits than those that inhabit permanently open estuaries, and these traits
may contribute to genetic isolation by limiting dispersal (Bilton et al. 2002, Watts and
Johnson 2004). For example, Dawson et al. (2002) found genetic evidence of limited
dispersal in the Tidewater Goby (Eucyclogobius newberryi), which inhabits seasonally
closed estuaries, whereas its sister species the Arrow Goby (Clevelandia ios), which
inhabits open systems, lacked regional genetic divergence. This difference is likely due to
greater marine larval dispersal and gene flow between populations of arrow gobies.
Larval duration has long been thought to influence dispersal, and differences in larval
duration could confound this interpretation.
Closed estuaries experience greater seasonal variation in environmental conditions
than is seen in estuaries perennially open to marine influence, and such variation is
known to significantly influence larval development (McCormick and Molony 1995,
Green and Fisher 2004). Along the California coast, many estuaries are partially or
completely isolated from tidal influence either seasonally or episodically (Jacobs et al.
2011). Opening or ‘‘breaching’’ is usually a function of streamflow (Rich and Keller
2013), which is driven by seasonal precipitation. Isolation, or closure, occurs when a sand
bar or raised beach berm impounds systems during periods of lowered ‘‘summer’’
streamflow, creating variable salinity. Such dynamic lagoonal systems are a product of
Bull. Southern California Acad. Sci.
113(3), 2014, pp. 165–175
ESouthern California Academy of Sciences, 2014
the Mediterranean climate that characterizes California. Similar lagoonal processes are
known to occur in other regions of the globe with similar climate (Jacobs et al. 2011).
The federally endangered Tidewater Goby is a California endemic that occurs in
estuaries that experience seasonal or episodic closure (Swift et al. 1989). The Tidewater
Goby is a small benthic fish that seldom exceeds 55 mm standard length (SL) (Miller and
Lea 1972). Lagoons with seasonally closing stream mouths on the outer coast are the
typical Tidewater Goby habitat, although they also occupy or historically occupied,
naturally closing, tide-gated, or marginal pond habitats around Humboldt, Tomales, and
San Francisco Bays (Swift et al. 1989, Moyle 2002, U.S. Fish and Wildlife Service 2005).
This species exhibits local genetic divergence at a finer geographic scale than any other
Pacific coast vertebrate (Barlow 2002, Dawson et al. 2002, Earl et al. 2010). Its preference
for small and isolated estuaries is one of the main reasons why it is predisposed to local
extirpation (Lafferty et al. 1999a,b).
Dispersal of the Tidewater Goby is associated with high streamflow events (Lafferty
et al. 1999a,b), which cause breaching of the estuary mouth, permitting dispersal (Earl et
al. 2010). Breaching events occur most frequently during winter months when
reproduction is limited and larvae are generally absent. As confirmed by genetic
divergence, marine larval dispersal appears to be extremely limited, if it occurs at all
(Barlow 2002, Dawson et al. 2002, Earl et al. 2010). This conclusion is supported by the
work of Hellmair and Kinziger (2014), which showed that small tidewater gobies
(,25 mm total length) experience higher mortality rates than larger individuals (.25 mm)
when salinity was raised from 6%to 26%. Presumably this intolerance to seawater in
young tidewater gobies is related to the isolated nature of their lagoonal habitats during
the summer peak reproductive months (Swenson 1999). Thus, dispersal appears limited
to adult movement over sandy substrate following breaching events (Earl et al. 2010).
The Arrow Goby is the sister species to the Tidewater Goby (Dawson et al. 2002), and
it ranges from Bahia Magdalena, Baja California Sur (C. Swift, pers. comm.) to British
Colombia (Miller and Lea 1972). Similar to the Tidewater Goby, the Arrow Goby is a
small benthic fish that rarely exceeds 60 mm SL (Miller and Lea 1972, Hart 1974). It
prefers more open, fully tidal estuaries and mudflats, which are typically cooler and
higher in salinity. This preference for open estuaries is thought to facilitate marine larval
Here, we compare the early life history of the Arrow Goby and the Tidewater Goby
collected from eighteen California estuaries. Larval traits were determined from daily
bands and settlement marks in lapillar otoliths of recently settled gobies. Larval duration,
size at settlement, and pre-settlement growth rates were compared between the two
Materials and Methods
Gobies were collected from eighteen estuaries in California (Fig. 1). Sites were chosen
based on the presence of healthy and abundant populations of the Arrow Goby or
Tidewater Goby, in addition to their mouth dynamics (closed vs. open estuary mouth).
The Tidewater Goby was collected at ten seasonally closing estuaries: Ten Mile River
(39u32943.860N, 123u45925.040W); Salmon Creek (38u21910.870N, 123u03957.190W);
Rodeo Lagoon (37u49954.410N, 122u31943.310W); San Gregorio (37u19914.290N,
122u24903.380W); Moore Creek (36u5794.500N, 122u03929.850W); San Luis Obispo Creek
(35u11913.350N, 120u43933.470W); Santa Ynez River (34u41930.570N, 120u35900.700W);
166 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
Arroyo Burro Lagoon (34u24911.770N, 119u44935.120W); Santa Clara River (34u14907.190N,
119u15927.460W); and Las Flores Marsh (33u17925.790N, 117u27953.910W). The Arrow Goby
was collected at eight estuaries that are typically fully tidal: Arcata Bay (40u51930.570N,
124u06900.080W); Bodega Bay (38u18959.420N, 123u02943.120W); San Lorenzo River
(36u57956.410N, 122u00945.460W); Elkhorn Slough (36u48940.140N, 121u44938.770W); Morro
Bay (36u57956.410N, 122u00945.460W); Carpinteria Salt Marsh (34u23952.970N,
119u32916.720W); Colorado Lagoon (33u45910.520N, 118u07947.370W); and Los Pen˜ asquitos
(32u55957.840N, 117u15929.110W). Due to the differences in habitat preference of the Arrow
Goby and Tidewater Goby, none of the eighteen study sites had both species present at the
time of collection.
Larval, transitional, and recently settled gobies (Fig. 2) were collected between August
and October of 2011. Both species were collected using a 3.7 x 1.2 m beach seine with a
1.6-mm mesh, and in some cases, a one-man push net with 1.6-mm mesh (Strawn 1954).
The Arrow Goby was collected at low tide in all eight study sites. The ten study sites
where the Tidewater Goby was collected were all completely closed to marine tidal
influence at the time of collection. Once collected, the fishes were euthanized and
preserved in 95%ethanol.
Fig. 1. Map of the 18 study sites located along the California coastline. The Arrow Goby (squares) was
collected at 8 sites, and the Tidewater Goby (circles) was collected at 10 sites.
LARVAL TRAITS OF TWO CALIFORNIA GOBIES 167
Otoliths were used to measure larval traits of the study species. Otoliths have been used
for these purposes in a wide variety of fishes, including many species in the family
Gobiidae (Sponaugle and Cowen 1994, Hernaman et al. 2000, Radtke et al. 2001,
Yamasaki and Maeda 2007, Wilson et al. 2009, Samhouri et al. 2009). Both the sagittal
and lapillar otoliths were extracted from all individuals using standard techniques
(Brothers 1987, Hellmair 2010) and placed in immersion oil for .30 days to clear
(Samhouri et al. 2009). For both species, lapilli were used because they were clearer and
easier to interpret than sagittae, and they did not require sectioning or polishing. Lapilli
were read whole from images on a computer monitor that were captured by a digital
camera mounted on a compound microscope at 2003magnification, with a polarizing
filter placed between the light source and the first stage. Increment measurements were
made along the longest axis, from the core to the outermost complete ring using Image-
Pro Plus image analysis software.
Larval duration, size at settlement, and growth rates were estimated from the otoliths.
Previous work has validated daily otolith increment deposition in the Tidewater Goby
(Hellmair 2010), and increments were assumed to be daily in the Arrow Goby. Settlement
was recorded in the otolith structure as a distinct transition in increment widths (Fig. 3),
as noted in other gobies by Sponaugle and Cowen (1994). Larval duration was
determined from a count of the rings from the hatch mark (first band from otolith core)
to the settlement mark. Average pre-settlement growth rates were estimated as average
Fig. 2. Photos showing the rapid development of the Tidewater Goby, exhibiting the transition from
(A) a 20-day-old postflexion larva (9.3 mm SL), to (B) a 25-day-old transitional juvenile captured prior to
settlement (10.8 mm), and (C) a 34-day-old settled juvenile (13.7 mm).
168 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
increment width between the hatch mark and the settlement mark (McCormick and
Molony 1995). All otoliths were read twice by one person (B.T. Spies), and all unclear
and abnormally shaped otoliths were discarded. If the two readings were more then 10%
different, then that otolith was not included in any analysis. If the two readings were less
than 10%different but not the same, then the second reading was used for the analysis.
To validate that the presumed settlement mark actually corresponded to settlement, the
number of presumed post-settlement bands was regressed against body length for both
species. The x-intercept of the linear regression equation estimates size at settlement,
which was compared to the size of fish known to have recently settled based on their
morphology. Further regression analyses examined the relationship between body length
and age (days) to determine whether otolith measurements provide accurate proxies of
somatic traits (Booth and Parkinson 2011). Back calculation was used to estimate body
size at settlement for each fish using the equation calculated by ordinary least square
linear regression of body size (mm SL) on otolith radius, of the form L 5mx +b; where L
represents the body length, m represents the slope, x represents the otolith radius at
settlement, and b represents the y-intercept. A nested ANOVA with the factors Species
(fixed effect) and Site nested within species (random effect) was used to test whether
larval traits differed between species and collection sites.
Otolith-based estimates of larval traits appeared to be appropriate for both the
Tidewater Goby and Arrow Goby due to the fact that body length was tightly related to
age as estimated from otoliths (Arrow Goby: r
50.75, n5317; Tidewater Goby: r
n5406; Fig. 4). The number of post-settlement otolith bands and body length were also
tightly related (Arrow Goby: r
50.79; Tidewater Goby: r
50.73) and the fitted lines
predicted realistic sizes at settlement (Fig. 5). This result indicates that the distinguishable
transition zone in the otolith is the settlement band (Fig. 3A), and that all bands between
the core and the settlement mark indicate the larval duration (Sponaugle and Cowen
Fig. 3. Lapillar otolith of a juvenile Tidewater Goby (A), and a larval Tidewater Goby (B) with visible
daily bands viewed at 200 x magnification. The black arrow indicates the settlement band. The white line
indicates pre-settlement bands, and the black line represents post-settlement bands.
LARVAL TRAITS OF TWO CALIFORNIA GOBIES 169
1994). Otolith radius was a good proxy for body length, as indicated by a strong linear
relationship between body length and otolith radius (Arrow Goby: r
50.78; Fig. 6). Based on back-calculation, mean 6SD size at settlement was
11.45 60.23 mm SL for the Tidewater Goby and 9.32 60.63 mm for the Arrow Goby,
slightly smaller than the estimates of size at settlement based on the x-intercept of the
regressions of post-settlement age on size (Fig. 5, Table 1).
The Arrow Goby had a significantly longer larval duration (F
larger otolith radius at settlement (F
528.4, p,0.0001), and slower larval growth rates
5399.8, p,0.0001) than the Tidewater Goby (Table 1). Size at settlement (back
calculated) of the Tidewater Goby was significantly larger than that of the Arrow Goby
55360.1, p,0.0001). All larval traits varied significantly among sites for both species
(larval duration: F
542.5, p,0.0001; otolith radius at settlement: F
Fig. 4. Relationship between body length and age for the Arrow Goby (A) and Tidewater Goby (B).
170 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
p,0.0001; pre-settlement growth rate: F
533.53, p,0.0001; back calculated body size
at settlement: F
Analysis of lapillar otoliths revealed that larval duration, size at settlement, and pre-
settlement growth rates of the two sister species, the Arrow Goby and Tidewater Goby,
were statistically different, though not dramatically different. The similarity between the
two species in larval duration is somewhat surprising given known differences in their
habitat usage and methods of dispersal. Despite inhabiting two different kinds of
estuaries, ‘‘open’’ versus ‘‘closed’’, and dispersing as larvae (Arrow Goby) versus adults
(Tidewater Goby), the duration of the larval phase was only 8%shorter in the Tidewater
Goby. Due to faster larval growth rates, the Tidewater Goby settled at larger size than
Fig. 5. Relationship between post-settlement age and body length for the Arrow Goby (A) and
Tidewater Goby (B).
LARVAL TRAITS OF TWO CALIFORNIA GOBIES 171
Fig. 6. Relationship between otolith radius and body length for the Arrow Goby (A) and Tidewater
Table 1. Estimated larval durations, settlement sizes (standard length), and growth rates of the Arrow
Goby (Clevelandia ios) and Tidewater Goby (Eucyclogobius newberryi) pooled over all study sites (n58
and 10 study sites for the two species, respectively).
Larval duration Settlement Growth rate
Species Range (days)
(mm) Length (mm)
E. newberryi 18 - 31 23.95 62.71 75.62 66.40 12.38 60.36 2.86 60.23 406
C. ios 20 - 33 26.11 62.40 77.61 66.37 10.00 60.71 2.60 60.22 317
172 SOUTHERN CALIFORNIA ACADEMY OF SCIENCES
did the Arrow Goby. It is likely that the Tidewater Goby has evolved the ability to settle
and reach maturity faster than the Arrow Goby due to its shorter, annual life cycle.
Larval durations of other related goby species (e.g. Gillichthys mirabilis,Ilypnus
gilberti,Lepidogobius lepidus,Quietula y-cauda), commonly referred to as the ‘‘bay
gobies’’ (Teleostei: Gobionellidae) (Thacker 2009, Ellingson et al. 2014) have not yet been
determined. In fact, detailed knowledge of early life history characteristics does not exist
for many native fishes that occupy estuaries and wetlands in California. Larval traits have
been studied more regularly in tropical gobies (Teleostei: Gobiidae), which on average,
tend to exhibit more prolonged larval stages than those of the species in this study.
Yamasaki and Maeda (2007) documented a 78-146 day larval duration in Stiphodon
percnopterygionus, a stream-associated goby found in Okinawa Island, southern Japan.
Lentipes concolor, an endemic Hawaiian amphidromous goby was found to have a larval
duration between 63-106 days (Radtke et al. 2001). The goldspot goby (Gnatholepis
thompsoni), commonly found on coral reefs in the Caribbean, was found to have a larval
duration between 45-80 days (Samhouri et al. 2009), with the potential of reaching
112 days (Sponaugle and Cowen 1994). Another common goby in the Caribbean
(Coryphopterus glaucofraenum), however, had a much shorter larval duration of 27 days
(Sponaugle and Cowen 1994), similar to that found in this study for the Arrow Goby.
Previous estimates of larval characteristics of these two study species have varied
widely, presumably due to methodological limitations. Estimates of larval duration for
the Tidewater Goby have ranged from as little as several days (Capelli 1997) to a few
weeks (Dawson et al. 2002). Past research on the Arrow Goby has provided a broad
range of estimates of larval duration using both field collected and laboratory-reared
specimens. However, none of these estimates were obtained from otolith analysis.
Brothers (1975) estimated larval duration at approximately 30 days from the timing of
annual reproduction and recruitment. Dawson et al. (2002) estimated a 2–4 week larval
duration by extrapolating the time to reach 7 mm (10 days; Hart 1973) to the time to
reach the average size at settlement of 13.1 61.3 mm (range 10.0–16.6 mm) observed by
Kent and Marliave (1997) for newly settled arrow gobies from British Colombia. Our
back-calculated estimate of size at settlement (10.00 60.6 mm) falls on the lower end of
Kent and Marliave’s (1997) range of size at settlement, but overlaps with measurements
of nearshore postlarvae collected by Brothers (1975). Morphological descriptions of
transitional juveniles (12.5 mm SL) from British Colombia are consistent with the
morphology of juveniles in southern California estuaries that settled at a smaller size,
perhaps indicating regional differences in size at settlement. It is possible, however, that
some of these apparent differences in size at settlement are related to how fish were
preserved in various studies, which can cause them to shrink (Hay 1982).
The larval traits investigated in this study varied among populations, possibly due to
differences among estuaries in environmental conditions, such as temperature, water
quality, or food resources. This could be one explanation for the different estimates of
settlement size from this study, done in southern California, and that of Kent and
Marliave (1997) conducted in British Colombia. Brothers (1975) found that the Arrow
Goby lives for approximately one year in southern California, similar to the annual
lifespan of the Tidewater Goby throughout its range, but has an extended lifespan in the
more northern portions of its range (2-3 years). Swenson (1999) found that the average
size of the Tidewater Goby was significantly larger in marsh habitats than in lagoons or
creeks. She speculated that this was due to the more stable physical conditions of the
marsh, which fosters improved growth, perhaps due to a more consistent or abundant
LARVAL TRAITS OF TWO CALIFORNIA GOBIES 173
supply of prey. It is also worth noting that, given genetic isolation of populations (e.g.
Earl et al. 2010), adaptation to local conditions is also possible in the Tidewater Goby.
Our findings provide baseline estimates of some of the early life history traits of the
Tidewater Goby and the Arrow Goby, but these attributes are likely to vary with
Thanks to Mike Rouse, Rhys Evans, Darren Fong, Kevin Lafferty, and Rikke Kvist
Preisle for their help gaining access to study sites, collections, and project logistics.
Thanks to Larry Allen, Camm Swift, David Jacobs, and Steve Dudgeon for their wealth
of knowledge and project guidance. We appreciate the help of Aaron Dufault in
collecting fish, and Brian Pen˜ a and Rando Has for their long hours spent in the lab
dissecting fish and preparing otoliths. This project was supported by the Department of
Biology at California State University, Northridge, and the International Women’s
Fishing Association (IWFA). Thanks to Chris Dellith and U.S. Fish and Wildlife Service
for their help, guidance, and permit support. This research was conducted under the
permitted support of California State Parks, National Parks Service, U.S. Fish and
Wildlife Service (recovery permit #TE-43944A-0), and the CA Department of Fish and
Wildlife (permit #SC-10750).
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