ArticlePDF Available

Larval Duration, Settlement, and Larval Growth Rates of the Endangered Tidewater Goby ( Eucyclogobius newberryi ) and the Arrow Goby ( Clevelandia ios ) (Pisces, Teleostei)


Abstract and Figures

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 µm/day−1). 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.
Content may be subject to copyright.
BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic
institutions, research libraries, and research funders in the common goal of maximizing access to critical research.
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
BioOne ( is a nonprofit, online aggregation of core research in the biological,
ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170
journals and books published by nonprofit societies, associations, museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your
acceptance of BioOne’s Terms of Use, available at
Usage of BioOne content is strictly limited to personal, educational, and non-commercial use.
Commercial inquiries or rights and permissions requests should be directed to the individual publisher
as copyright holder.
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
Study Sites
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);
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.
Collection Methods
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.
Otolith Analysis
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).
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.
Data 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.
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
50.80; Tidewater
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
5227.3, p,0.0001),
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).
p,0.0001; pre-settlement growth rate: F
533.53, p,0.0001; back calculated body size
at settlement: F
525.9, p,0.0001).
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).
Fig. 6. Relationship between otolith radius and body length for the Arrow Goby (A) and Tidewater
Goby (B).
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)
Mean 6SD
Otolith radius
(mm) Length (mm)
Mean 6SD
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
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
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
environmental parameters.
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).
Literature Cited
Barlow, M. 2002. Phylogeographic structure of the Tidewater Goby, Eucyclogobius newberryi (Teleostei:
Gobiidae), in the San Francisco Bay area and Ventura County: implications for conservation
management. Masters Thesis. University of California, Los Angeles.
Bilton, D. T., J. Paula, and J. D. D. Bishop. 2002. Dispersal, genetic differentiation and speciation in
estuarine organisms. Estuarine, Coastal and Shelf Science, 55:937–952.
Booth, D. J. and K. Parkinson. 2011. Pelagic larval duration is similar across 23uof latitude for two
species of butterflyfish (Chaetodontidae) in eastern Australia. Coral Reefs, 30:1071–1075.
Brothers, E. B. 1975. The comparative ecology and behavior of three sympatric California gobies. Ph.D.
Dissertation. University of California, San Diego.
———. 1987. Methodological approaches to the examination of otoliths in aging studies. Pp. 319–330 in
Summerfeldt R. C., and G. E. Hall, eds., The Age and Growth of Fish. Iowa State University Press,
Des Moines.
Capelli, M. H. 1997. Tidewater Goby (Eucyclogobius newberryi) management in California estuaries.
Proceedings, California and the World Ocean Conference, San Diego, American Society of Civil
Engineers, pp. 1247–1264.
Dawson, M. N., K. D. Louie, M. Barlow, D. K. Jacobs, and C. C. Swift. 2002. Comparative
phylogeography of sympatric sister species, Clevelandia ios and Eucyclogobius newberryi (Teleostei,
Gobiidae), across the California Transition Zone. Molecular Ecology, 11:1065–1075.
Earl, D.A., K. D. Louie, C. Bardeleben, C. C. Swift, and D. K. Jacobs. 2010. Rangewide microsatellite
phylogeography of the endangered Tidewater Goby, Eucyclogobius newberryi (Teleostei:
Gobiidae), a genetically subdivided coastal fish with limited marine dispersal. Conservation
Genetics, 11:103–114.
Ellingson, R. A., C. C. Swift, L. T. Findley, and D. K. Jacobs. 2014. Convergent evolution of
ecomorphological adaptations in geographically isolated Bay gobies (Teleostei: Gobionellidae) of
the temperate North Pacific. Molecular Phylogenetics and Evolution, 70:464–477.
Green, B. S. and R. Fisher. 2004. Temperature influences swimming speed, growth and larval duration in
coral reef fish larvae. Journal of Experimental Marine Biology and Ecology, 299:115–132.
Hart, J. L. 1974. Pacific Fishes of Canada. Bulletin of the Fisheries Research Board of Canada, 740 pp.
Hay, D. E. 1982. Fixation shrinkage of herring larvae: effects of salinity, formalin concentration, and
other factors. Canadian Journal of Fisheries and Aquatic Sciences, 39:1138–1143.
Hellmair, M. 2010. Manual for Otolith Based Age Determination of Tidewater Goby, Eucyclogobius
newberryi. Scientific Report. Humboldt State University, 22 pp.
——— and A. P. Kinziger. 2014. Increased extinction potential of insular fish populations with reduced
life history variation and low genetic diversity. PLoS ONE, in press.
Hernaman, V., P. L. Munday, and M. L. Schla¨ppy. 2000. Validation of otolith growth-increment
periodicity in tropical gobies. Marine Biology, 137:715–726.
Jacobs, D. K., E. D. Stein, and T. Longcore. 2011. Classification of California estuaries based on natural
closure patterns: templates for restoration and management. Southern California Coastal Waters
Research Project, 619.a, 72 pp.
Kent, D. I. and J. B. Marliave. 1997. Early life history of the Arrow Goby, Clevelandia ios (Jordan &
Gilbert), Gobiidae. Micronesica, 30:15–23.
Lafferty, K. D., C. C. Swift, and R. F. Ambrose. 1999a. Postflood persistence and recolonization of
endangered Tidewater Goby populations. North American Journal Fisheries Management, 19:
———, ———, and ———. 1999b. Extirpation and recolonization in a metapopulation of an endangered
fish, the Tidewater Goby. Conservation Biology, 13:1447–1453.
McCormick, M. I. and B. W. Molony. 1995. Influence of water temperature during the larval stage on size,
age and body condition of a tropical reef fish at settlement. Marine Ecology Progress Series, 118:
Miller, D. J. and R. N. Lea. 1972. Guide to the Coastal Marine Fishes of California. California
Department of Fish and Game. Fish Bulletin 157, 235 pp.
Moyle, P. B. 2002. Inland Fishes of California. University of California Press, Berkeley. 502 pp.
Radtke, R. L., R. A. Kinzie, and D. J. Shafer. 2001. Temporal and spatial variation in length of larval life
and size at settlement of the Hawaiian amphidromous goby Lentipes concolor. Journal of Fish
Biology, 59:928–938.
Rich, A. and E. A. Keller. 2013. A hydrologic and geomorphic model of estuary breaching and closure.
Geomorphology, 191:64–74.
Samhouri, J. F., M. A. Steele, and G. E. Forrester. 2009. Inter-cohort competition drives density
dependence and selective mortality in a marine fish. Ecology, 90:1009–1020.
Sponaugle, S. and R. K. Cowen. 1994. Larval durations and recruitment patterns of two Caribbean gobies
(Gobiidae): contrasting early life histories in demersal spawners. Marine Biology, 120:133–143.
Strawn, K. 1954. The pushnet, a one-man net for collecting in attached vegetation. Copeia, 1954:195–197.
Swenson, R. O. 1999The ecology, behavior, and conservation of the Tidewater Goby, Eucyclogobius
newberryi. Environmental Biology of Fishes, 55:99–114.
Swift, C. C., J. L. Nelson, C. Maslow, and T. Stein. 1989. Biology and distribution of the Tidewater Goby,
Eucyclogobius newberryi (Pisces: Gobiidae) of California. Natural History Museum of Los Angeles
County, Contributions in Science, No 404, 19 pp.
Thacker, C. 2009. Phylogeny of Gobioidei and placement within Acanthomorpha, with a new
classification and investigation of diversification and character evolution. Copeia, 2009:93–104.
U.S. Fish and Wildlife Service. 2005. Recovery plan for the Tidewater Goby (Eucyclogobius newberryi).
U.S. Fish and Wildlife Service, Portland Oregon, vi +199 pp.
Watts, R. J. and M. S. Johnson. 2004. Estuaries, lagoons and enclosed embayments: habitats that enhance
population subdivision of inshore fishes. Marine and Freshwater Research, 55:641–651.
Wilson, J. A., L. Vigliola, and M. G. Meekan. 2009. The back-calculation of size and growth from
otoliths: Validation and comparison of models at an individual level. Journal of Experimental
Marine Biology and Ecology, 368:9–21.
Yamasaki, N. and K. Maeda. 2007. Pelagic larval duration and morphology at recruitment of Stiphodon
percnopterygionus (Gobiidae: Sicydiinae). The Raffles Bulletin of Zoology, 14:209–214.
... al 2014), it was assumed that increments in otoliths of the arrow goby were also deposited daily. Furthermore, otolith-based estimates of larval traits appear to be suitable for both species due to a strong relationship between body length and age (Spies et al. 2014). ...
... The settlement band was interpreted as the point where the daily increment transitioned distinctly in band width (Wilson & McCormick 1999, Spies et al. 2014). Date of settlement was found by subtracting the number of post-settlement increments counted for each otolith from the date the fish was collected. ...
... The first dark ring outside the core was considered to be the hatch mark (Hellmair 2010), although this interpretation has not been validated. Average pre-settlement growth rates (larval growth rate) were estimated from otolith growth as a proxy for larval somatic growth, by dividing the otolith radius to the settlement mark by the age at settlement (McCormick & Molony 1995, Spies et al. 2014). ...
Full-text available
Variations in abiotic conditions across large latitudinal gradients can strongly influence the early life history of coastal marine organisms. We investigated the effects of temperature and latitude on the larval traits of 2 estuarine fish species. The arrow goby Clevelandia ios and the endangered tidewater goby Eucyclogobius newberryi were studied in 18 estuaries along the coast of California, spanning ~8 degrees of latitude. These 2 species were selected because of their dissimilar preferences for estuary type: the arrow goby prefers cooler, fully tidal systems, whereas the tidewater goby prefers warmer lagoons that experience some degree of seasonal isolation from the ocean. Recently settled individuals were collected from July to October 2011, and temperatures within each estuary were recorded to determine how temperature affected larval duration, settlement, and growth rates. Temperatures were more variable among estuaries inhabited by the tidewater goby (10°C range) than among those inhabited by the arrow goby (5°C range). Larval traits of both species differed among sites, but more so for the tidewater goby, a difference that was tied to the greater differences in temperatures among sites in the seasonally closed estuaries it inhabited. In both species, fish that experienced warmer temperatures had shorter larval durations and faster growth rates and were smaller at settlement. Since the length of the larval period has been related to survival and dispersal distance, future variations in temperature due to climate change could have predictable influences on population density and connectivity in estuarine species.
... Tidewater Gobies grow to approximately 5 cm and reach 1 year of age while Prickly Sculpins grow to approximately 10 cm and reach 3 years of age. Both species have pelagic larvae and are omnivorous as juvenile and adults feeding primarily on a variety of micro-and macrocrustaceans and insects (Swenson and McCray, 1996;Moyle, 2002;Feyrer et al., 2003;Spies et al., 2014). ...
Full-text available
Bar-built coastal lagoons are dynamic ecosystems at the land-sea interface that are important habitats for a variety of species. This study examined the habitat ecology of two lagoon species, the endangered Tidewater Goby (Eucyclogobius newberryi) and the Prickly Sculpin (Cottus asper) by reconstructing individual life histories from patterns in the concentration of the element Sr (as ratioed to Ca; Sr:Ca) in otoliths. Specific objectives were to (1) elucidate any movements of individual fishes among three primary habitat components of typical bar-built lagoon systems: coastal ocean, brackish lagoon, and freshwater watershed streams, and (2) determine if either species exhibited a consistent life history as defined by a stereotypical otolith Sr:Ca chronology, which could be indicative of a consistent range of salinity or temperature occupied through ontogeny. Results suggested that Tidewater Goby was a lagoon resident and that Prickly Sculpin exhibited migrations between lagoon and watershed stream habitats. There was no strong evidence in either species of ocean occupancy or of a stereotypical Sr:Ca chronology, the latter suggesting the full range of available lagoon habitat in terms of salinity and temperature was likely utilized at all life stages. These findings add to the body of evidence that bar-built lagoons are not isolated habitats, and holistic management of these habitats with adjoining watershed and marine environments could increase habitat connectivity across the landscape, with potential benefits to fishes.
... Pigmentation in preserved material of larval and early juvenile fish is very similar in both species and was described in Swift et al. [16] and Watson [93] from material of E. kristinae and by Spies et al. [94] for E. newberryi from San Gregorio Creek lagoon, San Mateo County. Nine to eleven large stellate melanophores uniformly spaced mid-dorsally from nape to caudal base in late larvae and small juveniles form 7-9 dorsal saddles or blotches in larger juveniles and adults. ...
Full-text available
A geographically isolated set of southern localities of the formerly monotypic goby genus Eucyclogobius is known to be reciprocally monophyletic and substantially divergent in mito-chondrial sequence and nuclear microsatellite-based phylogenies relative to populations to the north along the California coast. To clarify taxonomic and conservation status, we conducted a suite of analyses on a comprehensive set of morphological counts and measures from across the range of Eucyclogobius and describe the southern populations as a new species, the Southern Tidewater Goby, Eucyclogobius kristinae, now separate from the Northern Tidewater Goby Eucyclogobius newberryi (Girard 1856). In addition to molecular distinction, adults of E. kristinae are diagnosed by: 1) loss of the anterior supratemporal lateral line canals resulting in higher neuromast counts, 2) lower pectoral and branched caudal ray counts, and 3) sets of measurements identified via discriminant analysis. These differences suggest ecological distinction of the two species. Previous studies estimated lineage separation at 2–4 million years ago, and mitochondrial sequence divergence exceeds that of other recognized fish species. Fish from Santa Monica Artesian Springs (Los Angeles County) northward belong to E. newberryi; those from Aliso Creek (Orange County) southward constitute E. kristinae. The lagoonal habitat of Eucyclogobius has been diminished or degraded, leading to special conservation status at state and federal levels beginning in 1980. Habitat of the newly described species has been impacted by a range of anthropo-genic activities, including the conversion of closing lagoons to open tidal systems in the name of restoration. In the last 30 years, E. kristinae has only been observed in nine intermittently occupied lagoonal systems in northern San Diego County; it currently persists in only three sites. Thus, the new species is in imminent danger of extinction and will require ongoing active management.
... Pigmentation in preserved material of larval and early juvenile fish is very similar in both species and was described in Swift et al. [16] and Watson [93] from material of E. kristinae and by Spies et al. [94] for E. newberryi from San Gregorio Creek lagoon, San Mateo County. Nine to eleven large stellate melanophores uniformly spaced mid-dorsally from nape to caudal base in late larvae and small juveniles form 7-9 dorsal saddles or blotches in larger juveniles and adults. ...
Conference Paper
The endangered closed-estuary specialist goby genus Eucyclogobius, the tidewater gobies, is the most locally-differentiated vertebrate taxon on the Pacific coast. It is subdivided into regional clades, which are further subdivided into long-isolated entities. Clades and subclades exhibit regionally distinct metapopulation processes. In addition, the southern most clade is deeply divergent with a separation in the excess of a million years ago. It is reciprocally monophyletic in nuclear and mitochrondrial markers and morphologically distinct in counted lateral line attributes, fin rays and measured characters as determined by discriminant function analysis as well as rate of development. This distinctive southern entity is under review as a new species. It is critically endangered in having been reduced to three small lagoonal populations on Camp Pendleton, northern San Diego County. Currently, efforts are underway to establish captive populations derived from these to reduce the risk of extinction associated with the vulnerability to drought and flood. Due to their metapopulation process and subdivison, the tidewater gobies are of exceptional scientific interest. We hope that both species n the group are available for study in the future. We will outline the events that led to range reduction and endangerment of the southern species, as well as steps taken to increase the likelihood of persistence.
Full-text available
Theoretical work has shown that reduced phenotypic heterogeneity leads to population instability and can increase extinction potential, yet few examples exist of natural populations that illustrate how varying levels expressed diversity may influence population persistence, particularly during periods of stochastic environmental fluctuation. In this study, we assess levels of expressed variation and genetic diversity among demographically independent populations of tidewater goby (Eucyclogobius newberryi), show that reductions in both factors typically coincide, and describe how low levels of diversity contribute to the extinction risk of these isolated populations. We illustrate that, for this annual species, continuous reproduction is a safeguard against reproductive failure by any one population segment, as natural, stochastically driven salinity increases frequently result in high mortality among juvenile individuals. Several study populations deviated from the natural pattern of year-round reproduction typical for the species, rendering those with severely truncated reproductive periods vulnerable to extinction in the event of environmental fluctuation. In contrast, demographically diverse populations are more likely to persist through such periods through the continuous presence of adults with broader physiological tolerance to abrupt salinity changes. Notably, we found a significant correlation between genetic diversity and demographic variation in the study populations, which could be the result of population stressors that restrict both of these diversity measures simultaneously, or suggestive of a causative relationship between these population characteristics. These findings demonstrate the importance of biocomplexity at the population level, and assert that the maintenance of diversity contributes to population resilience and conservation of this endangered species.
Full-text available
Before-and-after surveys at several southern California sites indicated that populations of endangered tidewater goby Eucyclogobius newberryi persisted through heavy flooding in 1995. This was contrary to our expectations that flooding might have led to extirpation in some smaller wetlands. There was also no significant change in tidewater goby density before and after the flooding. Several apparent recolonization events coincided with the flood, suggesting that flooding may be important for the long-term persistence of the species.
To better understand how the hydrology of bar-built estuaries affects breaching and closing patterns, a model is developed that incorporates an estuary hydrologic budget with a geomorphic model of the inlet system. Erosion of the inlet is caused by inlet flow, whereas the only morphologic effect of waves is the deposition of sand into the inlet. When calibrated, the model is able to reproduce the initial seasonal breaching, seasonal closure, intermittent closures and breaches, and the low-streamflow (closed state) estuary hydrology of the Carmel Lagoon, located in Central California. Model performance was tested against three separate years of water-level observations. When open during these years, the inlet was visually observed to drain directly across the beach berm, in accordance with model assumptions. The calibrated model predicts the observed 48-h estuary stage amplitude with root mean square errors of 0.45 m, 0.39 m and 0.42 m for the three separate years. For the calibrated model, the probability that the estuary inlet is closed decreases exponentially with increasing inflow (streamflow plus wave overtopping), decreasing 10-fold in probability as mean daily inflow increases from 0.2 to 1.0 m3/s. Seasonal patterns of inlet state reflect the seasonal pattern of streamflow, though wave overtopping may become the main hydrologic flux during low streamflow conditions, infrequently causing short-lived breaches. In a series of sensitivity analyses it is seen that the status of the inlet and storage of water are sensitive to factors that control the storage, transmission, and inflow of water. By varying individual components of the berm system and estuary storage, the amount of the time the estuary is open may increase by 57%, or decrease by 44%, compared to the amount of time the estuary is open during calibrated model conditions for the 18.2-year model period. The individual components tested are: berm height, width, length, and hydraulic conductivity; estuary hypsometry (storage to stage relationship); two factors that control wave-swash sedimentation of the inlet; and sea level rise. The elevation of the berm determines the volume of water that must enter the estuary in order to breach, and it modulates the wave-overtopping flux and frequency. By increasing estuary storage capacity, the estuary will breach less frequently (− 27% change in time open for modeled excavation scenario) and store water up to 3 months later into the summer. Altering beach aquifer hydraulic conductivity affects inlet state, and patterns of breaching and water storage. As a result of sea-level rise of 1.67 m by 2100, and a beach berm that remains in its current location and accretes vertically, the amount of time the estuary remains open may decrease by 44%. Such a change is an end-member of likely scenarios given that the berm will translate landwards. Model results indicate that the amount of time the estuary is open is more sensitive to changes in wave run-up than the amount of sand deposited in the inlet per each overtopping wave.
Duration of the pelagic phase of benthic marine fishes has been related to dispersal distance, with longer pelagic larval duration (PLD) expected to result in greater dispersal potential. Here, we examine PLDs of 2 species of coral-reef butterflyfish (Chaetodon auriga and C. flavirostris) across latitudes (14°S–37°S) along the Great Barrier Reef into south-eastern Australia; we predict that PLD will be higher for fish collected below the breeding latitudes of 24°S. For C. auriga, apart from significantly longer PLDs at Lord Howe Island and Jervis Bay (means of 54 and 52 days, respectively), all locations had similar PLDs (mean 41 days). For C. flavirostris, there was no significant location effect on PLD (mean 41.5 days); however, PLD at Lord Howe Island was 58 days with high variance precluding significance. Also, there was no significant variation in PLD among years for either species despite considerable variation in East Australian Current strength.