Bayou ( pulvereus) Ecological Risk Screening Summary

U.S. Fish and Wildlife Service, June 2017 Revised, January 2018 Web Version, 9/12/2018

Image: A. H. Baldwin. Smithsonian Institution. Licensed under Creative Commons (BY-NC- SA). Available: http://eol.org/data_objects/18154701. (September 2018).

1 Native Range and Status in the United States Native Range From Froese and Pauly (2017):

“Western Atlantic: From the lower Chesapeake Bay to the St. Johns River in northern Florida; In northern coasts of the , from Alabama to Corpus Christi, Texas in the USA [Murdy and Musick 2013].”

Status in the United States From Bonner (2013):

“Species ranges from Corpus Christi, Texas to Mobile, Alabama, on Gulf Coast and from St. John’s River, Florida, to lower York River, Virginia, on Atlantic slope […]”

Means of Introduction into the United States This species is native to the United States. No introductions outside the native range have been reported.

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Remarks From Collette et al. (2015):

“The main threat to this species is the loss of coastal marsh habitat via coastal development and erosion throughout its restricted range. However, several projects are aiming to restore this habitat.”

From Bonner (2013):

“[…] wide zone of hybridization (or intergradation) between F. pulvereus and the marsh killifish (F. confluentus) on Gulf Coast and narrow zone in northeastern Florida (Hardy 1980).”

Froese and Pauly (2018) report several synonyms of F. pulvereus: Zygonectes pulvereus, Zygonectes funduloides, Fundulus funduloides, and Fundulus limbatus. These synonyms, along with the accepted scientific name, were used in searching for information for this report.

2 Biology and Ecology Taxonomic Hierarchy and Taxonomic Standing From ITIS (2017):

“Kingdom Animalia Subkingdom Bilateria Infrakingdom Deuterostomia Phylum Chordata Subphylum Vertebrata Infraphylum Gnathostomata Superclass Class Teleostei Superorder Acanthopterygii Suborder Fundulus Species (Evermann, 1892) – bayou killifish, sardinilla mulata”

“Taxonomic Status: valid”

Size, Weight, and Age Range From Froese and Pauly (2017):

“Maturity: Lm ? range ? - ? cm Max length : 6.5 cm TL male/unsexed; [Murdy and Musick 2013]”

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From Bonner (2013):

“Growth and Population structure: Simpson and Gunter (1956) examined 50 adult specimens, of which males ranged from 30-46 mm long, and females were 16-45 mm long.”

Environment From Froese and Pauly (2017):

“Marine; freshwater; brackish; benthopelagic; non-migratory. […] 15°C - 26°C [Baensch and Riehl 1991; assumed to represent recommended aquarium water temperatures based on source]”

From Bonner (2013):

“Macrohabitat: Occurs in brackish marshes, bayous, freshwater rivers, and rice fields on Gulf Coast, and barrier beach ponds on Atlantic slope (Hardy 1980). Found in bayous and marshes on the Gulf Coast of Mississippi, in fresh and brackish water (Cook 1959).”

“Mesohabitat: Found in medium to low salinity habitats of coastal marshes in salinities ranging from .3-47.6 ppt; able to survive at least two months in fresh water (Gunter 1950, Griffith 1974, Miller and Guillory 1980). In the Aransas National Wildlife Refuge, Texas, species seemed to prefer sheltered, brackish waters, and were collected in salinities ranging from 0.4-16.0 ppt (Gunter 1950). Specimens collected on the Texas coast in salinities of 30.1 ppt, 44.8 ppt, and 53.9 ppt (Simpson and Gunter 1956), in water temperatures ranging from 17-34.5 degrees C (Simpson and Gunter 1956, Nordlie 2006). In Alabama, experimental lower salinity limit was freshwater, and the experimental upper salinity limit ranged from 95.6-106.4 (Griffith 1974; Nordlie 2006). In MO, lower lethal DO limit, mg kgˉ¹ was 0.88 ± 0.23, 26 degrees C, fresh water (Smale and Rabeni 1995, Nordlie 2006). In TX, observed range of DO, mg kgˉ¹ was <5 (Gelwick et al. 2001, Nordlie 2006).”

Climate/Range From Froese and Pauly (2017):

“Temperate; […]”

Distribution Outside the United States Native The native range of F. pulvereus is fully within the contiguous United States.

Introduced There are no known introductions of F. pulvereus outside of the United States.

Means of Introduction Outside the United States There are no known introductions of F. pulvereus outside of the United States.

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Short Description From Bonner (2013):

“Coloration: Simpson and Gunter (1956) described coloration for specimens collected in Texas: Base color of females olive to dark gray-blue above, shading to gold on sides and to silvery or cream on belly; large discrete dark dots scattered over body, chiefly confined to area above midline or slightly below (dotting quite variable); some females with faint, broken horizontal stripes; fins pale to dusky and pectorals yellowish at base; suborbital occasionally golden. Males with 12-17 bluish vertical stripes, with silvery interfaces of same width; fainter horizontal stripes sometimes present; olive on back and silvery on belly; black and white horizontal stripes on posterior portion of dorsal; anal and dorsal fins dusky, or anal sometimes yellowish dusky; caudal fin dusky yellowish; dorsal ocellus clear, faint, or absent. Hubbs et al. (1991) noted the following information regarding coloration: predorsal stripe reaching occiput; dark markings on scales not concentrated at posterior edge; no noticeable cross-hatched pattern; no dark subocular bar; body mottled, barred, or irregularly spotted; body barred or not, but never with a dark spot on dorsal part of caudal peduncle; body without a distinct dark lateral band. Simpson and Gunter (1956) and Blackburn and Klee (1963) described a blue phase for the species in Texas, females having a dark gray blue above, and males with bluish vertical stripes.”

“Counts: 15+ scale rows from pelvic fin origin to isthmus; 30-40 longitudinal scale rows (Hubbs et al 1991); 5-7 gillrakers; 9-10 anal rays; 10 dorsal rays; 14-16 pectoral rays; 6 pelvic rays; 11 preoperculomandibular pores; 4 mandibular pores (Ross 2001).”

“Body shape: Small bodied with a strongly convex dorsal profile; body depth increases relative to standard length; head length goes into standard length 2.9-3.4 (Ross 2001).”

“Mouth position: Slightly superior (Ross 2001).”

“External morphology: Dorsal fin originating anterior to anal fin origin; eye contained fewer than one and one-half times in snout; gill slit extending dorsal to uppermost pectoral fin ray. Distance from origin of dorsal fin to end of hypural plate less than distance from origin of dorsal fin to preopercle or occasionally about equal to that distance (Hubbs et al. 1991).”

Biology From Bonner (2013):

“Spawning season: Based on data obtained from dissection of females collected in October, November, and January, species believed to spawn in fall and winter, and perhaps other seasons, on the Texas coast (Simpson and Gunter 1956; Nordlie 2006). On the Mississippi coast, during late March, specimens up to 29 mm SL were collected, containing fully developed eggs measuring 1.5 mm in diameter (Cook 1959). Greeley (1984) reported that Fundulus pulvereus apparently spawns only during the 3-day to 6-day periods of the biweekly or semilunar spring .”

“Spawning location: In aquaria, nylon mops utilized for spawning; eggs also deposited in large pores of corks (Blackburn and Klee 1963; Boschung and Mayden 2004).” 4

“Reproductive strategy: Low frequency sound is produced by male as he circles female from a distance; somewhat later, higher frequency sounds are produced when male positions his body either above or below that of the female in an effort to direct the female to suitable substrate for spawning (Drewry 1962; Foster 1967). In aquaria, nylon mops were used as suitable spawning location; male was observed to swim beneath the female, directing the female toward the mop; female would occasionally turn her vent upward as she neared the mop, while male remained below her; many eggs drifted to bottom, being only moderately adherent (Blackburn and Klee 1963; Boschung and Mayden 2004).”

“Fecundity: Average diameter of ripe eggs 1.6 mm (Greely 1984). Eggs from southern Louisiana stock often require 3-4 weeks before hatching (Foster 1967). In aquaria, 40 eggs laid in 1 day; eggs moderately adhesive (Blackburn and Klee 1963; Boschung and Mayden 2004).”

“Age at maturation: Based on collection of fish on the Mississippi coast, no longer than 29 mm SL, and containing fully developed eggs, sexual maturity is reached by 1 year old (Cook 1959; Boschung and Mayden 2004).”

“Food habits: Stomach contents of two specimens collected on the Texas coast yielded aquatic insects from one specimen, and two isopods (each about 1.0 mm in length) from the other (Simpson and Gunter 1956).”

From Gaines (2015):

“Dominant prey items consumed by F. pulvereus were copepods, mites, chironomids, and terrestrial and aquatic insects. The bayou killifish is an opportunistic omnivore that utilizes marsh floodplains to forage for prey, and is a link in the trophic interactions taking place within estuaries. Parasitological observations resulted in the documenting of fourteen species; all but two of which were not previously reported from F. pulvereus. The presence of trophically transmitted parasites indirectly indicates consumption of prey items not observed during the examination of stomach contents within bayou killifish (e.g., Cyrenoida floridana); as well as, the role of this marsh fish as trophic intermediate host for piscivorous predators such as alligators, herons, and raccoons. Analysis using the Gonadosomatic Index indicated a spawning season that began in February and extended through August.”

From Collette et al. (2015):

“Not a Migrant”

Human uses From Collette et al. (2015):

“This species is not utilized.”

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Diseases From Gaines (2015):

“The parasite assemblage in all 445 fish in this study was typically comprised of digenean trematodes, with at least nine different species of parasites infected in and attached to the stomachs of F. pulvereus (Table 4). Forty percent of all specimens examined contained parasites in and attached to the stomach (n=445). The most common parasite found was the monorchid digenean, L. minuta (prevalence = 27.42% (Figure 15); mean intensity = 6.27 ± 0.33) (Figure 16). Cestodes, nematodes, and two species of the digenean Homalometron spp. were also common. One species was identified as Homalometron palmeri. The other species, Homalometron sp. A remains unidentified and may be undescribed.”

No OIE-reportable diseases have been documented in this species.

Threat to humans From Froese and Pauly (2017):

“Harmless.”

3 Impacts of Introductions There are no known introductions of this species. Therefore, there are no known studies of the impacts of introductions of this species.

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4 Global Distribution

Figure 1. Known global distribution of Fundulus pulvereus, reported from the Gulf and southern Atlantic Coasts of the United States. Map from GBIF Secretariat (2017). A point in South Carolina was removed from this image and the climate matching analysis because it was a record for a different species. A point inland in Texas was removed from this image and the climate matching analysis because the coordinates given did not match the verbal description of the collection location. Limited georeferenced occurrences were available for the Atlantic coastal portion of the species range.

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5 Distribution within the United States

Figure 2. Known distribution of Fundulus pulvereus in the United States. Map from BISON (2017). A point inland in Texas was removed from this image and the climate matching analysis because the coordinates given did not match the verbal description of the collection location. Limited georeferenced occurrences were available for the Atlantic coastal portion of the species range.

6 Climate Matching Summary of Climate Matching Analysis

The climate match presented here refers only to where the species can survive in freshwater and brackish water environments and not in marine environments.

The Climate 6 score (Sanders et al. 2014; 16 climate variables; Euclidean distance) for the contiguous United States was 0.172, which is classified as a high climate match. Climate scores over 0.103 indicate high match. The highest climate matches in the contiguous United States are along the coastline of the southeastern United States, exclusive of southern peninsular Florida, and matches decrease with increasing distance from the southeastern region. Medium matches occur in southern peninsular Florida, and the interior Mid-Atlantic and Southeast from New Jersey to Texas, and small patches in Arizona. There is a low match in the Northern and Western states.

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Figure 3. RAMP (Sanders et al. 2014) source map showing weather stations in the southeastern United States selected as source locations (red; coastal Texas, Louisiana, Mississippi, Alabama, Florida, and Georgia) and non-source locations (gray) for Fundulus pulvereus climate matching. Source locations from GBIF Secretariat (2017).

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Figure 4. Map of RAMP (Sanders et al. 2014) climate matches for Fundulus pulvereus in the contiguous United States based on source locations reported by GBIF Secretariat (2017). 0= Lowest match, 10=Highest match. Counts of climate match scores are tabulated on the left.

The High, Medium, and Low climate match categories are based on the following table:

Climate 6: Proportion of Climate Match (Sum of Climate Scores 6-10) / (Sum of total Climate Scores) Category 0.000≤X≤0.005 Low 0.005

7 Certainty of Assessment Information on the biology and distribution of this species is readily available. There is some uncertainty in the climate matching analysis because this non-migratory species can be found in a wide range of salinities, and the analysis is only valid for freshwater and brackish water environments. There are no known introductions of this species and therefore no studies of the impacts of introduction. Certainty of assessment for this species is low.

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8 Risk Assessment Summary of Risk to the Contiguous United States Bayou Killifish (Fundulus pulvereus) is a fish that can be found in waters with a wide range of salinity. It is native to the Gulf of Mexico, from Alabama to Corpus Christi, Texas, and the Atlantic Coast from the Chesapeake Bay to northern Florida. F. pulvereus has a high climate match with the contiguous United States, with highest matches in its current range along the Southeast coast from Texas to North Carolina excluding southern Florida. However, RAMP (Sanders et al. 2018) was not developed for use in assessing climate match for marine species so the climate match in a marine environment cannot be assessed with an ERSS. There have been no documented introductions of F. pulvereus so no information is available on the impacts of introduction and the history of invasiveness is uncertain. The overall risk categorization for F. pulvereus is uncertain, and certainty of this assessment is low.

Assessment Elements  History of Invasiveness (Sec. 3): Uncertain  Climate Match (Sec.6): High  Certainty of Assessment (Sec. 7): Low  Overall Risk Assessment Category: Uncertain

9 References Note: The following references were accessed for this ERSS. References cited within quoted text but not accessed are included below in Section 10.

Bonner, T. 2013. Fundulus pulvereus. Fishes of Texas. Available: http://www.fishesoftexas.org/taxa/fundulus-pulvereus. (January 2018).

Collette, B. B., D. Grubbs, F. Pezold, J. Simons, J. Caruso, J. Carlson, J. D. McEachran, J. Brenner, L. Tornabene, P. Chakrabarty, R. Robertson, H. Jelks, H. Espinosa-Perez, J. Tolan, and L. Chao. 2015. Fundulus pulvereus. The IUCN Red List of Threatened Species. Available: http://www.iucnredlist.org/details/196679/0. (January 2018).

Froese, R., and D. Pauly, editors. 2017, 2018. Fundulus pulvereus. FishBase. Available: http://fishbase.org/summary/Fundulus-pulvereus.html. (June 2017, September 2018).

Gaines, L. C. 2015. Observations on the life history of the Bayou Killifish, Fundulus pulvereus (Everman 1899) in Mississippi tidal marshes. Masters thesis. University of Southern Mississippi, Hattiesburg, Mississippi.

GBIF Secretariat. 2017. GBIF backbone : Fundulus pulvereus. Global Biodiversity Information Facility, Copenhagen. Available: https://www.gbif.org/species/5203698 (January 2018).

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ITIS (Integrated Taxonomic Information System). 2017. Fundulus pulvereus (Evermann, 1892). Integrated Taxonomic Information System, Reston, Virginia. Available: https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=165 656#null. (June 2017).

Sanders, S., C. Castiglione, and M. H. Hoff. 2014. Risk Assessment Mapping Program: RAMP. U.S. Fish and Wildlife Service.

10 References Quoted But Not Accessed Note: The following references are cited within quoted text within this ERSS, but were not accessed for its preparation. They are included here to provide the reader with more information.

Akin, S., K. O. Winemiller, and F. P. Gelwick. 2003. Seasonal and spatial variations in fish and macrocrustacean assemblage structure in Mad Island Marsh estuary, Texas. Estuarine, Coastal and Shelf Science 57:269-282.

Baensch, H. A., and R. Riehl. 1991. Aquarien atlas, volume 3. Mergus, Verlag für Natur-und Heimtierkunde, Melle, Germany.

Blackburn, A., and A. J. Klee. 1963. A beautiful backyard “import”, the bayou killifish. Aquarium 32(8):28-31.

Boschung, H. T., Jr., and R. L. Mayden. 2004. Fishes of Alabama. Smithsonian Books, Washington, D.C.

Cook, F. A. 1959. Freshwater fishes in Mississippi. Mississippi Game and Fish Commission, Jackson, Mississippi.

Drewry, G. E. 1962. Some observations of courtship behavior and sound production in five species of Fundulus. Masters thesis. University of Texas, Austin.

Evermann, B. W. 1892. A report upon investigations made in Texas in 1891. Bulletin of the U.S. Fish Commission 11:61-90.

Evermann, B. W. 1899. Report on investigations by the U.S. Fish Commission in Mississippi, Louisiana, and Texas, in 1897. Report of the U.S. Fish Commission 24:287-310.

Fournie, J. W., and R. M. Overstreet. 1993. Host specificity of Calyptospora funduli (Apicomplexa: Calyptosporidae) in Atheriniform fishes. Journal of Parasitology 79(5):720-727.

Foster, N. R. 1967. Comparative studies on the biology of (Pisces: Cyprinodontidae). Doctoral dissertation. Cornell University, Ithaca, New York.

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Gelwick, F. P., S. Akin, D. A. Arrington, and K. O. Winemiller. 2001. Assemblage structure in relation to environmental variation in a Texas Gulf Coastal wetland. Estuaries 24(2):285- 296.

Greeley, M. S., Jr. 1984. Spawning by Fundulus pulvereus and Adinia xenica (Cyprinodontidae) along the Alabama Gulf Coast is associated with the semilunar tidal cycles. Copeia 3:797-800.

Griffith, R. W. 1974. Environment and salinity tolerance in the genus Fundulus. Copeia 2:319- 331.

Gunter, G. 1950. Distributions and abundance of fishes on the Aransas National Wildlife Refuge, with life history notes. Publication of the Institute of Marine Science, University of Texas 1(2):89-101.

Hardy, J. D., Jr. 1980. Fundulus pulvereus (Evermann), Bayou killifish. Pages 525 in D. S. Lee et al. Atlas of North American freshwater fishes. North Carolina Museum of Natural History, Raleigh, North Carolina.

Hubbs, C. 1957. Distributional patterns of Texas fresh-water fishes. The Southwestern Naturalist 2(2/3):89-104.

Hubbs, C., R. J. Hubbs, and G. P. Garrett. 1991. An annotated checklist of the freshwater fishes of Texas, with keys to the identification of species. Texas Journal of Science, Supplement 43(4):1-56.

Kinsella, J. M., and R. W. Heard, III. 1974. Morphology and life cycle of Stictodora cursitans n. comb. (Trematoda: Heterophyidae) from mammals in Florida salt marshes. Transactions of the Microscopical Society 93(3):408-412.

Miller, C. and V. Guillory. 1980. A comparison of marsh fish communities using the Wegener ring. Proceedings of the Southeastern Association of Fish and Wildlife Agencies 34:223- 233.

Murdy, E. O., and J. A. Musick. 2013. Field guide to fishes of the Chesapeake Bay. JHU Press, Baltimore, Maryland.

Nordlie, F. G. 2006. Physiochemical environments and tolerances of cyprinodontoid fishes found in estuaries and salt marshes of eastern . Reviews in Fish Biology and Fisheries 16(1):51-106.

Ross, S. T. 2001. The inland fishes of Mississippi. University Press of Mississippi, Jackson, Mississippi.

Simpson, D. G., and G. Gunter. 1956. Notes on habitats, systematic characteristics and life histories of Texas salt water Cyprinodontes. Tulane Studies in Zoology 4(4):115-134.

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Smale, M. A., and C. F. Rabeni. 1995. Hypoxia and hyperthermia tolerances of headwater stream fishes. Transactions of the American Fisheries Society 124:698-710.

Warren, M. L., Jr., B. M. Burr, S. J. Walsh, H. L. Bart, Jr., R. C. Cashner, D. A. Etnier, B. J. Freeman, B. R. Kuhajda, R. L. Mayden, H. W. Robison, S. T. Ross, and W. C. Starnes. 2000. Diversity, distribution, and conservation status of the native freshwater fishes of the southern United States. Fisheries 25(10):7-29.

Wiley, E. O. 1986. A study of the evolutionary relationships of Fundulus topminnows (Teleostei: Fundulidae). American Zoology 26:121-130.

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