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Annotated checklist and distribution of New Jersey freshwater fishes, with comments on abundance.

ABSTRACT: An investigation of the freshwater, anadromous, catadromous, and currently existing and long occurring introduced species that comprise the freshwater fish fauna of New Jersey indicates the occurrence of 86 species in 24 families. Containing the most species (20) is the carps and minnows) family, followed by the sunfish family (14 species), and the North American catfishes family (7 species); these throe families include 47.6% of the total species. Ten families contain only one species each. Included in the state fish fauna are: one native species (longnose gar) that has been extirpated from New Jersey waters; 19 species that are known to be not native and to have been introduced to New Jersey, of which three (goldfish, common carp, grass carp) are native to Asia and Eurasia and 16 to other areas of the eastern United States and adjacent Canada; and 66 species that are presumably native, although three (bowfin, eastern mosquitofish, warmouth) of these species are so listed tenuously because one or more of them might have occurred, or been introduced (as the first two of these three species are known to have been), or both. Some 21 of the 86 species are extensively coastal forms and occur to a large degree in the saltwater, brackish water, and freshwater tidal waters of the state. This group contains most of the commercially important species, and a number of these forms have suffered decreases in abundance from overexploitation and habitat destruction over the past three centuries.

Of the total of 86 species, two are native and riverine, of which one form (longnose gar) has now been extirpated in New Jersey, and the second fibrin (quillback) is poorly known in the state. At least two additional native species were apparently originally restricted to rivers in New Jersey, but they have been introduced elsewhere in the state. Nine other forms occur primarily in rivers and are also part of the coastal forms category mentioned above. Fourty-three species of native and introduced species are widely distributed in the state and found at many localities, although many of these species in southern New Jersey are restricted to drainages tributary to the Delaware River and/or Delaware Bay. Another 13 species occur statewide, but at relatively few localities compared to the 43 forms. Nine species are found only or primarily in northern New Jersey, in the Piedmont, Highlands and Ridge and Valley Provinces, but are common. Six species are found in northern New Jersey, but each form is represented by relatively few localities of occurrence, although some species are common at a particular locality. Three species are restricted, or largely so, to the southern half of New Jersey, in the Coastal Plain Province. One form is found only at very few, sites in southern New Jersey. Some five species of the total are now known from very few sites in New Jersey, and two of these species now occur at considerably fewer sites than formerly.

I list 49 species (and do not include two stocked, non-reproducing, ephemerally-occurring trout species) that occur in the Pinelands region of the state. Thirteen of these forms are known to be, or may be, introductions into this region, of which three species are now widespread and often common. Of the total of 49 forms, 10 occur on the margin of the Pinelands and are known from few localities; 15 species are restricted to the main rivers in the region; five additional forms are found mostly in rivers but also outside them; 16 species are widespread and often common in the Pinelands; and three species are recorded from a few scattered localities within the Pinelands. One species in the last category has decreased greatly in the Pinelands. Of the total of 49 species, only some 12 species are native and occur widespread in the acidic and dystrophic waters of the Pinelands interior.

KEY WORDS: New Jersey, freshwater fishes, fish distributions, anadromous fishes, catadromous fishes, introduced fishes, New Jersey Pinelands

INTRODUCTION

The number of species reported to comprise the freshwater fish fauna of New Jersey varies considerably. For example, a list prepared by the Bureau of Fisheries Laboratory at Lebanon, New Jersey, in 1974, contains 77 species in 18 families. Stiles (1978) identified 114 species in 27 families; Anonymous (1993) documented 116 species in 24 families; and Niles et al. (undated) and Anonymous (2004) registered 81 species in 21 families. Some of these differences are a result of intervening changes in the fauna, and some no doubt to diverse definitions of "freshwater" species. Some of these lists, however, contain numerous species, and some families, that have never been recorded near New Jersey. In this work, I have compiled a list of freshwater fish species with annotations and distribution figures for all native, extirpated, and introduced (from outside the United States or from other states) forms with currently existing and long occurring populations. There is no existing publication on New Jersey freshwater fishes that provides such a comprehensive treatment of their distributions. This paper is part of an ongoing effort to produce a larger and more detailed document on the highly dynamic freshwater fish fauna of New Jersey.

In compiling the list of freshwater species, I include those forms that spend all of their life in freshwater; one species that begins its life in saltwater and then lives for years in freshwater or brackish water before returning to the sea to reproduce (i.e., a catadromous species, specifically the American eel); species that reproduce in freshwater but then spend most of their life in brackish water or saltwater (i.e., anadromous and semi-anadromous fishes, e.g., many of the herrings, white perch, striped bass, rainbow smelt, and hogchoker); and those forms that are often considered to be estuarine or marine but have representatives that occur regularly and extensively in freshwater in New Jersey (e.g., the mummichog, inland silverside, three-spine stickleback, and four-spine stickleback). I do not include species that occur on a limited basis in freshwater on the edges of their largely brackish water and saltwater habitats (e.g., the bay anchovy and Atlantic silverside). No introduced freshwater species is included unless: (1) it is known to have reproducing populations in New Jersey (e.g., the smallmouth bass, largemouth bass, and bluegill); or (2) it is not reproductive, or usually not reproductive, but is such a highly desirable species that it is introduced, typically by state biologists, so extensively as to maintain populations (e.g., the grass carp, northern pike, and walleye). Thus, I have not included rare marine fish waifs and those records of introduced or escaped non-native species which typically have been represented by only one or a few individuals, and which have no chance of long-term survival due to the lack of suitable habitats and water temperatures.

This paper gives a broad overview of the distribution and abundance of the New Jersey freshwater fishes. The format is designed primarily for those individuals who have considerable knowledge of these fishes because it would not be possible to give detailed descriptions of so many species. Thus, in order to help maintain focus on the overview, I use only common names in the text. The general reader will also be more familiar with these common names. The scientific names of the fish species can be found in Tables 1 and 2, in the species distribution figures, and in the references mentioned.

In regard to taxonomy, I avoid comment on name changes that have occurred recently (e.g., the use of the common name of cutlip, rather than of cutlips, minnow; the use of Sander instead of Stizostedion for the genus name of the walleye; the use of Atherinopsidae instead of Atherinidae for the family name of the inland silverside; and the use of Achiridae instead of Soleidae for the family name of the hogchoker). Similarly, I avoid consideration of the recent name changes of the mosquitofishes Gambusia affinis affinis and G. affinis holbrooki to G. affinis and G. holbrooki, respectively, the questions of whether or not G. holbrooki is native to New Jersey or was introduced long ago or both, of where and when G. affinis was introduced, and of possible hybridization between the two species. More research on Gambusia in New Jersey is needed.

I continue to add to the data base on New Jersey freshwater fishes and welcome comments and contributions from fishermen, scientists, and others who can expand our knowledge of this topic.

MATERIALS AND METHODS

Data Sources

Preserved Specimens

Critically important data on New Jersey fish holdings were obtained from the following institutions listed in alphabetical order and with the name(s) of the collection curator(s) or manager(s) given in parentheses: The Academy of Natural Sciences of Philadelphia (W. Saul), American Museum of Natural History (S. Schaefer), Cornell University (J. P. Friel and C. M. Dardia), Harvard University (K. Hartel), National Museum of Natural History (J. T. Williams), North Carolina State Museum of Natural Sciences (W. Starnes and G. M. Hogue), and Tulane University (M. Doosey). J. Dighton, Rutgers University, Camden, contributed an extensive and extremely valuable collection of preserved New Jersey fishes and an accompanying typed catalogue prepared by former professor R. W. Hastings. L. Sarner, New Jersey Department of Environmental Protection (NJDEP), Division of Fish, Game and Wildlife, contributed a collection of fishes from the Nacote Creek Field Research Station, Galloway Township, Atlantic County.

Unpublished Data

H. Carberry and L. Barno of the NJDEP, Freshwater Fisheries, kindly allowed access to a very extensive and highly useful computer database on the New Jersey freshwater fishes. D. Byrne, NJDEP, Marine Fisheries, very generously provided data from his years of sampling fishes in ocean waters along the New Jersey coast. B. Margolis and some of his colleagues shared their extensive data collected mostly in northern New Jersey in an Index of the Biological Diversity sampling program. J. Normant, NJDEP, Nacote Creek Field Station, Galloway Township, New Jersey, provided valuable fish data collected in his sampling program in Delaware Bay.

Field Sampling

I have made several hundred collections of freshwater fishes in New Jersey over the past 30 years, many with my Stockton College students in field exercises. Collections were made mostly in southeastern New Jersey, and less so in several tributaries of the Delaware River in southwestern New Jersey. In order to become familiar with more distant areas of the state, I collected data in the non-tidal portion of the Delaware River in recent years, in waters in the mountains and hills of northwestern New Jersey, and on the fringes of the northeastern New Jersey metropolitan area. I also surveyed fishermen and bait and tackle shop workers along portions of the Delaware River. Some sites were sampled specifically in my attempt to fill in gaps in distribution between known sites for a species. Additional data were obtained by accompanying the following biologists (listed alphabetically) during some of their field work: F. and J. Akers, D. Byrne, H. Carberry, S. Eisenhauer, and B. Margolis.

Literature Sources

Many data are derived from the published literature. Some relatively limited areas of the state have been sampled extensively, as follows: Smith (1971), Alloway and Stowe Creeks; Hastings and Good (1977), Woodbury Creek; Hastings (1979), Pinelands; Hastings (1984), Mullica River; Kraus et al. (1987) and Kraus and Bragin (1989, 1990), Hackensack River; Able and Fahy (1998), Great Bay and Little Egg Harbor and adjacent areas; Zampella et al. (2001), Mullica River Basin; and Zampella et al. (2003), Rancocas Creek Basin. Many data are derived from the "gray" literature. This literature is an eclectic collection, as I used any reference that contained information on fishes identified to species with collection locations precise enough to allow me to plot the data. Included in the literature sources are graduate degree theses, publications with localities of interest to fishermen, fish collection sites for studies of heavy metals, environmental impact studies, and data from conservation groups for certain protected areas. The list of references examined is extensive, and the data from these and other sources likely give an accurate picture of the distribution of the freshwater fishes of New Jersey. For brevity, locality citations by species are not provided.

Methods of Fish Collection

I made collections, often with students, mostly with 3 m x 1.2 m and 4.5 m x 1.2 m small mesh flat seines, sometimes with 6 m x 1.2 m or larger small mesh bag seines, and in the Mullica River and adjacent bay waters with a 4.9 m bottom trawl. I also obtained specimens, and valuable sight records, by snorkeling with an aquarium net in summer in the non-tidal Delaware River. Fishes reported in the literature were collected with a variety of equipment. Commonly used in small streams and shoreline habitats were seines of several sizes, backpack electroshockers, and dip nets. Away from shore habitats, commonly used equipment was 4.9 m and 2.7 m bottom trawls and boat electroshockers. In ocean habitats, large boats and large bottom trawls were used. Some records were also derived from fishes collected on the intake screens of electric generating stations.

Data Organization

Data in the tables are listed in phylogenetic sequence, that is, listed from the most "primitive" to the most "advanced." The sequence used, and the class, order, family, genus, species, and common names used herein are from Gilbert and Williams (2002). Names not given in this reference follow Page and Burr (1991). Author(s) of scientific names and the years of their publication are from Robins et al. (1991). The species accounts by individual authors in the compendium by Lee et al. (1980) and in the book by Page and Burr (1991) have been especially valuable in the preparation of this paper.

The annotated list for each species includes:

1. The general habitat(s) of the form (i.e., whether it is a freshwater, brackish water, or saltwater species, listed in the order of primary occurrence), and if it is catadromous or anadromous);

2. If the form is native to New Jersey or is introduced (introduced forms are also identified with an asterisk);

3. The New Jersey drainage(s) in which the species has been recorded (i.e., in waters that drain directly into the Atlantic and/or into the Delaware River and/or into the Delaware Bay);

4. If the form occurs in the Pinelands, which is a region of special interest because of its unusual habitats and species, with some forms common here and restricted elsewhere in New Jersey;

5. If it is an introduced species, the area to which it is native;

6. The relative abundance of the form in New Jersey (i.e., categorized as rare, occasional, common, or abundant, based on my personal observations and on data from other sources).

Locations of occurrence for landlocked anadromous species are depicted on the figures, but only the usual condition (e.g., whether it is an anadromous, brackish water, or a saltwater species) of the form is listed in the pertinent table.

My comments on the abundance of a species are subjective. Usually, "abundant" is indicated if a species occurs at many localities. However, a species with few known localities might occur in large numbers at one or more of these sites and, consequently, can also be considered "abundant." Subjectivity can be a result of a number of other factors. One of these factors is uneven sampling effort expended in some habitats or in some drainages. Another is that some species are secretive or otherwise difficult to sample, and the capture of small numbers may not reflect the actual numbers that occur. Some non-native species may be common or abundant at or near points of introduction, and less so in areas subsequently invaded. Some species are apparently numerous when they school, especially when they migrate, and thus may give the impression of generally occurring in abundance.

Other factors contribute to subjectivity in abundance estimates. Some commercially important species (e.g., the sturgeons and rainbow smelt) were abundant centuries or decades ago, based on data in the older literature, but are uncommon or rare today, although apparently slowly increasing in numbers. Further, the numbers of some species (e.g., the striped bass) in New Jersey and adjacent waters have fluctuated greatly, from being common, to rare, to abundant, all in the most recent few decades. Likewise, a number of species appear to be abundant at this point in time, but they are not nearly as abundant as the older literature seems to indicate. Abundance is also a function of the trophic level of a fish. For example, large piscivorous species are typically never as abundant as their smaller, faster-maturing and faster-growing prey fish species.

Distribution Figures

The species distribution figures show locations of recorded occurrence. A dot represents a locality at which any life history stage (e.g., juvenile, young, and adult) of a species has been found (i.e., not all the stages may occur at a given locality, although for the majority of species, they do). A dot may represent a locality at which a single specimen was captured or observed, or dozens of individuals, or even more, and made at one point in time or over time. Larger dots are used on some figures to clarify localities (e.g., those that lie on the irregular back bay Atlantic coast), and to render more conspicuous the localities when only a few exist for a species. The number of dots on a figure should ideally reflect all localities at which a species occurs. This is probably more likely for the larger species, notably the game species, which have been sampled by state biologists over many years, and consequently the number of localities is generally accurately known. It is much less likely for the non-game species, particularly the smaller and less distinctive forms, which have typically been sampled less intensively and/or only incidentally to research on other species, and thus the actual number of localities of occurrence may be considerably greater than is represented by the number of dots. A large number of dots shown for some species on some rivers and streams (e.g., the Mullica River and the Rancocas Creek systems, Woodbury Creek, and the Hackensack River), as opposed to nearby waters, probably only signifies that more extensive sampling has taken place there, and does not indicate major differences in natural occurrence, for example, due to differences in habitat. A few imprecise locality records (perhaps a total of 10 records for all species combined) were found for some species, and an approximate midpoint location was plotted for these records. Solid triangles show localities plotted from small range maps presented in Lee et al. (1980). The triangles reflect approximate locations. The reader is thereby alerted to the presence of records that are located at some distance from more precise locations. For some species, locations are also plotted in adjacent portions of New York State, including the Hudson and Delaware Rivers. Some of these localities are in the same drainages that feed or drain New Jersey waters, and fishes reported from them could thus also be expected to occur in New Jersey. I consider a few of the plotted records, often isolated from other records, to be of questionable accuracy. Nevertheless, by including such records, I believe that they are more likely subject to study and possible correction in the future. I have usually given less weight to such records in my comments in other parts of this paper. A few obviously misidentified records in some databases were not plotted. Note that a species may no longer occur at an indicated locality. Likewise, a species may occur at sites that are not indicated. Every effort has been made to be accurate, but I was not able to examine all the specimens collected by others.

The fish distribution figures are arranged in phylogenetic sequence. This should enable the reader to quickly locate the figure for a given species by leafing through the distribution figures, if its phylogenetic placement is known. However, use of this scheme means that the figure numbers in the text do not follow consecutively, because the fishes in the text are not referred to in taxonomic order. The number of the figure that shows the distribution of a species is usually given only the first time that the species is referred to in the "Results" and the "Species Distribution and Abundance" sections.

Voucher specimens and series have been deposited in The North Carolina State Museum of Natural Sciences, Raleigh.

RESULTS AND DISCUSSION

Area Descriptions

Figure 1 shows New Jersey counties superimposed on a drainage base map. This figure facilitates site location by county.

[FIGURE 1 OMITTED]

Figures 2 and 3 identify all the New Jersey rivers and the larger streams, and show for selected waters the approximate upper limits of tidal water (t) and brackish water (b) occurrence (R. G. Arndt, personal observations). Considerable lengths of tributaries to the Delaware River downstream of Trenton, and even more so to Delaware Bay (Figure 2), are tidal. In contrast, except for the Mullica and the Egg Harbor Rivers and their lower tributaries, streams that flow directly into the Atlantic Ocean or its back bays are characterized by short tidal reaches (Figure 3).

[FIGURES 2-3 OMITTED]

Figure 4 shows the locations of the five major physiographic provinces (i.e., Outer Coastal Plain, Inner Coastal Plain, Piedmont, Highlands, and Ridge and Valley) in New Jersey (Robichaud and Buell, 1983). The physiography of each, including the elevation, slope, and substrate, greatly influences the types and sizes of water bodies that can occur (e.g., it affects the absence or presence of lakes and their depths, shapes and sizes; the directions of stream flow and the speed of their currents; the stream substrates and their physicochemical characteristics, etc.), and thus the life forms, including the fishes, found there. The Coastal Plain Province (incorporating the Outer and Inner Coastal Plains) consists of unconsolidated soils and sediments. It is generally fiat, with elevations usually less than 33 m, but with some unusual and local high points that reach to 114 m. The Piedmont has a soft rock substrate, gently hilly terrain, and a regional slope from its highest elevation of about 122 m in its northwestern section to about 30 m in its southeastern section along the Delaware River. The Highlands Province consists of roughly parallel mountains and valleys, with the mountains at about 305 m elevation but reaching a maximum of about 457 m in the northwest. The Ridge and Valley Province consists of relatively widely separated mountains with broad intervening valleys at elevations of about 122 m to 305 m, and with the highest point of the state at about 550 m in its northwest corner (Robichaud and Buell, 1983).

[FIGURE 4 OMITTED]

Figure 5 shows the location of the Pinelands (gray shaded) (from the NJDEP Geographic Information Systems base map) in relation to drainages and counties. Many other references (e.g., McCormick, 1970) indicate that the Pinelands region is larger than that shown in Figure 5.

[FIGURE 5 OMITTED]

Species Distribution and Abundance

Figure 6 displays all New Jersey sites combined at which one or more species of fishes were recorded.

[FIGURE 6 OMITTED]

Figures 7-91 show the distribution of New Jersey freshwater fishes, by species, and they are arranged in phylogenetic order.

[FIGURES 7-91 OMITTED]

In sum, I record a total of 86 species in 24 families of native, extirpated, and introduced freshwater fish forms with long occurring populations in New Jersey (Table 1). This number is larger by five species (i.e., the quillback, inland silverside, Atlantic tomcod, warmouth, and hogchoker) and three families (Atherinopsidae, Gadidae, and Achiridae) than that listed by Niles et al. (undated) and Anonymous (2004). Containing the most species (20) is the family of carps and minnows, followed by the family of sunfish (14 species), and the family of North American catfishes (7 species). These three families include 47.6% of the total species. Ten families contain only one species each.

Of the 86 species, one native form (longnose gar) has been extirpated in New Jersey (Table 2); its former distribution is shown in Figure 11. Three species (i.e., bowfin, warmouth, and eastern mosquitofish) are tenuously listed as native, for all three species could have occurred in the state because known populations of each occur relatively close to New Jersey. The bowfin (Figure 12) has been recorded from a few widely scattered localities. The bowfin is also known to have been stocked in the state. The warmouth (Figure 80) has been recorded from two known New Jersey localities, and one unknown locality. The two localities, and probably also the third, are close to populations of the warmouth located to the south. Whether the warmouth occurred in New Jersey naturally, was introduced, or both, or found its way here from where it was supposedly introduced into northern Delaware, or from elsewhere, has not been established. The eastern mosquitofish presumably occurs at widely scattered localities throughout the state. It was introduced into the state, originally about a century ago. The form occurs along the coast of Delaware, and suitable habitat exists for it in southern New Jersey. It might be a New Jersey native. However, since I have not identified the eastern mosquitofish from New Jersey, and since there is apparent confusion on its taxonomy in New Jersey, I include no distribution figure for this form.

Of the remaining 82 species, 19 forms (22.0% of all 86 species) are not native to New Jersey and are known to have been introduced, usually many decades ago. (For documentation that these species--identified subsequently--have been introduced, see the respective species accounts in Lee et al., 1980.) Introductions were for the purpose of increasing the quality of life by providing species for food for humans, and/or for sport, for insect and plant pest control, as ornamental species, and as prey for other fish species of value to man. Some introduced species may have originated from "bait bucket releases." Nine species (i.e., the goldfish, common carp, rock bass, green sunfish, bluegill, smallmouth bass, largemouth bass, white crappie, and black crappie; respectively, Figures 19, 23, 73, 78, 81, 82, 83, 84, and 85) of the 19 introduced forms are now so widely distributed in the state and have such strong self-reproducing populations that most persons probably think of them as being natives. Through further invasion, releases, and stocking, many of these species will be even more widespread in the future. Another four of the introduced species (i.e., the northern pike, rainbow trout, brown trout, and walleye; respectively, Figures 51, 56, 57, and 90) are popular sport fishes that are relatively long-lived, long have been and continue to be widely introduced, and have benefited from the creation of suitable habitat, and therefore have relatively large distributions. There is no documented reproduction of northern pike and walleye in the wild in New Jersey, and of only a few populations of the rainbow trout and the brown trout (P. L. Hamilton, personal communication, 2004). Thus, their presence in New Jersey depends only or mostly on continued stocking (for example, more than 600,000 individuals of three trout species were stocked per year in recent years by state personnel, and many more by private parties) (P. L. Hamilton, personal communication, 2004). The remaining six species (i.e., the grass carp, bluntnose minnow, fathead minnow, black bullhead, lake trout, and western mosquitofish; respectively, Figures 20, 33, 34, 44, 59, and 65) of the 19 introduced forms are little known to the average person because these forms occur at few localities, some are small in size, and none is a major sport fish in New Jersey. The continued presence of the grass carp and the lake trout is probably due to stocking (although there is some spawning of the lake trout); the other four species are probably naturalized. Future increases in the distributions of members of this group will likely be modest.

Of the remaining 63 and native species, some 21 forms (24.4% of all 86 species) are extensively coastal fishes (i.e., with at least part of the population found commonly, at least at some stage of their life, along the edge of the state in tidal waters, and with a tolerance to a wide range of salinity, including of freshwater). Twelve of these species (i.e., the sea lamprey, shortnose sturgeon, Atlantic sturgeon, blueback herring, hickory shad, alewife, American shad, gizzard shad, rainbow smelt, Atlantic tomcod, white perch, and striped bass; respectively, Figures 8, 9, 10, 14, 15, 16, 17, 18, 55, 61, 70, and 71) are anadromous or semi-anadromous, and one (i.e., the American eel; Figure 13) is catadromous. The American eel is also widespread inland, and much less so the alewife and white perch; the landlocked localities for the last two species are probably a result of stocking.

Of the total of 21 extensively coastal species, 12 forms (i.e., the shortnose sturgeon, Atlantic sturgeon, American eel, blueback herring, hickory shad, alewife, American shad, rainbow smelt, Atlantic tomcod, white perch, striped bass; and white catfish, Figure 43) are or have been commercially and/or recreationally important, primarily as food for humans. One additional species (the mummichog; Figure 64) is important because it is collected and sold commercially for use as a fish bait. Four species (i.e., the shortnose sturgeon, Atlantic sturgeon, rainbow smelt, and Atlantic tomcod) of this total of 13 commercially important species show pronounced declines in abundance, and the first two forms of the four mentioned are now protected. Four other species (i.e., the American shad, blueback herring, alewife, and striped bass) of these 13 forms have shown less severe declines in population size and are now afforded some protection (e.g., commercial fishing licenses are no longer being issued to harvest the American shad in the Delaware River; reduced daily catch limits have been implemented for the next two species; and size limits and catch limits have been implemented for the striped bass). The American eel and white perch are widespread and common; nevertheless, both are also afforded some protection. Populations of the white catfish, mummichog, hickory shad, and gizzard shad appear to be healthy (i.e., these fishes are still common, although uncorroborated decreases in numbers of the mummichog have been reported).

Of the six as yet unidentified extensively coastal species (i.e., the inland silverside, banded killifish, fourspine stickleback, threespine stickleback, ninespine stickleback, and hogchoker; respectively, Figures 62, 63, 66, 67, 68, and 91), five are small in size, widely distributed, and probably have robust populations. The one exception is the ninespine stickleback, which probably occurs in small and localized populations, since there are few localities and individuals recorded.

Of the 42 native species not yet identified, one (i.e., the quillback; Figure 39) is riverine, and is native to the Delaware River (quillback, Platania and Jenkins, 1980), and it has been recorded from the Raritan River. It is seldom recorded and apparently is rare. The longnose gar, previously mentioned, was also riverine in New Jersey. The channel catfish (Figure 47) in New Jersey is apparently native only to the Delaware River, and is a generally riverine species (channel catfish, Glodek, 1980). However, this catfish has been introduced extensively elsewhere in New Jersey as a sport fish. The white catfish, also mentioned earlier, occurs in large rivers in New Jersey, but also in small rivers and streams, and this form has a wide native distribution in this state. It is a riverine species in New Jersey, and is largely so in other parts of its range (white catfish, Glodek, 1980). However, it has also been introduced in other parts of New Jersey. I have found it to be abundant.

Of the remaining 40 native species not yet considered, 15 forms are widely distributed in the state. Of these species, five (i.e., the white sucker, creek chubsucker, redfin pickerel, chain pickerel, and tessellated darter; respectively, Figures 40, 41, 50, 53, and 87) have robust distributions and occur essentially statewide. They are often common. A sixth species (i.e., the common shiner, Figure 26) has a somewhat less robust distribution, but is also often common. Five species (i.e., the yellow bullhead, eastern mudminnow, mud sunfish, bluespotted sunfish, and banded sunfish; respectively, Figures 45, 54, 72, 75, and 76) are somewhat less widely distributed statewide and are apparently better adapted to habitats present in the Coastal Plain. All are often common to abundant. Four species (i.e., the golden shiner, brown bullhead, pumpkinseed, and yellow perch; respectively, Figures 27, 46, 79, and 88) are widespread in the state, probably at least partly as a result of intrastate introductions. They are often common to abundant. (A sixteenth species, the banded killifish, is also widely distributed in the state and is often common. This species has already been considered in another category.)

Of the remaining 25 native species not yet mentioned, 11 forms (i.e., the American book lamprey, satinfin shiner, spotfin shiner, eastern silvery minnow, comely shiner, spottail shiner, swallowtail shiner, fallfish, tadpole madtom, margined madtom, and redbreast sunfish; respectively, Figures 7, 21, 22, 25, 28, 31, 32, 38, 48, 49, and 77) are widely distributed in the state, but with fewer known localities than those species that have been identified in the preceding paragraph. The American brook lamprey and the comely shiner are only occasionally observed, but they are small and easily confused with other species and may be more common than records indicate; the other species of the H are often common. Five more species (i.e., the cutlip minnow, blacknose dace, longnose dace, creek chub, and brook trout; respectively, Figures 24, 35, 36, 37, and 58) are widespread in the northern half of the state, with the blacknose dace, creek chub, and brook trout also weakly represented in the Coastal Plain. All are sometimes common. Three additional species occur only in the Coastal Plain, but they are widespread there. These forms are the pirate perch (Figure 60), blackbanded sunfish (Figure 74), and swamp darter (Figure 86). All are often common and sometimes abundant. (The sea lamprey, mentioned earlier, is also only occasionally observed because of its secretive life history, and it may be more widespread and common than records indicate.)

Of the remaining six native species not yet mentioned, three (i.e., the muskellunge, slimy sculpin, and shield darter; respectively, Figures 52, 69, and 89) are restricted to the hilly and rocky part of the state, mainly the northern and western portion of New Jersey. The muskellunge localities east of the Delaware River are almost certainly a result of stocking. The muskellunge is occasional to rare, as is usual for a top-level predator, and the sculpin and the shield darter are usually occasional. This may be the typical condition for the darter, but the sculpin is also sometimes locally common. Two species (i.e., the bridle shiner and ironcolor shiner; respectively, Figures 29 and 30) are known from relatively few localities, but these are widely distributed in the state. Only a handful of localities, however, for these two species occur now, probably as a result of habitat degradation and destruction. I know of only one locality where the ironcolor shiner is abundant. Both of these species deserve protection. The last of these six species (i.e., the northern hog sucker; Figure 42) is known from only two localities in the hilly and rocky Delaware River area. Two other localities (triangles) indicated on Figure 42 need verification. The northern hog sucker is apparently rare in New Jersey, but is better represented on the Pennsylvania side of the Delaware River. It needs more study in New Jersey.

Pinelands Region

Species (n = 49) of the Pinelands (Figure 5), located in the Coastal Plain (Figure 4), are listed in Tables 2 and 3. Those species (n = 10) (Table 3) that are referred to as being marginal are found more or less only on the edge of the Pinelands, and are not typical of the interior. The sea lamprey (Figure 8) apparently occurs only at a few localities on the Pinelands' edge, and the introduced goldfish, grass carp, common carp, black bullhead, and white crappie (respectively, Figures 19, 20, 23, 44, and 84) probably occur largely in man-made ponds and lakes, and as washouts in their outflows. The fallfish (Figure 38) occurs at least at one site on the northern edge of the Pinelands, and the brook trout (Figure 58) at one site on its western edge. Other localities for trouts in the Pinelands region result from introductions and none are reproducing. The eastern mosquitofish and the western mosquitofish (Figure 65) presumably have been found, or would be expected to occur, at only a few localities on the margin of the Pinelands.

Of those fish species (n = 15) that have been recorded only in rivers in the Pinelands (Table 3), I include the bowfin (Figure 12). I have only a few river records of the blueback herring (Figure 14). Even fewer records exist for the hickory shad, American shad, and gizzard shad (respectively, Figures 15, 17, 18). These records are so few that it is likely that all are waifs, derived from where these species are often common and widely distributed nearby in inshore ocean waters and Delaware Bay. This hypothesis is strengthened by the fact that these last four species are migratory. Nevertheless, they may also occur in the Pinelands in larger numbers than is now known. The American shad (Figure 17) apparently once had a reproducing population in the Mullica River, but it has been extirpated. Of the remaining 10 river species, the alewife, white perch, striped bass, mummichog, inland silverside, fourspine stickleback, and hogchoker (respectively, Figures 16, 70, 71, 64, 62, 66, and 91) occur in the downstream portions of rivers in much of the salty tidal water and upstream to freshwater, and all probably do so seasonally. The white sucker, white catfish, and banded killifish (respectively, Figures 40, 43, and 63) inhabit primarily upstream freshwater habitat and the low salinity reaches of tidal waters. Five additional species (i.e., the golden shiner, brown bullhead, tessellated darter, yellow perch, and black crappie; respectively, 27, 46, 87, 88, and 85; Table 3) occur largely in rivers in the Pinelands. These five forms are also found at other locations scattered through the Pinelands, some of these, I suspect, as a result of being introduced there.

Of the 16 species listed as widespread and often common in the Pinelands (Table 3), two forms (i.e., the bluegill and largemouth bass; respectively, Figures 81 and 83) are introduced, and along with the pumpkinseed and tadpole madtom (respectively, Figures 79 and 48), are apparently somewhat limited in distribution. All are absent from the "heart" of the Pinelands. Considering also the three species listed as occurring "scattered" (Table 3), the channel catfish (Figure 47) has been introduced in the Pinelands, and probably this is true also for the redbreast sunfish (Figure 77). The native ironcolor shiner (Figure 30) is now rare in the Pinelands. Thus, only some 12 species of fishes are native and occur widespread in the relatively acidic and dystrophic waters of the Pinelands. None of these 12 species is restricted to the Pinelands in New Jersey, but all also occur outside of this region, as well as in other states. Based on preliminary sampling, I have also found many of these species in the small streams used to regulate water levels in active commercial cranberry bogs.

CONCLUSIONS

The freshwater fish fauna of New Jersey exhibits considerable variation. One species (longnose gar; Figure 11) has been extirpated in New Jersey, two species (i.e., the bridle shiner and ironcolor shiner; respectively, Figures 29 and 30) apparently now have many fewer populations than previously recorded, and a number of species (i.e., the shortnose sturgeon, Atlantic sturgeon, American shad, blueback herring, alewife, rainbow smelt, and Atlantic tomcod; respectively, Figures 9, 10, 17, 14, 16, 55, and 61) exhibit greatly reduced numbers as a result of overfishing and pollution. Another category of human impact on the natural waters is the planned and unplanned introduction of many species of non-native fishes, of which about 13 species are now established and widespread in the state; another four species are fairly widespread, especially in the northern half of the state, and survive largely through continuing introductions; and another two species are known only from a few localities for each. Such decreases in the numbers of some native species, and the increases of others, especially of introduced species, is probably linked to the vast changes in the terrestrial and aquatic habitats that have occurred in New Jersey over the last three centuries. The impacts of land clearing and resulting sedimentation, stream damming, stream channelization, industrialization, pollution (from industrial, residential and agricultural sources), and overfishing probably have significantly impacted some native species. However, since no relevant baseline data exist on these anthropogenic impacts, it is not possible to accurately assess them. More surveys and research must be conducted to effectively assess the effects of natural and anthropogenic factors on the distribution and abundance of the freshwater fishes in New Jersey.
Table 1. List of native, extipated, and currently existing and long
occuring introduced (*) species, in phylogenetic order, considered
herein to comprise the freshwater fish fauna of New Jersey.

Lampreys Family Petromyzontidae

American brook lamprey Lampetra appendix
Sea lamprey Petromyzon marinus

Sturgeons Family Acipenseridae

Shortnose sturgeon Acipenser brevirostrum
Atlantic sturgeon Acipenser oxyrhynchus

Gars Family Lepisosteidae

Longnose gar Lepisosteus osseus

Bowfins Family Amiidae

Bowfin Amia calva

Freshwater Eels Family Anguillidae

American eel Anguilla rostrata

Herrings Family Clupeidae

Blueback herring Alosa aestivalis
Hickory shad Alosa mediocris
Alewife Alosa pseudoharengus
American shad Alossa sapidissima
Gizzard shad Dorosoma cepedianum

Carps & Minnows Family Cyprinidae

Goldfish * Carassius auratus
Grass carp * Ctenopharyngodon idella
Satinfin shiner Cyprinella analostana
Spotfin shiner Cyprinella spiloptera
Common carp * Cyprinus carpio
Cutlip minnow Exoglossum maxillingua
Eastern silvery minnow Hybognathus regius
Common shiner Luxilus cornutus
Golden shiner Notemigonus crysoleucas
Comely shiner Notropis amoenus
Bridle shiner Notropis bifrenatus
Ironcolor shiner Notropis chalybaeus
Spottail shiner Notropis hudsonius
Swallowtail shiner Notropis procne
Bluntnose minnow * Pimephales notatus
Fathead minnow * Pimephales promelas
Blacknose dace Rhinichthys atratulus
Longnose dace Rhinichthys cataractae
Creek chub Semotilus atromaculatus
Fallfish Semotilus corporalis

Suckers Family Catostomidae

Quillback Carpiodes cyprinus
White sucker Catostomus commersonnii
Creek chubsucker Erimyzon oblongus
Northern hog sucker Hypentelium nigricans

N Amer. Catfishes Family Ictaluridae

White catfish Ameiurus calus
Black bullhead * Ameiurus melas
Yellow bullhead Ameiurus natalis
Brown bullhead Ameiurus nebulosus
Channel catfish Iclalurus punctatus
Tadpole madtom Noturus gyrinus
Margined madtom Noturus insignis

Pikes Family Esocidae

Redfin pickerel Esox americanus
Northern pike * Esox lucius
Muskellunge Esox masquinongy
Chain pickerel Esox niger

Mudminnows Family Umbridae

Eastern mudminnow Umbra pygmaea

Smelts Family Osmeridae

Rainbow smelt Osmerus mordax

Trouts & Salmons Family Salmonidae

Rainbow trout * Oncorhynchus mykiss
Brown trout * Salmo trutta
Brook trout Salvelinus fontinalis
Lake trout * Salvelinus namaycush

Pirate Perches Family Aphredoderidae

Pirate perch Aphredoderus sayanus

Cods Family Gadidae

Atlantic tomcod Microgadus tomcod

New World Silversides Family Atherinopsidae

Inland silverside Menidia beryllina

Topminnows Family Fundulidae

Banded killifish Fundulus diaphanus
Mummichog Fundulus heteroclitus

Livebearers Family Poeciliidae

Western mosquitofish * Gambusia affinis
Eastern mosquitofish Gambusia holbrooki

Sticklebacks Family Gasterosteidae

Fourspine stickleback Apeltes quadracus
Threespine stickleback Gasterosteus aculeatus
Ninespine stickleback Pungitius pungitius

Sculpins Family Cottidae

Slimy sculpin Cottus cognatus

Temperate Basses Family Moronidae

White perch Morone americana
Striped bass Morone saxatilis

Sunfishes Family Centrarchidae

Mud sunfish Acantharchus pomotis
Rock bass * Ambloplites rupestris
Blackbanded sunfish Enneacanthus chaetodon
Bluespotted sunfish Enneacanthus gloriosus
Banded sunfish Enneacanthus obesus
Redbreast sunfish Lepomis auritus
Green sunfish * Lepomis cyanellus
Pumpkinseed Lepomis gibbosus
Warmouth Lepomis gulosus
Bluegill * Lepomis macrochirus
Smallmouth bass * Micropterus dolomieu
Largemouth bass * Micropterus salmoides
White crappie * Pomoxis annularis
Black crappie * Pomoxis nigromaculatus

Perches Family Percidae

Swamp darter Etheostoma fusiforme
Tessellated darter Etheostoma olmstedi
Yellow perch Perca flavescens
Shield darter Percina peltata
Walleye * Sander vitreum

American Soles Family Achiridae

Hogchoker Trinectes maculatus

Table 2. Annotated list of native, extirpated, and currently existing
and long occurring introduced (*) species, in phylogenetic order,
considered herein to comprise the freshwater fish fauna of New Jersey.
Taxonomy follows primarily Carter and Williams (2002), and Page and
Burr (1991) for those species not listed in the former. Authors and
years of publication of scientific names follow Robins et al. (1991).

Class CEPHALASPIDOMORPHI--LAMPREYS

Order Petromyzontiformes

Family Petromyzontidae--Lampreys 2 species

Lampetra appendix (DeKay, 1842). American brook lamprey (Figure 7)
Freshwater; native; widespread in the Atlantic, Delaware River and
Delaware Bay drainages; larvae (=ammocoetes) and adults occasional.

Petromyzon marinus Linnaens, 1758. Sea lamprey (Figure 8)
Freshwater, brackish, marine, anadromous; native; largely coastal
and widespread in the Atlantic, Delaware River, Delaware Bay and
Hudson River drainages; occurs in the Pinelands, but is marginal
there; larvae (=ammocoetes) and adults occasional.

Class ACTINOPTERYGII--RAY-FINNED FISHES

Order Acipenseriformes

Family Acipenseridae--Sturgeons 2 species

Acipenser brevirostrum Lesueur, 1818. Shortnose sturgeon (Figure 9)
Freshwater, brackish, occasionally marine; native; coastal in the
tidal portions of the Delaware River, upper Delaware Bay and Hudson
River; now an endangered species and protected by laws.

Acipenser oxyrhynchus Mitchill, 1814. Atlantic sturgeon (Figure 10)
Freshwater, brackish, marine: native; largely coastal in the Delaware
River, Delaware Bay and Hudson Rivers, and the inshore Atlantic Ocean;
formerly common but through overfishing and habitat degradation now
uncommon; protected by laws.

Order Lepisosteiformes

Family Lepisosteidae--Gars 1 species

Lepisosteus osseus (Linnaeus, 1758). Longnose gar (Figure 11)
Freshwater; native; formerly present in the Delaware River (and
perhaps portions of Delaware Bay) but now extirpated in New Jersey.

Order Amiiformes

Family Amiidae--Bowfins 1 species

Amia calva Linnaeus, 1766. Bowfin (Figure 12)
Freshwater; in Atlantic and Delaware River drainages, mostly in rivers
and at mouths of large tributary creeks, and in the lower Hudson
River; native?, introduced at some localities; a Mullica River record
appears highly unlikely but is documented by one adult specimen;
apparently rare, and secretive.

Order Anguilliformes

Family Anguillidae--Freshwater Eels 1 species

Anguilla rostrata (Lesueur, 1817). American eel (Figure 13)
Freshwater, brackish, marine, catadromous; native; coastal and
statewide in Atlantic, Delaware River and Delaware Bay drainages, back
bays and inshore Atlantic Ocean, and Hudson River; the paucity of
records from the north-central part of state at least partly reflects
the effect of dams and waterfalls on upstream migration; occurs in the
Pinelands; small juveniles to adults often abundant.

Order Clupeiformes

Family Clupeidae--Herrings 5 species

Alosa aestivalis (Mitchill, 1814). Blueback herring (Figure 14)
Marine, brackish, freshwater, anadromous; native; coastal, migrating
and spawning adults and larvae and juveniles seasonally widespread in
Atlantic, Delaware River and Delaware Bay drainages in rivers, and in
Hudson River, otherwise young to adults found in inshore Atlantic
Ocean; occurs in the Pinelands, but limited to lower portions of
rivers; often common and sometimes even abundant, but probably not as
much as formerly; receives protection through laws.

Alosa mediocris (Mitchell, 1814). Hickory shad (Figure 15)
Marine, brackish, freshwater, anadromous; native; coastal, widespread
in Atlantic, Delaware River, Delaware Bay and Hudson River drainages,
in rivers and large streams, and in inshore Atlantic Ocean; occurs in
the Pinelands, but known from there on the basis of very few records
from tidal portions of rivers; occasional in freshwater, often common
to abundant in the ocean.

Alosa pseudoharengus (Wilson, 1811). Alewife (Figure 16)
Marine, brackish, freshwater, anadromous; native; coastal, migrating
and spawning adults and larvae and juveniles seasonally widespread in
Atlantic, Delaware River, Delaware Bay and Hudson River drainages in
rivers, otherwise young to adults found in inshore Atlantic Ocean;
landlocked (lake) populations present in northern New Jersey; occurs
in the Pinelands, but only in tidal portions of rivers; often common
and even abundant, but probably not as much as formerly; receives
protection through laws.

Alosa sapidissima (Wilson, 1811). American shad (Figure 17)
Marine, brackish, freshwater, anadromous; native; coastal, migrating
and spawning adults and larvae and juveniles seasonally widespread in
Atlantic, Delaware River, Delaware Bay and Hudson River drainages in
rivers, in winter young to adults found in inshore Atlantic Ocean;
extirpated as a reproducing population in the Mullica River system;
sometimes common, but was formerly abundant; receives protection
through laws.

Dorosoma cepedianum (Lesueur, 1818). Gizzard shad (Figure 18)
Freshwater, brackish, marine; native; somewhat coastal, widespread in
Atlantic, Delaware River, Delaware Bay and Hudson River drainages in
rivers and large creeks and in inshore Atlantic Ocean; introduced into
some lakes; occurs in the Pinelands, on the basis of a few records
from the lower portions of two rivers; often common.

Order Cypriniformes

Family Cyprinidae--Carps and Minnows 20 species

* Carassius auratus (Linnaeus, 1758). Goldfish (Figure 19)
Freshwater; introduced, naturalized; widely distributed in Atlantic,
Delaware River and Hudson River drainages; native to Eurasia; often
common.

* Ctenopharyngodon idella (Valenciennes, 1844). Grass carp (Figure 20)
Freshwater; introduced; scattered few localities in Atlantic, Delaware
River and Delaware Bay drainages, and no doubt found at other
localities not shown; a permit is required to introduce it and only
non-reproductive animals are allowed in New Jersey, but there might
be some reproducing populations; native to Asia; sometimes locally
common.

Cyprinella analostana Girard, 1859. Satinfin shiner (Figure 21)
Freshwater; native; widespread in Atlantic and Delaware River
drainages; sometimes abundant.

Cyprinella spiloptera (Cope, 1868). Spotfin shiner (Figure 22)
Freshwater; native; fairly widespread in Atlantic and Delaware River
drainages; sometimes abundant.

* Cyprinus carpio Linnaeus, 1758. Common carp (Figure 23)
Freshwater; introduced, naturalized; widespread in Atlantic, Delaware
River, Delaware Bay and Hudson River drainages; native to Eurasia;
often common.

Exoglossum maxillingua (Lesueur, 1817). Cutlip minnow (Figure 24)
Freshwater; native; widespread in Atlantic and Delaware River
drainages in northern half of New Jersey: locally common.

Hybognathus regius Girard, 1856. Eastern silvery minnow (Figure 25)
Freshwater, sometimes in slightly brackish water; native; widespread
in Atlantic and Delaware River drainages; often abundant.

Luxilus cornutus (Mitchill, 1817). Common shiner (Figure 26)
Freshwater; native; widespread in Atlantic and Delaware River
drainages; often common.

Notemigonus crysoleucas (Mitchill, 1814). Golden shiner (Figure 27)
Freshwater, sometimes in slightly brackish water; native; widespread
in Atlantic, Delaware River, Delaware Bay and Hudson River drainages;
occurs in the Pinelands, but there largely limited to rivers and
man-made ponds; often abundant.

Notropis amoenus (Abbott, 1874). Comely shiner (Figure 28)
Freshwater; native; fairly widespread in Atlantic and Delaware River
drainages; occasional.

Notropis bifrenatus (Cope, 1869). Bridle shiner (Figure 29)
Freshwater; native; fairly widespread in Atlantic, Delaware River
and Delaware Bay drainages, but found at only a few and scattered
localities; now apparently occurs at fewer localities than formerly;
occasional to rare.

Notropis chalybaeus (Cope, 1869). Ironcolor shiner (Figure 30)
Freshwater; native; scattered records in Atlantic, Delaware River and
Delaware Bay drainages, but mostly in southern New Jersey; there has
apparently been a pronounced decrease in the number of populations;
occurs in the Pinelands, but at only a few localities, and it has been
extirpated in the Mullica River system; occurrence is probably
occasional, but there is at least one site where this species is
abundant.

Notropis hudsonius (Clinton, 1824). Spottail shiner (Figure 31)
Freshwater, and sometimes in slightly brackish water; native;
widespread in Atlantic, Delaware River and Delaware Bay drainages;
often common.

Notropis procne (Cope, 1865). Swallowtail shiner (Figure 32)
Freshwater; native; fairly widespread in Atlantic and Delaware River
drainages; often common.

* Pimephales notatus (Rafinesque, 1820). Bluntnose minnow (Figure 33)
Freshwater; introduced; recorded at scattered localities in Atlantic
and Delaware River drainages; native to much of eastern and central
United States and to adjacent parts of Canada; occasional.

* Pimephales promelas Rafinesque, 1820. Fathead minnow (Figure 34)
Freshwater; introduced; recorded at scattered localities in Atlantic
and Delaware River drainages, mostly in the northern half of New
Jersey; native to much of eastern and central United States and
adjacent parts of Mexico and Canada; occasional.

Rhinichthys atratulus (Hermann, 1804). Blacknose dace (Figure 35)
Freshwater; native; widespread in Atlantic and Delaware River
drainages; disjunct populations occur in southwestern New Jersey;
common to locally abundant.

Rhinichthys cataractae (Valenciennes, 1842). Longnose dace (Figure 36)
Freshwater; native; widespread in Atlantic and Delaware River
drainages in the northern half of the state; occasional to locally
common.

Semotilus atromaculatus (Mitchill, 1818). Creek chub (Figure 37)
Freshwater; native; widespread in Atlantic and Delaware River
drainages in the northern half of the state; occasional to locally
common.

Semotilus corporalis (Mitchill, 1817). Fallfish (Figure 38)
Freshwater; native; widespread in Atlantic and Delaware River
drainages, mostly in the northern part of state; in southeastern
Monmouth County occurs disjunct from rest of populations; occasional
to locally abundant.

Family Catostomidae--Suckers 4 species

Carpiodes cyprinus (Lesueur, 1817). Quillback (Figure 39)
Freshwater; native; recorded from the Delaware River and from one
locality in the Raritan River; apparently rare in New Jersey.

Catostomus commersonnii (Lacepede, 1803). White sucker (Figure 40)
Freshwater, sometimes in slightly brackish water; native; widespread
in Atlantic and Delaware River drainages; in the southeastern part of
state it occurs only in the larger rivers; occurs in the Pinelands,
but only in tidal freshwater of rivers; common to locally abundant.

Erimyzon oblongus (Mitchill, 1814). Creek chubsucker (Figure 41)
Freshwater; native; statewide in Atlantic, Delaware River and Delaware
Bay drainages; occurs in the Pinelands; occasional to common.

Hypentelium nigricans (Lesueur, 1817). Northern hog sucker (Figure 42)
Freshwater; native; Delaware River drainage; there are only a very few
New Jersey records, some of them many decades old; present data
indicate that it is rare in New Jersey.

Order Siluriformes

Family Ictaluridae--North American Catfishes 7 species

Ameiurus catus (Linnaeus, 1758). White catfish (Figure 43)
Freshwater, often in slightly brackish water; native; widespread in
Atlantic, Delaware River, Delaware Bay and Hudson River drainages,
mainly in rivers and large streams; introduced in other waters in New
Jersey; occurs in the Pinelands, there limited to the tidal portions
of rivers; often abundant.

* Ameiurus melas (Rafinesque, 1820). Black bullhead (Figure 44)
Freshwater; introduced; scattered records in Atlantic, Delaware River
and Delaware Bay drainages; native to much of eastern and central
United States and parts of adjacent Canada; apparently uncommon to
rare.

Ameiurus natalis (Lesueur, 1819). Yellow bullhead (Figure 45)
Freshwater; native; widespread in Atlantic, Delaware River and
Delaware Bay drainages: occurs in the Pinelands; usually occasional.

Ameiurus nebulosus (Lesueur, 1819). Brown bullhead (Figure 46)
Freshwater, sometimes in slightly brackish water; native; widespread
in Atlantic, Delaware River and Delaware Bay drainages; occasional to
common.

Ictalurus punctatus (Rafinesque, 1818). Channel catfish (Figure 47)
Freshwater, sometimes in slightly brackish water; probably native only
in the Delaware River in New Jersey; now widely distributed in
Atlantic, Delaware River and Delaware Bay drainages as a result of
extensive introductions; occurs in the Pinelands, where it has been
introduced in at least two localities; usually occasional.

Noturus gyrinus (Mitchill, 1817). Tadpole madtom (Figure 48)
Freshwater, and rarely in slightly brackish water; native; widespread
in Atlantic, Delaware River and Delaware Bay drainages; occurs in the
Pinelands; occasional to locally common.

Noturus insignis (Richardson, 1836). Margined madtom (Figure 49)
Freshwater; native; widespread in Atlantic and Delaware River
drainages, mostly in northern part of state; occasional to locally
common.

Order Esociformes

Family Esocidae--Pikes 4 species

Esox americanus Gmelin, 1788. Redfin pickerel (Figure 80)
Freshwater, and rare in slightly brackish water: native: widespread
statewide in Atlantic, Delaware River and Delaware Bay drainages,
occurs in the Pinelands; occasional to locally common.

* Esox lucius Linnaeus, 1758. Northern pike (Figure 51)
Freshwater; introduced; scattered records in Atlantic, Delaware River
and Delaware Bay drainages; native to much of eastern and central and
northwestern North America; occasional.

Esox masquinongy Mitchill, 1824. Muskellunge (Figure 52)
Freshwater; native in non-tidal Delaware River and perhaps its
tributaries in northwestern New Jersey; through introductions, now
scattered records in Atlantic, Delaware River and Delaware Bay
drainages, almost all in the northern half of New Jersey; occasional.

Esox niger Lesueur, 1818. Chain pickerel (Figure 53)
Freshwater, and not uncommonly found in brackish water; native:
widespread statewide in Atlantic, Delaware River and Delaware Bay
drainages; occurs in the Pinelands; occasional to abundant.

Family Umbridae--Mudminnows 1 species

Umbra pygmaea (DeKay, 1842). Eastern mudminnow (Figure 54)
Freshwater, and occasionally in slightly brackish water; native;
widespread in Atlantic, Delaware River and Delaware Bay drainages,
but apparently absent from much of the Raritan River and the
central-west part of the state; occurs in the Pinelands; common to
abundant.

Order Salmoniformes

Family Osmeridae--Smelts 1 species

Osmerus mordax (Mitchill, 1814). Rainbow smelt (Figure 55)
Marine, brackish, freshwater, anadromous; native; largely coastal,
occurs in the Atlantic drainage in the lower Hackensack, Hudson,
Passaic and Raritan Rivers and in the lower Delaware River during
spawning migrations, and otherwise found in inshore Atlantic Ocean,
New York Harbor estuary, and probably in Delaware Bay; formerly
abundant, now occasional to rare.

Family Salmonidae--Trouts and Salmons 4 species

* Oncorhynchus mykiss (Walbaum, 1792). Rainbow trout (Figure 56)
Freshwater; introduced; widespread in Atlantic, Delaware River and
Delaware Bay drainages, especially in northern portion of the state;
most records represent stockings, although there is limited
reproduction in the wild; native to far western North America;
occasional.

* Salmo trutta Linnaeus, 1758. Brown trout (Figure 57)
Freshwater, rare in saltwater; introduced; widespread in Atlantic,
Delaware River and Delaware Bay drainages, especially in northern
portion of the state; most records represent stockings, although
there is limited reproduction in the wild; native to Eurasia;
occasional.

Salvelinus fontinalis (Mitchill, 1814). Brook trout (Figure 58)
Freshwater; native; widespread in Atlantic, Delaware River and
Delaware Bay drainages, mostly in the northern part of the state;
natives occur north of the latitude of the mainstem Raritan River and
in one stream near Camden; the species is maintained by stockings at
many localities in northern New Jersey, near metropolitan New Jersey,
and in southern New Jersey, but excluding those localities where there
is reproduction by natives; occasional.

* Salvelinus namaycush (Walbaum, 1792). Lake trout (Figure 59)
Freshwater; introduced; in Atlantic and Delaware River drainages at
three localities in northern New Jersey; reproduction occurs in New
Jersey; native to almost all of North America south to southern New
York State; occasional.

Order Percopsiformes

Family Aphredoderidae--Pirate Perches 1 species

Aphredoderus sayanus (Gilliams, 1824). Pirate perch (Figure 60)
Freshwater; native; widespread in Atlantic, Delaware River and
Delaware Bay drainages south of the latitude of the mainstem Raritan
River; occurs in the Pinelands; occasional.

Order Gadiformes

Family Gadidae--Cods 1 species

Microgadus tomcod (Walbaum, 1792). Atlantic tomcod (Figure 61)
Marine, brackish water, freshwater; native; coastal, in the lower
Hudson, Hackensack, and Passaic Rivers and New York Harbor area and
the lower Hudson River, where it spawns, and there are records from
the lower Delaware and Raritan rivers and Barnegat gay; sometimes
locally common in the New York Harbor area and in the lower Hudson
River, and rare (now absent?) in southern New Jersey waters.

Order Atheriniformes

Family Atherinopsidae--New World Silversides 1 species

Menidia beryllina (Cope, 1866). Inland silverside (Figure 62)
Brackish water, freshwater, marine: native; widespread along the
coast in Atlantic, Delaware River and Delaware Bay drainages and in
the tidal portions of rivers; occurs in the Pinelands in tidal waters;
occasional to common.

Order Cyprinodontiformes

Family Fundulidae--Topminnows 2 species

Fundulus diaphanus (Lesueur, 1817). Banded killifish (Figure 63)
Freshwater, and often in brackish water; native; largely coastal,
widespread in Atlantic, Delaware River and Delaware Bay, drainages;
some of the records on the immediate Atlantic shore are perhaps of
misidentified other killifishes, especially of the mummichog; occurs
in the Pinelands in some tidal waters; often common or abundant.

Fundulus heteroclitus (Linnaeus, 1766). Mummichog (Figure 64)
Brackish water, freshwater, marine; coastal, it is mostly estuarine
and often in tidal freshwaters, but numbers also occur in some
non-tidal freshwaters; native; widespread in Atlantic, Delaware River
and Delaware Bay drainages; isolated locality dots in lakes and ponds
in Sussex and Morris counties, and some in southern New Jersey,
represent introduced individuals; occurs in the Pinelands in tidal
waters or adjacent; often abundant.

Family Poeciliidae--Livebearers 2 species

* Gambusia affinis (Baird and Girard, 1853). Western mosquitofish
(Figure 65) (and see also next account) Freshwater, often in
brackish; introduced; scattered in Atlantic, Delaware River and
Delaware Bay drainages; probably at more localities than shown through
introductions by New Jersey counties mosquito control commissions;
native to south-central United States; often common. (All New Jersey
Gambusia I have identified have been of this species. The species may
persist through continued introduction, through reproduction in the
wild, or both.)

Gambusia holbrooki Girard, 1859. Eastern mosquitofish (No Figure)
Freshwater, often in brackish; native?; probably occurs at scattered
localities throughout New Jersey, reported as long introduced by New
Jersey counties' mosquito control commissions; native to central and
southeastern United States north to Delaware, and perhaps in New
Jersey; often abundant in Delaware and south. (It is uncertain which
species (singular or plural) occur(s) in New Jersey. This is because
specimens were first introduced into New Jersey about 100 years ago,
and then subsequently, and before its natural occurrence or lack of
it here had been established. A recent taxonomic change, in which two
subspecies of the species affinis have been elevated to the status of
species, adds to the confusion. Native populations, if there were
any, probably continue to be heavily augmented by stockings of captive
bred populations which appear to have been derived from a number of
original out-of-state sources over the years. However, I have not
yet identified specimens from New Jersey.)

Order Gasterosteiformes

Family Gasterosteidae--Sticklebacks 3 species

Apeltes quadracus (Mitchill, 1815). Fourspine stickleback (Figure 66)
Brackish water, saltwater, freshwater: native: coastal in Atlantic,
Delaware River and Delaware Bay drainages, in tidal Delaware River as
far upstream as Trenton area; occasional to abundant.

Gasterosteus aculeatus Linnaeus, 1758. Threespine stickleback (Figure
67) Brackish water, inshore marine, freshwater; native; coastal in
Atlantic, Delaware River and Delaware Bay drainages, in tidal Delaware
River as far upstream as Trenton area; occasional.

Pungitius pungitius (Linnaeus, 1758). Ninespine stickleback (Figure
68) Brackish water and adjacent freshwater; recorded at five
localities on New Jersey Atlantic coast, from Cape May Peninsula north
to New York Harbor; apparently rare.

Order Scorpaeniformes

Family Cottidae--Sculpins 1 species

Cottus cognatus Richardson, 1836. Slimy sculpin (Figure 69)
Freshwater: native; in Atlantic and Delaware River drainages in
northwestern New Jersey; occasional, sometimes common.

Order Perciformes

Family Moronidae--Temperate Basses 2 species

Morone americana (Gmelin, 1789). White perch (Figure 70)
Primarily brackish water, freshwater, occasionally marine; native;
widespread coastal in Atlantic, Delaware River and Delaware Bay
drainages; in rivers and larger creeks, especially in their tidal
lower portions, and inshore Atlantic Ocean: landlocked lake
populations are introduced; in Pinelands in tidal waters; often
abundant.

Morone saxatilis (Walbaum, 1792). Striped bass (Figure 71)
Primarily marine, brackish water, freshwater; native; widespread
coastal in Atlantic, Delaware River and Delaware Bay drainages; in
rivers and larger creeks, especially their lower portions, and in
inshore Atlantic Ocean; in Pinelands in tidal waters; often common.

Family Centrarchidae--Sunfishes 14 species

Acantharchus pomotis (Baird, 1855). Mud sunfish (Figure 72)
Freshwater; native; widespread in Atlantic, Delaware River and
Delaware Bay drainages, especially south of latitude of mainstem
Raritan River; in Pinelands; occasional.

* Ambloplites rupestris (Rafinesque, 1817). Rock bass (Figure 73)
Freshwater; introduced; widespread in Atlantic and Delaware River
drainages, in northern half of state; native to much of central and
eastern United States and adjacent Canada; occasional to common.

Enneacanthus chaetodon (Baird, 1855). Blackbanded sunfish (Figure 74)
Freshwater; native; widespread in Atlantic and Delaware River and
Delaware Bay drainages south of mainstem Raritan River; in Pinelands;
occasional to common.

Enneacanthus gloriosus (Holbrook, 1855). Bluespotted sunfish (Figure
75) Freshwater; native; widespread in Atlantic, Delaware River and
Delaware Bay drainages; in Pinelands; occasional to common.

Enneacanthus obesus (Girard, 1854). Banded sunfish (Figure 76)
Freshwater; native; widespread in Atlantic, Delaware River and
Delaware Bay drainages, especially in southern half of state;
in Pinelands; occasional to common.

Lepomis auritus (Linnaeus, 1758). Redbreast sunfish (Figure 77)
Freshwater; native; widespread in Atlantic, Delaware River and
Delaware Bay drainages, especially throughout northern half of the
state; Atlantic drainage localities south of mainstem Raritan River
probably the result of introductions; in Pinelands probably
introduced; occasional to common.

* Lepomis cyanellus Rafinesque, 1819. Green sunfish (Figure 78)
Freshwater; introduced; widespread in Atlantic and Delaware River
drainages, mostly in northern half of state; native to much of central
and eastern United States and adjacent Mexico and Canada; occasional
to common.

Lepomis gibbosus (Linnaeus, 1758). Pumpkinseed (Figure 79)
Freshwater; native; statewide in Atlantic, Delaware River and Delaware
Bay drainages, although perhaps more widespread than formerly as a
result of intrastate introductions; in Pinelands, but absent from
"deep Pinelands"; occasional to abundant.

Lepomis gulosus (Cuvier, 1829). Warmouth (Figure 80)
Freshwater; native?; one known population in Salem County, one record
from Delaware River, and one other Delaware River specimen with no
locality; native to much of central and eastern United States to the
northern Delmarva Peninsula; rare.

* Lepomis macrochirus Rafinesque, 1819. Bluegill (Figure 81)
Freshwater; introduced; statewide in Atlantic, Delaware River and
Delaware Bay drainages; in Pinelands, but absent from "deep
Pinelands"; native to much of eastern and central United States and
adjacent Mexico and Canada; occasional to abundant.

* Micropterus dolomieu Lacepede, 1802. Smallmouth bass (Figure 82)
Freshwater; introduced; widespread in Atlantic, Delaware River and
Delaware Bay drainages in northern half of state, a few records from
the southern portion; native to much of north-central United States
and adjacent Canada; occasional to common.

* Micropterus salmoides (Lacepede, 1802). Largemouth bass (Figure 83)
Freshwater; introduced; statewide in Atlantic. Delaware River and
Delaware Bay drainages; in Pinelands, although absent from "deep
Pinelands"; native to much of central United States and adjacent
Mexico and Canada; often common.

* Pomoxis annularis Rafinesque, 1818. White crappie (Figure 84)
Freshwater, seldom in very slightly brackish water; introduced;
scattered in Atlantic, Delaware River and Delaware Bay drainages; much
less widely distributed and common in New Jersey than black crappie;
native to much of eastern and central United States and adjacent
Mexico and Canada; occasional.

* Pomoxis nigromaculatus (Lesueur, 1829). Black crappie (Figure 85)
Freshwater; introduced; widespread in Atlantic, Delaware River and
Delaware Bay drainages; in Pinelands, but limited; native to much of
eastern and central United States and adjacent Canada; occasional.

Family Percidae--Perches 5 species

Etheostoma fusiforme (Girard, 1854). Swamp darter (Figure 86)
Freshwater; native; widespread in Atlantic, Delaware River and
Delaware Bay drainages south of latitude of mainstem Raritan River;
in Pinelands; often abundant.

Etheostoma olmstedi Storer, 1842. Tessellated darter (Figure 87)
Freshwater, occasionally in brackish water; native; almost statewide
in Atlantic, Delaware River, Delaware Bay and Hudson River drainages;
in Pinelands, but there largely limited to rivers; often abundant.

Perca flavescens (Mitchill, 1814). Yellow perch (Figure 88)
Freshwater, occasionally in slightly brackish water; native; almost
statewide in Atlantic, Delaware River and Delaware Bay drainages; in
Pinelands, but there largely limited to rivers; sometimes common.

Percina peltata (Stauffer, 1864). Shield darter (Figure 89)
Freshwater; native; in mainstem non-tidal Delaware River and lower
portions of some of its tributaries, and in portions of upper Raritan
River system; occasional.

* Sander vitreum (Mitchill, 1818). Walleye (Figure 90)
Freshwater; introduced; fairly widespread in Atlantic and Delaware
River drainages, mostly in northern half of New Jersey, in southern
half of state a few records from Delaware River and its tributaries;
native to much of eastern and central North America; occasional.

Order Pleuronectiformes

Family Achiridae--American Soles 1 species

Trinectes maculatus (Bloch and Schneider, 1801). Hogchoker (Figure 91)
Freshwater, brackish water, marine; native; along Atlantic coast and
lower portions of tributary rivers and back bays from Hudson River to
tip of Cape May Peninsula, throughout Delaware Bay and lower portions
of its tributary rivers, and in tidal Delaware River; in Atlantic
Ocean to distances offshore; in Pinelands, but there limited to tidal
portions of rivers; sometimes common.

Table 3. List of 49 native and apparently established introduced
freshwater fish species recorded from the Pinelands of New Jersey
(Figure 5), in phylogenetic order by category. Marginal = found at
very few sites, usually on the edge of the Pinelands; In Rivers =
found exclusively or almost exclusively in rivers in the Pinelands;
In Rivers and Scattered = found mostly in rivers in the Pinelands,
but also at a number of other sites throughout the Pinelands;
Widespread and Often Common = found widespread in the Pinelands, and
often with many individuals at a locality; Scattered = found at few
and widely scattered sites in the Pinelands. (I = introduced to
Pinelands; few recs. = very few specimens recorded.)

MARGINAL IN RIVERS

sea lamprey bowfin (I?)
goldfish (I) blueback herring (few recs.)
grass carp (I) hickory shad (very few recs.)
common carp (I) alewife
fallfish American shad (?, very few recs.)
black bullhead (I) gizzard shad (very few recs.)
brook trout white sucker
western mosquitofish (I) white catfish
eastern mosquitofish (occurs?, I?) inland silverside
white crappie banded killifish
 mummichog
 fourspine stickleback
 white perch
 striped bass
 hogchoker

 IN RIVERS AND WIDESPREAD AND
 SCATTERED OFTEN COMMON SCATTERED

golden shiner American eel ironcolor shiner
brown bullhead creek chubsucker channel catfish (I)
tessellated darter yellow bullhead redbreast sunfish (I?)
yellow perch tadpole madtom
black crappie (I) redfin pickerel
 chain pickerel
 eastern mudminnow
 pirate perch
 mud sunfish
 blackbanded sunfish
 bluespotted sunfish
 banded sunfish
 pumpkinseed
 bluegill (I)
 largemouth bass (I)
 swamp darter


ACKNOWLEDGMENTS

This paper represents some five years of work, with much help from others, to locate data and to plot localities of fish occurrence. Obviously, this paper could not have been written without their cooperation. I am indebted to the curators, collection managers, professors, biologists, environmentalists, and computer specialists named in the preceding paragraphs for their valuable data and help. It is only in the interest of brevity that I do not repeat their names here.

I am also indebted to V. K. Farris, former president of The Richard Stockton College of New Jersey, who generously provided release time from my teaching duties and additional financial support for this research, and who created a pleasant atmosphere which made it possible for me to pursue this laborious task. Members of the Stockton College Board of Trustees are thanked for endorsing the release time recommendation of President Farris.

The following New Jersey Department of Environmental Protection personnel (in alphabetical order) likewise are thanked for the information they provided: R. Babb, T. Baum, M. Boriek, S. Crouse, P. L. Hamilton, P. Himchak, D. Keller, J. Matthews, W. Murawsky, S. G. Piotrowski, C. Smith, and R. Soldwedel. Data were also kindly provided by the following individuals (in alphabetical order): J. H. Balletto, Public Service Electric and Gas Company, New Jersey; R. Blye, Pennsylvania Audubon Society; J. F. Bunnell, Jr. and R. Zampella, Pinelands Commission, New Jersey; D. Carlson, New York State Department of Environmental Conservation; M. Danko, New Jersey Marine Sciences Consortium; M. H. Doosey, Tulane University; S. Eisenhauer, Natural Lands Trust, New Jersey; M. Fabrizio and J. Pessutti as well as the scientists and technicians of the Behavioral Ecology Branch, Northeast Fisheries Science Center, NOAA-Fisheries, Sandy Hook, New Jersey; P. Harmon, environmental consultant; D. Littlehale, New Jersey State Aquarium; R. Miller, Delaware Fish and Game; L. T. Penny, Easthamton Township, Long Island, New York; and M. Vavra, National Park Service, Pennsylvania.

The following persons (in alphabetical order) were also helpful: F. and J. Akers, The Great Egg Harbor Watershed Association, New Jersey, assisted in the field; S. L. Allen, Port Republic, New Jersey, proofed parts of the manuscript; A. B. Bragin, answered many questions and provided references; D. Dorfman, Monmouth University, New Jersey, provided specimens; P. Feldbauer, Stockton College, provided computer help; Z. M. G. S. Jahangir, Kingsborough Community College, Brooklyn, New York, assisted in the field; C. Macfarlan-Dickson, New Jersey, proofed parts of the manuscript; J. C. O'Herron, Jr., New Jersey, environmental consultant, loaned publications; R. Horwitz, Academy of Natural Sciences of Philadelphia, provided general information; K. L. Kenley, Stockton College graduate, organized data; M. Mihalasky, Stockton College, provided computer help; A. Modjesky, New Jersey, provided reference material; F. C. Robde, Division of Marine Fisheries, North Carolina, provided a copy of his M. S. thesis; and M. Sebetich, William Paterson University, New Jersey, assisted in the field.

Many Stockton College students/graduates assisted me in the field, and there are too many to acknowledge them individually. However, several were especially helpful including (in alphabetical order): A. B. Bragin, who also provided comfortable lodging on some field trips; E. W. Gaine; A. M. Teti, who also helped with computer work; and C. Woolcott, whose family also provided comfortable lodging on some field trips.

Stockton College students/graduates who searched through data sources and entered locality data on paper base maps are (in alphabetical order):

R. Denton, A. Franz, E. W. Gaine, B. Hixon, J. P. Hooper, C. Hostetter, S. Hurder, J. Judas, B. Kirkland, M. Pickering, S. Piotrowski, K. Schaffer, and D. Zima. Stockton College students/graduates who entered data into a computer GIS arc-view program to produce the final figures are E. W. Gaine, J. Judas, and M. Pickering; C. Giordano, a Rutgers University graduate, also assisted in this effort.

W. Fan, Stockton College, produced the state-base drainage figure on which fish localities are plotted by combining county drainage maps prepared by the New Jersey Department of Environmental Protection. She also kindly answered many of my questions about GIS arc-view.

New Jersey state scientific collecting permits were provided over the years by W. J. Eisele, Jr., New Jersey Bureau of Marine Water Classification and Analysis; B. A. Halgren, New Jersey Marine Fisheries; T. McCloy and M.J. McHugh, New Jersey Division of Fish and Wildlife; and R. A. Cookingham and R. McDowell, New Jersey Department of Environmental Protection. D. L. Beall, T. Casselman, and A. Scherer of the U.S. Fish and Wildlife Service, Forsythe National Wildlife Refuge, Oceanville, New Jersey, issued special-use permits. L. S. Hales, Jr., now with the U.S. Fish and Wildlife Service, included me on a research permit issued to him.

I thank three anonymous reviewers for their carefully considered and helpful comments, and M. J. Kennish for his work as editor.

Any contributor to this project not acknowledged here is assured that this is due only to my oversight.

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RUDOLF G. ARNDT

DIVISION OF NATURAL SCIENCES AND MATHEMATICS

THE RICHARD STOCKTON COLLEGE OF NEW JERSEY

POMONA, NEW JERSEY 08240-0195

Professor Arndt can be contacted at Rudolf.Arndt@stockton.edu.
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