SYSTEMATICS OF SOUTHERN FLORIDA (PISCES: ENGRAULIDAE)1

RICHARD J. DALY Bureau of Commercial Fisheries, Tropical Atlantic Biological Laboratory, Miami, Florida 33149

ABSTRACT Counts of gill rakers, vertebrae, and dorsal-, anal-, and pectoral-fin rays of the five common of the family Engraulidae in southern Florida were analyzed and found to vary little within the species. Pigmentation and proportional characters are described and compared, and a key to speci- mens measuring 40 to 70 mm SL is presented. The five species- lamprotaenia Hildebrand, (Linnaeus), Anchoa nasuta Hildebrand & Carvalho, (Valenciennes), and Anchoviella perfasciata (Poey)-are discontinuously distributed in southern Florida and reach great seasonal abundance. The fishes I observed fed in currents at night and formed large schools (com- posed of a mixture of species) in protected areas to escape predation during the day.

INTRODUCTION Since Hildebrand (1943) reviewed the American anchovies, no major systematic studies have been made of this important family in eastern North America. Hildebrand's revisionary study was republished in part, with changes (principally editorial) by George S. Myers in "Fishes of the West- ern North Atlantic" in 1964, and reviewed briefly by Berry (1964). The present study was undertaken to evaluate variation among species of Engraulidae commonly collected in southern Florida, and to determine the value of various characters in distinguishing species of anchovies within a restricted geographic range. The particular area to which this study was restricted extends from Fort Pierce to Naples, Florida, and includes the Florida Keys. Anchovies are extremely abundant throughout most of southern Florida's coastal waters. Although not now of commercial importance locally, fishes of this family are used throughout the world for human consumption and in fish meal and similar products. The five species considered in this paper are the dominant species by far in southern Florida, but occasionally other engraulid species were collected during my study, especially on the fringes of the area described. These species are listed under "Additional Species" and are also included in the key. Selected specimens of the five common

1 Contribution No. 103, Bureau of Commercial Fisheries. Tropical Atlantic Biological Laboratory, Miami, Florida 33149; and Contribution No. 1150 from the University of Miami, Rosenstiel School of Marine and Atmospheric Sciences. The results of this investigation were submitted to the faculty of the Graduate School of the University of Miami in partial fulfillment of the requirements for the degree of Master of Science. 1970] Daly: Southern Florida Anchovies 71 species were examined from areas outside southern Florida to make the study of intraspecific variation more comprehensive.

METHODS Measurements were made with dividers and a millimeter ruler, and are expressed as a percentage of standard length (SL). All counts were re- peated at least once. The terms used are discussed below. 1. Dorsal and anal rays: All rays were counted, including the anterior two, which are much shorter than the third. In both fins, the posterior ray is divided to its base, but is counted as one ray. 2. Pectoral rays: The first ray is counted as one, even though it often ap- pears to be composed of two elements. 3. Gill rakers: Rakers were counted on the first arch without removing it from the fish. These counts are presented as a gill-raker formula, which consists of the range for the upper limb plus the range for the lower limb. 4. Vertebrae: Vertebral counts were made from radiographs, except for some collections of Anchoa lamprotaenia and Anchoa hepsetus in which vertebral number was determined by dissection. All vertebrae were counted, including the hypural plate. 5. Pelvic rays: The rays of the right pelvic fin were counted in 445 speci- mens. This fin is often missing or malformed; one specimen had four com- plete pelvic fins. 6. Standard length (SL): Measured frotTI the tip of the snout to the pos- terior edge of the hypural plate. 7. Head length: Taken along the axis of the body from the tip of the snout to a perpendicular through the most posterior point on the bony opercular margin. 8. Body depth: The greatest vertical depth of the body. 9. Snout length: Taken along the body axis from the tip of the lower jaw to a perpendicular from the tip of the snoUt. 10. Eye diameter: The greatest horizontal distance across the exposed por- tion of the orbit. 11. Width of lateral silver stripe: Taken above the center of the anal-fin base. 12. Longest dorsal ray: The distance from the base to the extremity of the longest ray. 13. Location of anal-fin origin: The relative position of the dorsal and anal fins was determined by a perpendicular to the axis of the body at the anal- fin origin. Specimens were fixed in 10 per cent formalin solution buffered with hexamethylarnine and transferred to very dilute formalin or to 40 per cent isopropanol after about 7 days. Specimens left in 10 per cent formalin longer than 2 weeks deteriorate rapidly and often lose all pigmentation. Most of the collections were made with a 4-foot cast net and a 30-foot shore seine, or a dip net and an attracting light of 60 to 500 watts. The following key is limited in its application to specimens ranging from 40 to 70 mm SL. This limitation allows comparison of characters that change during growth. 72 Bulletin of Marine Science [20(1 )

FIGURE 1. Anchoa lamprotaenia Hildebrand, UMML 20186, 55.3 mm SL.

KEY TO SOUTHERN FLORIDA SPECIES OF ENGRAULIDAE 1. Maxilla prolonged past angle of jaw to opercular margin 4 Maxilla not prolonged past angle of jaw to opercular margin 2 2. Head short, less than 25 per cent SL Anchoviella perfasciata Head long, greater than 25 per cent SL .______3 3. Lower gill rakers 29 or fewer, upper 25 or fewer; head less than 28 per cent SL .._. Engraulis eurystole Lower gill rakers 30 or more; upper 26 or more; head more than 28 per cent SL . ._._._.. Engraulis estauquae 4. Anal-fin origin under anterior half of dorsal-fin base; snout short, less than 4.5 per cent SL .__._ . ._.__. . ._. 5 Anal-fin origin under or behind center of dorsal-fin base; snout moderate to long, usually greater than 4.5 per cent SL __. ._ 6 5. Anal-fin rays 24 or fewer; body slender, less than 18 per cent SL . . ._..__._. . ._. . A nchoa cubana Anal-fin rays 25-30; body deep, greater than 19 per cent SL . . A nchoa mitchilli 6. Snout long, greater than 6.5 per cent SL . . .._ 7 Snout moderate in length, less than 6.5 per cent SL 8 7. Lower gill rakers 24 or more ._.. ..__. .__Anchoa nasuta Lower gill rakers 23 or fewer _. .__. . A nchoa lyolepis 8. Lower gill rakers 18 or more . . . .______9 Lower gill rakers 17 or fewer ...... _._A nchoa cayorum 1970] Daly: Southern Florida Anchovies 73 9. Upper gill rakers 17 or fewer, pectoral rays 14 or fewer; lateral silvery stripe slightly expanded over anal fin A nchoa lampr0taenia Upper gill rakers 17 or more, pectoral rays sometimes 14 but usually more than 14; lateral silvery stripe not noticeably expanded over anal fin Anchoa hepsetus

Genus Anchoa Jordan & Evermann Anchoa Jordan & Evermann, 1927: 501 (type-species by original designation, Engraulis compressus Girard, 1858: 336, San Diego, California).-Hilde- brand, 1943: 29 (redefined; synonymy); 1964: 162. Hildebrand distinguished the from Anchoviella by the long, pointed maxilla extending well past the angle of the jaw. Anchoa lamprotaenia Hildebrand Figs. 1, 6 Anchoa lamprotaenia Hildebrand, 1943: 62-64, fig. 23 (Type: USNM 117661, 65 mm SL, Key West, Florida); 1964: 192-194, fig. 39. Description.-CouNTS (meristic data are summarized in Tables 2-7): Dor- sal rays 14-16; anal rays 21-26; pectoral rays 12-15; vertebrae 39-42; gill rakers 14-17 + 18-21. MORPHOMETRY(10 specimens 53 to 61 mm SL, collected at Virginia Key, Miami, Florida, UMML 20186): Head large (27-30), body fairly deep (21-23). Snout long (5.4-6.1), eye large (7.6-9.5). Lateral silver stripe expanded (width 4.9-6.0) over anal fin. Anterior rays of depressed dorsal reaching past posterior rays. Fins long (longest dorsal ray 18-21). Vent closer to depressed pelvic rays than to anal fin. Anal-fin origin under eleventh or twelfth dorsal ray. Gill rakers widely spaced (about 0.5 mm apart) and tapering rapidly (width at midpoint % of width at base). PIGMENTATION:In life, the ground color is a translucent gray on the snout and dorsal aspect; the opercular area, eye, jaw, and scales are silvery. In contrast, a preserved specimen devoid of scales is usually gray or yel- lowish with guanine on the eye, opercular area, jaw, and in a lateral stripe reaching the base of the caudal fin. The peritoneum is also silvery. In the living , iridescent reds, blues, yellows, and greens are visi- ble on top of the head and in a thin line above the lateral stripe. The dor- sum of the orbits is golden when viewed through the transparent flesh. Posterior to the orbits, a large green and gold patch shields the brain dor- sally; behind this lies a small violet patch over the occipital region of the skull. The snout is a noniridescent yellow. After fixation, tracts of melano- phores underlying the iridescent areas become evident. Large melanophores in a bandlike arrangement impart a dusky appear- 74 Bulletin ot Marine Science [20( 1) TABLE 1 CHARACTERS USEFUL IN SEPARATING 50- TO 60-MM SPECIMENS OF Anchoa lamprotaenia AND A. hepsetus

Character A nc/1oa lamprotaenia A nchoa hepsetlls Lateral silver stripe Expanded over anal fin Fairly uniform in width Position of the vent Nearer to depressed pelvics Nearer to anal ongm Melanophore size Easily visible as dots Very fine, hard to see with unaided eye Dorsal melanophores Irregular oblique rows or Outline scale pockets if bands, if present present Melanophores on No median separation Me~lian separation at this the nape size Melanophores above Form O.5-mm-wide line Absent silver stripe anteriorly Gonadal condition1 Gonads obvious, eggs Gonads difficult to discern visible in ovary

1 During any season of the year. ance to the dorsal aspect of living and preserved specimens (Fig. 6). These dorsal melanophores are arranged in oblique rows proceeding from a thin line of melanophores running the length of the back. Large melanophores underlying the lateral silver stripe usually show above the stripe as a dark line about 0.5 mm wide. Heavy stippling above the skull continues for- ward in a band around the snout to the forward ends of the premaxillae. A dark area of continuous pigmentation appears on the dorsum of the or- bits and also on the skull bones shielding the brain. A dense concentration of melanophores on the nape is barely separated from the dark patch on the skull. Melanophores are also present along the rays of the dorsal and caudal fins and around the bases of the dorsal and anal fins. Comparisons.-Hildebrand (1943) noted that this species is often misiden- tified as Anchoa hepsetus. Adults of lamprotaenia usually range from 50 to 65 mm SL, whereas adults of hepsetus reach 85 mm SL or more at ma- turity. Specimens of hepsetus commonly fall within the 50- to 60-mm size range, however, and may be separated most readily from lamprotaenia by a combination of meristic characters (Tables 2-7). Additional distin- guishing characteristics are given in Table 1. Distribution.-Hildebrand (1943: 64) reported the range as "Southern Florida to Panama and the West Indies." I found this species abundant from Jupiter, Florida, southward along the Atlantic coast and through the 1970] Daly: Southern Florida Anchovies 75 Florida Keys. The closely related Anchoa hepsetus replaces A. lamprotaenia north of Jupiter in the Indian River and on the west coast of Florida in Florida Bay. Only two specimens of lamprotaenia were found among the hundreds of thousands of anchovies sorted from collections made in But- tonwood Canal, Flamingo, Everglades National Park; none was taken at Fort Pierce, just north of Jupiter. Specimens were examined from as far south as Puerto Rico and Venezuela. Anchoa hepsetus (Linnaeus) Figs. 2, 6 Esox hepsetus Linnaeus, 1758: 314 (type-locality: America). Anchoa hepsetus hepsetus, Hildebrand, 1943: 57-59, fig. 21; 1964: 194-200, fig. 40. Description.-COUNTS (meristic data are summarized in Tables 2-7): Dor- sal rays, 14-17; anal rays, 20-24; pectoral rays, 14-17; vertebrae, 41-44; gill rakers, 17-20 + 20-23. MORPHOMETRY(10 specimens 90 to 106 mm SL, from Flamingo, Flor- ida, herein termed "adults," UMML 20191; 10 specimens 51 to 61 mm SL, from Ft. Pierce, Florida, herein termed "juveniles," UMML 20188): Head moderate (22-28), body fairly deep (19-22). Snout moderately long (4.8-5.9); eye moderate in adult (6.5-7.3), large in juvenile (7.6-8.4). Lateral silver stripe not expanded posteriorly except at caudal base (width over anal 3.4-5.2). Anterior rays of depressed dorsal falling short of tips of posterior rays. Fins moderate (longest dorsal ray 15-17). Vent closer to anal fin than to depressed pelvic rays. Anal-fin origin under twelfth to fourteenth dorsal ray in juveniles, 1 or 2 mm posterior to dorsal in adults. Gill rakers fairly close, slender, tapering to half the basal width at their midpoint. PIGMENTATION:The ground color in life is translucent gray on the snout and dorsum, and silver on the opercular area, eye, jaw, and scales. Scales are easily shed in juveniles, but fairly adherent in adults. Beneath the scales, a narrow silvery stripe extends from the head to the caudal base; the peri- toneum is also silvery. The ground color becomes a light gray to yellow after fixation. Although living adults were not studied, juveniles observed in the field showed iridescence on the dorsum of the head and above the lateral stripe. Yellow and green predominate; noniridescent yellow is very noticeable around the head, lateral stripe, and cleithral area in living and freshly pre- served specimens. Melanophores are small. They outline the scale pockets dorsally, most noticeably posterior to the dorsal fin. Individuals at 55 mm SL are nearly devoid of speckles on the predorsal area, except for a thin middorsal stripe. 76 Bulletin of Marine Science [20(1)

FIGURE 2. Anchoa hepsetus (Linnaeus), UMML 20189, 95.0 mm SL.

Large dots underlying the lateral silver stripe usually show above the stripe as a thin, dark line in adults, but not in 60-mm specimens. Melanophores on the head continue forward in a band around the snout, reaching the an- terior ends of the premaxillae. Dark pigmentation shields the dorsum of each orbit and the brain. Two prominent, ovate spots formed of fine melanophores lie on the nape just posterior to the large, dark area on the skull. A narrow, light area lies between these spots at the midline; it is often wider anteriorly than posteriorly. In half-grown fishes (ca. 55 mm SL), the median separation is clearly evident and the skull pigmentation is also di- vided, giving the appearance of four closely appressed dark blotches lying along the midline (Fig. 6). Melanophores are also present along the fin rays of the dorsal and caudal fins and around the bases of the dorsal and anal fins.

Comparisons.-In the juvenile stage, this species is often confused with Anchoa lamprotaenia. A combination of meristic characters will readily distinguish the species; additional characters are given in Table 1. Distribution.-Hildebrand (1943: 58) reported the range of Anchoa hepsetus hepsetus as "Massachusetts, rarely to Nova Scotia, through the West Indies, and southward as far as Montevideo. It is interesting that only one speci- men of this is included in the 12 collections at hand from Key West and Tortugas, Fla., made at different times and by several collectors." I found this species common as far south on the east coast of Florida as 1970] Daly: Southern Florida Anchovies 77 TABLE 2 FREQUENCYDISTRIBUTIONSOF NUMBER OF DORSALRAYSIN ENGRAULID FISHES FROM SOUTHERNFLORIDAANDADJACENT REGIONS

Species and locality 13 14 15 16 17 x Anchoa lamprotaenia Jupiter, Fla. (UMML 20770) 1 11 3 15.1 Virginia Key, Miami, Fla. (UMML 20197) - 13 34 3 14.8 Ragged Keys, Fla. (UMML 20632) 2 1 15.3 Flamingo, Fla. (UMML 20192, 20193) 2 15.0 Indian Key, Fla. Keys (UMML 13041, 6631) - 4 11 14.7 Pigeon Key, Fla. Keys (UMML 20187) 4 7 4 15.1 Dry Tortugas (UMML 20676) 3 1 15.3 Los Roques, Venezuela (UMML 13034) 1 11 3 15.1 Anchoa hepsetus St. Augustine, Fla. (UMML 13461, 8113, 6940) 2 8 1 15.9 Ft. Pierce, Fla. (UMML 20188) - 13 2 16.1 Flamingo, Fla. (UMML 20191, 20190) 1 6 37 6 16.0 Bonita Springs, Fla. (UMML 8127, 4573, 4554, 3073) 3 9 1 15.8 Northern Gulf of Mexico (UMML 10403) - 5 6 15.5 Anchoa nasuta Ft. Pierce, Fla. (UMML 3326) - 15 15.0 Lake Worth, Fla. (UMML 20179) - 11 4 15.3 Virginia Key, Miami, Fla., 1965 (UMML 20198) - 14 1 15.1 Virginia Key, Miami, Fla., 1955 (UMML 691) 6 39 5 15.0 Anchoa mitchilli St. Augustine, Fla. (UMML 8454) 1 8 1 15.0 Stuart, Fla. (UMML 20205) 4 8 2 1 15.0 Lake Worth, Fla. (UMML 20178, 3148) 4 6 3 2 15.2 Virginia Key, Miami, Fla. (UMML 20199) 1 11 32 6 14.9 Key Largo, Fla. Keys (UMML 247) 1 3 11 14.7 Lower Matecumbe Key, Fla. Keys (UMML 8233, 3303) 1 2 14.7 Flamingo, Fla. (UMML 20196, 20195) 5 9 1 14.7 Everglades City, Fla. (UMML 2051, 1891) - 7 1 15.1 Bonita Springs, Fla. (UMML 11692, 5567, 4553) 2 13 14.9 Northern Gulf of Mexico (UMML 16639, 10402) 1 4 14.8 Anchoviella perjasciata Virginia Key, Miami, Fla. (UMML 20185, 20202) 7 36 7 15.0 Ragged Keys, Fla. (UMML 20203) 1 8 3 15.2 Seven Mile Bridge, Fla. Keys (UMML 998) - 1 3 1 ] 5.0 78 Bulletin of Marine Science [20( 1) TABLE 2-Continued

Speciesand locality 13 14 15 16 17 x SUMMARY Anchoa lamprotaenia - 23 81 15 14.9 Anchoa hepsetus I 16 73 10 15.9 Anchoa nasuta 6 79 10 15.0 Anchoa mitchilli 2 32 100 14 3 14.9 A nchoviella perfasciata 9 47 11 15.0

Fort Pierce, and along the entire west coast, including Flamingo. I saw no specimens of Anchoa hepsetus from the Florida Keys or the lower east coast of Florida. The species is common in the Gulf of Mexico (Gunter & Hall, 1965; Gunter, 1945, 1963; Springer & Woodburn, 1960); the systematic status of Anchoa hepsetus colonensis Hildebrand, reportedly common in the Gulf of Mexico, is unclear.

Anchoa nasuta Hildebrand & Carvalho Figs. 3, 6 (not Evermann & Marsh), (in part) Hildebrand, 1943: 65- 67; 1964: 200-202. Anchoa nasuta Hildebrand & Carvalho, 1948: 288-290, fig. 1 (type-specimen: USNM 143903, 54 mm SL, Sao Sebastiao Island, Sao Sebastiao, Brazil). Description.-CouNTS (meristic data are summarized in Tables 2-7): Dor- sal rays, 14-16; anal rays, 21-24; pectoral rays, 12-15; vertebrae, 40-42; gill rakers, 19-23 + 24-28. MORPHOMETRY(10 specimens 49 to 61 mm SL, collected at Virginia Key, Miami, Florida, UMML 20198): Head long (27-29), body slender (18-20). Snout long (6.7-7.9), eye small (6.7-7.6). Lateral silver stripe expanded over anal fin (width of stripe 5.4-6.6). Fins long (longest dorsal ray 15-19). Vent just anterior to anal-fin origin. Anal-fin origin slightly posterior to dorsal fin. Gill rakers crowded, slender, tapering only slightly (midpoint width more than half basal width). PIGMENTATION:In life the ground color is dark translucent gray on the snout and dorsal aspect; the opercular area, eye, jaw, and scales are silvery. Scales are often lost in fixation. Beneath the scales, a broad silvery stripe extends from the head to the caudal base; the peritoneum is also silvery. The ground color becomes a light gray to yellow after fixation. In living and preserved specimens, violet iridescence is prominent on the head and body, especially above the operculum. A juvenile specimen reared in an aquarium developed this characteristic color at about 30 mm SL. In living adults, the dorsum of the skull shows an iridescent golden spot; green 1970] Daly: Southern Florida Anchovies 79 TABLE 3 FREQUENCYDISTRIBUTIONSOF NUMBER OF PECTORALRAYSIN ENGRAULID FISHES FROM SOUTHERNFLORIDAANDADJACENTREGIONS

Species and locality 10 11 12 13 14 15 16 17 x Anchoa lamprotaenia Jupiter, Fla. (UMML 20770) 2 10 3 13.1 Virginia Key, Miami, Fla. (UMML 20197) - 10 24 16 13.1 Ragged Keys, Fla. (UMML 20632) 2 12.7 Flamingo, Fla. (UMML 20192, 20193) 1 1 13.0 Indian Key, Fla. Keys (UMML 13041, 6631) 4 6 4 13.1 Pigeon Key, Fla. Keys (UMML 20187) 4 8 3 12.9 Dry Tortugas (UMML 20676) 4 13.0 Los Roques, Venezuela (UMML 13034) 4 10 1 12.8 Anchoa hepsetus St. Augustine, Fla. (UMML 13461, 8113, 6940) 2 2 6 1 15.5 Ft. Pierce, Fla. (UMML 20188) 7 6 2 15.7 Flamingo, Fla. (UMML 20191, 20190) 2 29 19 15.0 Bonita Springs, Fla. (UMML 8217, 4573, 4554, 3073) 1 7 5 16.3 Northern Gulf of Mexico (UMML 10403) 5 5 15.6 Anchoa nasuta Ft. Pierce, Fla. (UMML 3326) 1 8 6 13.3 Lake Worth, Fla. (UMML 20179) 3 8 4 13.1 Virginia Key, Miami, Fla., 1965 (UMML 20198) - 10 4 1 13.4 Virginia Key, Miami, Fla., 1955 (UMML 691) - 15 28 7 12.8 Anchoa m itch illi St. Augustine, Fla. (UMML 8454) 3 6 1 11.8 Stuart, Fla. (UMML 20205) 7 5 2 11.5 Lake Worth, Fla. (UMML 20178, 3148) 2 11 11.8 80 Bulletin of Marine Science [20(1 ) TABLE 3-Continued

Species and locality 10 11 12 13 14 15 16 17 Virginia Key, Miami, Fla. (UMML 20199) - 14 28 8 11.9 Key Largo, Fla. Keys (UMML 247) 5 9 11.5 Lower Matecumbe Kev, Fla. Keys (UMML 8233; 3303) 2 1 11.3 Flamingo, Fla. (UMML 20196, 20195) 2 11 3 11.1 Everglades City, Fla. (UMML 2051, 1891) 1 5 3 11.2 Bonita Springs, Fla. (UMML 11692, 5567, 4553) 1 6 7 11.5 Northern Gulf of Mexico (UMML 16639, 10402) 3 2 11.4 Anchoviella perfasciata Virginia Key, Miami, Fla. (UMML 20185, 20202) 3 20 24 3 15.5 Ragged Keys, Fla. (UMML 20203) 2 10 14.8 Seven Mile Bridge, Fla. Keys (UMML 998) 1 3 15.0 SUMMARY Anchoa lamprotaenia - 26 64 28 13.0 Anchoa hepsetus 4 44 43 9 15.6 Anchoa nasuta - 19 54 21 1 13.0 Anchoa mitchilli 9 57 74 12 11.6 Anchoviella perfasciata 6 33 25 3 15.4 and golden iridescence is present on the skull and the upper border of the lateral silver stripe. The dorsum of the snout is noniridescent yellow. Melanophores are numerous on the dorsum. They are arranged in oblique rows, forming bands or a reticulated pattern. Lateral melanophores form a broad, well-defined stripe extending from the head to the caudal base in preserved specimens. In specimens retaining guanine, the lateral silver stripe covers the melanophores only partially; a prominent dark line shows above the silver stripe. Melanophore markings above the skull form a thin tract over the eyes; this band widens on top of the snout, continues forward around the tip of the snout, and ends far short of the premaxillae. No large spot or pair of spots appears on the nape; pigment here is diffuse. A small spot or pair of spots only 1 or 2 mm wide is usually present, how- ever, above the posterior part of the skull (Fig. 6). Melanophores are also distributed along the rays of the dorsal and caudal fins and around the bases of the dorsal and anal fins. 1970J Daly: Southern Florida Anchovies 81

FIGURE3. Anchoa nasuta Hildebrand & Carvalho, UMML 20198, 65.3 mm SL.

Comparisons.-This species closely resembles Anchoa lyolepis (Evermann & Marsh); it may be distinguished from lyolepis by its higher gill-raker count. Also, the dorsal pigmentation of lyolepis is usually much heavier than in nasuta. These two species may be separated from other species of Anchoa by their distinctively long head and snout, and slender body. Distribution.-Anchoa nasuta was originally described on the basis of six specimens from Sao Sebastiao and Vitoria, both of which are on the south- ern coast of Brazil near Rio de Janeiro. I have examined specimens from as far north as Cape Hatteras and as far south as Venezuela. Large popu- lations seem to occur sporadically in southeastern Florida, but I have not seen examples of nasuta from the Antillean Islands, and I have seen only one specimen (57 mm SL) from the gulf coast of Florida (Pinellas County). Remarks.-Hildebrand (1943, 1964) noted that his specimens of Anchoa lyolepis from the Gulf of Mexico have a consistently higher gill-raker count than those in other collections of that species, which were of Caribbean origin. It seems likely that these specimens from the Gulf of Mexico are referable to Anchoa nasuta, since the higher gill-raker count of nasuta is the principal means I have found for distinguishing it from lyolepis. This subpopulation in the Gulf of Mexico could then account for the unusually wide range of gill-raker counts which Hildebrand obtained for Anchoa lyolepis (Hildebrand, 1943: 36, 37; 1964: 166). 82 Bulletin of Marine Science [20(1 )

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FIGURE 4. Anchoa mitchilli (Valenciennes), UMML 20183,41.4 mm SL.

A nchoa mitchilli (Valenciennes) Figs. 4, 6 Engraulis mitchilli Valenciennes, in Cuvier & Valenciennes, 1848: 50 (type- locality: New York). Anchoa mitchilli, Hildebrand, 1943: 87-94, figs. 37 and 38 (two subspecies described); 1964: 176-181, figs. 31 and 32. Description.-CouNTS (meristic data are summarized in Tables 2-7): Dor- sal rays, 13-17; anal rays, 23-30; pectoral rays, 10-13; vertebrae, 38-41; gill rakers, 15-20 + 20-25. MORPHOMETRY (10 specimens 41 to 54 mm SL, collected at Virginia Key, Miami, Florida, UMML 20199,20178): Head short (24-28), body deep (19-24). Snout short (3.1-4.5), eye large (7.4-9.0). Lateral silver stripe narrow and often faded (width approximately 2.6-3.5). Fins mod- erate (longest dorsal ray 15-19). Vent close behind tips of depressed pel- vic rays. Anal-fin origin under fifth or sixth dorsal ray. Gill rakers slender, crowded, not rapidly tapering. PIGMENTATION: In life the ground color is a translucent gray on the snout and dorsal aspect; the opercular area, eye, jaw, and scales are silvery. The scales are easily shed; when present, they mask a narrow silver stripe extending from the head to the caudal base. Carefully preserved specimens retain guanine on the head for some time, but the lateral silver stripe usu- ally fades quickly. Ground color becomes a light gray to yellow after fixa- tion. The peritoneum is silvery. 86 Bulletin of Marine Science [20(1) TABLE 5 FREQUENCYDISTRIBUTIONSOF NUMBEROF GILL RAKERSONTHE UPPER LIMB OF THE FIRST ARCH IN ENGRAULIDFISHES FROMSOUTHERNFLORIDAAND ADJACENTREGIONS

Species and location 14 15 16 17 18 19 20 21 22 23 24 Anchoa lamprotaenia Jupiter, Fla. (UMML 20770) 2 10 3 15.1 Virginia Key, Miami, Fla. (UMML 20197) 1 22 19 8 15.7 Ragged Keys, Fla. (UMML 20632) - 3 15.0 Flamingo, Fla. (UMML 20192, 20193) - 2 - - - - - 15.0 Indian Key, Fla. Keys (UMML 13041, 6631) - 10 4 15.4 Pigeon Key, Fla. Keys (UMML 20187) 3 9 3 14.9 Dry Tortugas (UMML 20676) - 2 2 15.5 Los Rogues, Venezuela (UMML 13034) - 4 9 2 15.9 Anchoa hepsetus St. Augustine, Fla. (UMML 13461, 8113, 6940) 461 18.7 Ft. Pierce, Fla. (UMML 20188) - -11 4 - 18.3 Flamingo, Fla. (UMML 20191, 20190) - - - 6 24 16 3 - - - - 18.4 Bonita Springs, Fla. (UMML 8217, 4573, 4554, 3073) - - - 8 4 - - - 18.2 Northern Gulf of Mexico (UMML 10403) - - - 4 5 1 18.5 Anchoa nasuta Ft. Pierce, Fla. (UMML 3326) - - - 9 3 2 20.4 Lake Worth, Fla. (UMML 20179) 1 6 5 3 - 21.7 Virginia Key, Miami, Fla., 1965 (UMML 20198) - - - - 2 7 5 - 21.3 Virginia Key, Miami, Fla., 1955 (UMML 691) - - - - 7 21 21 - 21.3 Anchoa mitchilli St. Augustine, Fla. (UMML 8454) - - - 3 7 - - - - 17.7 Stuart, Fla. (UMML 20205) - 9 5 - - - 17.5 Lake Worth, Fla. (UMML 20178, 3148) - 1 3 7 3 - - - 17.7 Virginia Key, Miami, Fla. (UMML 20199) - 1 3 17 20 8 17.7 1970] Daly: Southern Florida Anchovies 87 TABLE 5-Continued

Speciesand location 14 15 16 17 18 19 20 21 22 23 24 x Key Largo, Fla. Keys (UMML 247) 7 5 3 ------17.7 Lower Matecumbe Key, Fla. Keys (UMML 8233, 3303) 2 - 1 - - - - 16.7 Flamingo, Fla. (UMML 20196, 20195) 6 7 3 ------16.8 Everglades City, Fla. (UMML 2051, 1891) 2 5 2 17.0 Bonita Springs, Fla. (UMML 11692, 5567, 4553) 7 7 1 ------16.6 Northern Gulf of Mexico (UMML 16639, 10402) - - - - 3 1 - - - - 18.6 Anchoviella perfasciata Virginia Key, Miami, Fla. (UMML 20185, 20202) 11 15 20 3 - 21.3 Ragged Keys, Fla. (UMML 20203) 3 5 4 20.1 Seven Mile Bridge, Fla. Keys (UMML 998) 2 - 2 19.6 SUMMARY Anchoa lamprotaenia 6 62 40 11 - - - - - 15.5 Anchoa hepsetus - - - 8 51 35 5 - - 18.4 Anchoa nasuta - - - - - 1 19 37 33 5 - 21.2 Anchoa mitchilli - 2 28 56 52 13 2 - - - - 17.3 Anchoviella perfasciata - - - - 1 6 16 21 20 3 - 20.8

In living specimens, an iridescent green-and-gold patch on the skull shields the brain dorsally; the same colors are on the dorsum of each orbit. No iridescence was observed above the silver stripe in this species. The snout is noniridescent yellow in some individuals and transparent in others. Melanophore pigmentation is higWy variable. Dorsal melanophores are arranged in a reticulated pattern which is oriented vertically. In preserved specimens, lateral melanophores often form a narrow, vague stripe extend- ing forward from the caudal base, but not reaching the head. This dark stripe is completely covered by the lateral silver stripe in newly fixed speci- mens. The portion of the head anterior to the eyes mayor may not be stippled; in fullest development, melanophores are continuous around the snout and onto the lower jaw. A double row of melanophores extends from the dorsal-fin origin to the caudal base, and a similar double row lies along the anal-fin base. Together, these rows outline the posterior half of the body when a specimen is viewed laterally. A prominent black spot is pres- 88 Bulletin of Marine Science [20(1 ) ent on top of the skull; melanophores are not concentrated on the nape. Melanophores are also distributed along the rays of the dorsal and caudal fins. Comparisons.-This species is most easily distinguished by the anterior placement of the anal fin. Other distinguishing characters include the very short snout; the absence of either a wide, silvery lateral stripe or a complete melanophore stripe; the short head and deep body; and the pronounced middorsal stripe between the dorsal-fin origin and the caudal base. Distribution.-Hildebrand (1943: 90,94) reported the range as Massachu- setts to Yucatan, Mexico, but not the West Indies. His collections did not include specimens from the Florida Keys; I found that A. mitchilli ranges to Long Key at least, and it is probably present throughout the Florida Keys. The species is common on both coasts of Florida.

Genus Anchoviella Fowler Anchoviella Fowler, 1911: 211 (type-species by original designation, Engrau- lis perfasdata Poey, 1860: 312).-Hildebrand, 1943: 108 (redefined; synonymy); 1964: 204. The genus is used here following the redescription of Hildebrand (1943: 108); the principal distinguishing character is the short, rounded maxilla, which does not extend past the angle of the jaw.

A nchoviella perfasciata (Poey) Figs. 5, 6 Engraulis perfasdatus Poey, 1860: 312 (type-locality: Cuba). Anchoviella perfasdata, Hildebrand, 1943: 116-118, fig. 49; 1964: 213-216, fig. 46. Description.-CouNTS (meristic data are summarized in Tables 2-7): Dor- sal rays, 14-16; anal rays, 16-19; pectoral rays, 14-17; vertebrae, 42-44; gill rakers, 18-23 -+- 26-29. MORPHOMETRY(10 specimens 50 to 77 mm SL, collected at Virginia Key, Miami, Florida, UMML 20185,20202): Head short (23-24), body slender (15-16). Snout short (3.7-4.5), eye (6.0-6.7) appears large in small head. Lateral silver stripe broad and posteriorly expanded (4.0-4.8). Vent just anterior to anal-fin origin. Anal-fin origin under or just behind two posteriormost dorsal rays. Gill rakers crowded, tapering to half the basal width at midpoint. PIGMENTATION:In life the ground color is translucent gray on the snout and dorsal aspect; the opercular area, eye, jaw, and scales are silvery. Scales, though present on the ventral two-thirds of the body, are usually lost 1970] Daly: Southern Florida Anchovies 89

FIGURE 5. Anchoviella perfasciata (Poey), UMML 20184, 49.1 mm SL. in fixation. Beneath the scales a broad, silvery stripe extends from the head to the caudal base; the peritoneum is also silvery. The ground color be- comes a light gray to yellow after fixation. The dorsum of the orbits is bright red-orange. A pair of characteristic bright blue spots lies just posterior to the eyes; between them lies part of a large green and gold patch which expands posteriorly to shield the brain dorsally. A thin green and gold stripe runs above the lateral silver stripe. Preserved specimens are pallid; melanophores are present in a middorsal line and may extend laterally to form a reticulated pattern. Melanophores show above the silvery lateral stripe as an extremely narrow line in small specimens, and as a 0.5-mm line in larger individuals (longer than 70 mm SL). Melanophores on the head may extend only part way onto the snout. Dark pigmentation shields the dorsum of the orbits and the brain. The pos- terior dorsum of the skull bears two large black spots. Just posterior to these spots, the melanophores on the nape form a second pair of dark spots, to give four closely grouped dark blotches along the midline (Fig. 6). Melanophores are also present along the rays of the dorsal and caudal fins and around the bases of the caudal and anal fins. A crescent-shaped dark area is apparent within the caudal base (Fig. 5). A middorsal line of large melanophores forms a distinctive short, dark stripe on the dorsum of the caudal peduncle. 90 Bulletin of Marine Science [20( 1)

TABLE 6 FREQUENCY DISTRIBUTIONS OF NUMBER OF GILL RAKERS ON THE LOWER LIMB OF THE FIRST ARCH IN ENGRAULID FISHES FROM SOUTHERN FLORIDA AND ADJACENT REGIONS

Species and location 18 19 20 21 22 23 24 25 26 27 28 29 x Anchoa lamprotaenia Jupiter, Fla. (UMML 20770) 411 ------18.7 Virginia Key, Miami, Fla. (UMML 20197) 4 29 16 ------19.3 Ragged Keys, Fla. (UMML 20632) 2 ------18.3 Flamingo, Fla. (UMML 20192, 20193) 2 ------18.0 Indian Key, Fla. Keys (UMML 13041, 6631) 3 6 5 1 ------19.3 Pigeon Key, Fla. Keys (UMML 20187) 6 8 1 ------18.6 Dry Tortugas (UMML 20676) 1 2 1 ------19.0 Los Roques, Venezuela (UMML 13034) - 7 7 ------19.6 Anchoa hepsetus St. Augustine, Fla. (UMML 13461, 8113, 6940) - - - 3 7 21.8 Ft. Pierce, Fla. (UMML 20188) - - - 4 8 3 ------21.9 Flamingo, Fla. (UMML 20191, 20190) 7 32 11 - - - - - 21.1 Bonita Springs, Fla. (UMML 8217, 4573, 4554, 3073) 2 6 5 - - - - - 21.2 Northern Gulf of Mexico (UMML 10403) 3 7 1 - - - - - 20.8 Anchoa nasuta Ft. Pierce, Fla. (UMML 3326) ------5 8 - 2 - 25.9 Lake Worth, Fla. (UMML 20179) - - - - - 3 8 4 26.1 Virginia Key, Miami, Fla., 1965 (UMML 20198) ------2 10 3 26.1 Virginia Key, Miami, Fla., 1955 (UMML 691) ------4 7 31 8 25.9 Anchoa mitchilli St. Augustine, Fla. (UMML 8454) - - - - - 4 5 1 - - - - 23.7 1970] Daly: Southern Florida Anchovies 91 TABLE ~Continued

Species and location 18 19 20 21 22 23 24 25 26 27 28 29 x Stuart, Fla. (UMML 20205) - - - - 2 7 5 - - - - 23.3 Lake Worth, Fla. (UMML 20178, 3148) 1 1 1 8 3 1 22.9 Virginia Key, Miami, Fla. (UMML 20199) 1 1 12 20 11 5 23.1 Key Largo, Fla. Keys (UMML 247) - - - 1 2 8 3 1 - - - - 23.1 Lower Matecumbe Key, Fla. Keys (UMML 8233, 3303) 1 - 1 ------21.7 Flamingo, Fla. (UMML 20196, 20195) - 2 10 3 - - - 22.1 Everglades City, Fla. (UMML 2051, 1891) 3 6 - - - - 22.7 Bonita Springs, Fla. (UMML 11692, 5567, 4553) - - - 1 10 2 2 - - - 22.3 Northern Gulf of Mexico (UMML 16639, 10402) - - - - 3 2 24.4

Anchoviella perfasciata Virginia Key, Miami, Fla. (UMML 20185, 20202) - - 10 23 13 4 27.2 Ragged Keys, Fla. (UMML 20203) 47- 1 26.8 Seven Mile Bridge, Fla. Keys (UMML 998) 3 2 - - 26.4

SUMMARY Anchoa lamprotaenia 22 64 30 3 ------19.1 Anchoa hepsetus 12 52 32 4 - - - - 21.3 Anchoa nasuta ------4 17 57 15 2 - 25.9 Anchoa mitchilli 3 6 41 59 33 11 - - - - 22.9 Anchoviella perfasciata ------17 32 13 5 27.1

Comparisons.- This species may be distinguished from other members of the genus found in Florida by its short head and the relatively high place- ment of the lateral stripe. Distribution.-Hildebrand (1943: 118) reported the range as "The West Indies, and northward to southern Florida and Mississippi. This species apparently is not as abundant as many other anchovies." I found this spe- cies to be seasonally abundant in the Florida Keys and at Miami, where it is much more easily collected by night-light than by shore seining. The 92 Bulletin of Marine Science [20(1) apparent rarity of the species may be due to its tendency to remain at a distance from shore during the day, thus avoiding seines.

ECOLOGY Huge concentrations of many species of clupeoid fishes are frequently seen along protected shorelines in southern Florida. Shore seining in Sep- tember in the Miami area may yield, for example, many individuals of Anchoa lamprotaenia and Anchoa mitchilli, a few specimens of Anchoa lyolepis and Anchoviella perfasciata, and a great many other clupeoids such as Sardinella anchovia Valenciennes, Harengula spp., and lenkinsia spp. These schools are truly composite; the varied iridescent colorations charac- teristic of the different species can be seen intermingling freely. Feeding jacks (Caranx spp.) are often seen driving the fishes close to shore, and at times the fleeing c1upeoids leap onto the shore, then wriggle back into the water. Frequent cast-net collecting during twilight hours showed that these schools close to shore always dispersed or moved away from shore just be- fore the onset of darkness. In December, adult anchovies often disappear from shore-seine collec- tions until the following spring. In his study of Anchoa hepsetus and An- choa mitchilli in the more northerly waters of Delaware Bay, Stevenson (1958) collected no anchovies by shore seine during the colder half of the year, despite a specific attempt at year-round sampling of the two species. In the Miami area, Anchoa lamprotaenia remained abundant in Biscayne Bay during the winter of 1965-66, but had not during the previous two winters. Juveniles of various species were collected throughout the year at Miami from 1963 to 1966. The seasonal fluctuations in abundance are complex and undoubtedly result from a network of environmental influ- ences. Collecting by dip net under a night light in passes or cuts yields a catch which differs from a shore-seine collection. For example, Anchoviella per- fasciata was relatively rare in shore collections, but the species appeared in great numbers under a night light. Individuals of all the species studied were often observed feeding on swarms of zooplankton under night lights. Schools were not formed on these occasions unless predators, such as the carangid Selene vomer, were active. The anchovies then ceased feeding and formed small schools close to the surface. From these observations I tentatively conclude that the species of an- chovies studied feed nocturnally in areas of current, and return close to shore to escape predation during the day. Anchovies kept in aquaria were fed zooplankton or the nauplii of Arte- mia. These fishes seemed to have an ability to locate individual nauplii; they did not strain the water as indiscriminately as juvenile specimens of 1970] Daly: Southern Florida Anchovies 93

FIGURE 6. Dorsal views of the five species: a, Anchoa mitchilli, adult, 48.6 mm SL, UMML 20205; b, Anchoa lamprotaenia, adult, 60.0 mm SL, UMML 20186; c, Anchoviella perfasciata, juvenile, 58.0 mm SL, UMML 20202; d, Anchoa nasuta, adult, 63.3 mm SL, UMML 20198; e, Anchoa hepsetus, juve- nile, 64.1 mm SL, UMML 20188; f, Anchoviella perfasciata, adult, 82.2 mm SL, UMML 20204; g, Anchoa hepsetus, adult, 114 mm SL, UMML 8113. 94 Bulletin of Marine Science [20(1) TABLE 7 FREQUENCYDISTRIBUTIONSOF NUMBER OF VERTEBRAEIN ENGRAULIDFISHES FROM SOUTHERNFLORIDAANDADJACENTREGIONS

Species and location 38 39 40 4] 42 43 44 x Anchoa lamprotaenia Jupiter, Fla. (UMML 20770) 4 9 2 40.9 Virginia Key, Miami, F]a. (UMML 20197) 5 36 9 41.1 Ragged Keys, Fla. (UMML 20632) - 2 ] 41.3 Flamingo, Fla. (UMML 20192, 20193) 2 41.0 Indian Key, Fla. Keys (UMML 13041, 6631) 5 7 3 40.9 Pigeon Key, F]a. Keys (UMML 20187) 1 3 8 3 40.9 Dry Tortugas (UMML 20676) 1 3 40.8 Los Roques, Venezuela (UMML 13034) 2 ]2 1 40.9 Anchoa hepsetus 51. Augustine, Fla. (UMML 13461, 8113, 6940) 1 9 42.7 Ft. Pierce, Fla. (UMML 20188) - 12 3 43.2 Flamingo, Fla. (UMML 20191, 20190) - 15 28 2 42.7 Bonita Springs, Fla. (UMML 8217, 4573, 4554, 3073) 3 9 1 42.8 Northern Gulf of Mexico (UMML 10403) 2 9 42.8 Anchoa nasuta Ft. Pierce, Fla. (UMML 3326) - 12 3 41.2 Lake Worth, Fla. (UMML 20179) - 14 41.1 Virginia Key, Miami, Fla., 1965 (UMML 20198) 1 11 3 41.1 Virginia Key, Miami, Fla., 1955 (UMML 691) - 38 11 40.4 Anchoa mitchilli St. Augustine, Fla. (UMML 8454) - 1 6 3 40.2 Stuart, Fla. (UMML 20205) 1 11 2 39.9 Lake Worth, Fla. (UMML 20178, 3148) 4 7 3 39.9 Virginia Key, Miami, Fla. (UMML 20199) 5 30 15 38.6 Key Largo, Fla. Keys (UMML 247) 1 7 7 40.4 Lower Matecumbe Key, Fla. Keys (UMML 8233, 3303) 2 40.3 1970] Daly: Southern Florida Anchovies 95 TABLE 7-Continued

Species and location 38 39 40 41 42 43 44 x Flamingo, Fla. (UMML 20196, 20195) 2 12 2 40.0 Everglades City, Fla. (UMML 2051, 1891) 3 5 39.8 Bonita Springs, Fla. (UMML 11692, 5567, 4553) 3 10 2 39.9 Northern Gulf of Mexico (UMML 16639, ]0402) 5 40.0 Anchoviella perfasciata Virginia Key, Miami, Fla. (UMML 20]85, 20202) 3 39 8 43.1 Ragged Keys, Fla. (UMML 20203) - 2 8 2 43.0 Seven Mile Bridge, Fla. Keys (UMML 998) 5 43.0 SUMMARY Anchoa lamprotaenia 20 79 19 41.0 Anchoa hepsetus 1 21 67 6 42.8 Anchoa nasuta 1 75 18 41.2 Anchoa mitchilli 20 95 36 40.1 Anchoviella perfasciata 5 52 10 43.1

Sardinella anchovia did under the same conditions. The complex anterior lateralis system, with its hundreds of pores and anastomosing canals (Tretiakov, 1950: fig. 7), may provide the sensing mechanism necessary to enable the anchovies to feed in this manner.

DISCUSSION The species described were separated most readily by using meristic characters. When the means of these characters were computed, the sam- ple means were consistent with one another, even when samples from widely separated areas were compared (see Tables 2-7). Of those studied in detail, the two species closest to one another in all respects are Anchoa lamprotaenia and Anchoa hepsetus. Despite the close resemblances, use of both upper gill-raker and pectoral-ray counts allowed separation of more than 99 per cent of the specimens examined. Proportional measurements become more important in forms with shorter maxillae (the genera Engraulis and Anchoviella). At least among southern Florida forms, these fishes are extremely close meristically and are separa- ble chiefly on the basis of slight differences in characters such as head length and pigmentation. Bodily proportions are more consistent than other non- 96 Bulletin of Marine Science [20 (1) meristic characters, but, as with meristics, a combination of characters may be needed to eliminate overlap. Characters such as melanophore pattern are usually difficult to use in distinguishing species of anchovies. Four factors contribute to this prob- lem: (1) intraspecific variation is high even among members of a single school; (2) interspecific differences are rarely obvious; (3) the distinguish- ing character is occasionally absent from some individuals; and (4) changes with growth are usually greater than interspecific differences. Differences in melanophore pattern enable an experienced person to sort species, but sorting can only be done with fairly large specimens (ca. 45 mm or more) in good condition. The size and extent of the lateral guanine stripe can be very useful in sorting, especially since effects of fixation on the guanine dif- fer among species. Geographical differences in nonmeristic characters were not detected within the limited scope of this study. Hence, fairly fine distinctions did not break down when collections from different areas were studied.

MATERIAL EXAMINED Specimens examined are listed by species and major locality. The catalog number is followed in parentheses by the number of specimens and, where measurements were taken, by the range of standard lengths. When only part of a collection was measured, the number of additional specimens not measured is included within the parentheses and preceded by a plus sign. Unprefixed catalog numbers below 100,000 are "UMML" numbers of the Rosenstiel School of Marine and Atmospheric Sciences, University of Miami; numbers exceeding 100,000 are catalog numbers of the Tropical Atlantic Biological Laboratory, Bureau of Commercial Fisheries, Miami. The prefix "FSU" refers to the Florida State University Collection, Tallahassee. Anchoa Zamprotaenia.-DADE COUNTY: 20632 (12) Ragged Keys, pass between two southernmost islands, RD-F-8, 24 Sept. 1964.-20509 (37) Bear Cut at Virginia Key, RD-F-41, 11 Aug. 1965.-20506 (34) Bear Cut at Key Biscayne, RD-F-37, 21 May 1965.-20505 (12) Bear Cut at Virginia Key, RD-F-35, 8 April 1965.-20502 (1, 45) Biscayne Bay at NE. 123 St., RD-F-22, 6 Dec. 1964.-20500 (8) Bear Cut at Virginia Key, CRR-F-354, C. R. Robins, 4 Oct. 1963.-20200 (11,31-39) Bear Cut at Virginia Key, RD-F-l, 13 Nov. 1963.-20197 (50, 54-67) Bear Cut at Virginia Key, RD-F-5, 19 Oct. 1964.-20186 (19, 53-61) Bear Cut at Key Biscayne, RD-F-44, 20 Aug. 1965.-11621 (640) Beach at Cape Florida, Key Biscayne, CRR-F-343, D. de Sylva et aZ., 18 Nov. 1961.- 8246 (18) Bear Cut at Virginia Key, CRR-F-239, C. R. Robins and B. Collette, 25 Oct. 1960.-8184 (1) Virginia Key, CRR-F-246, C. R. Robins and class, 8 Nov. 1960.-2270 (2) Key Biscayne, N. end of Crandon Park, 1970] Daly: Southern Florida Anchovies 97 CRR-F-48, C. R. Robins et al., 22 Jan. 1957.-1051 (41) Bear Cut at Virginia Key, CRR-F-90, C. R. Robins et at., 11 Aug. 1957. MONROECOUNTY,FLORIDAKEYS: 20504 (33) Indian Key land fill on Overseas Highway, bay side, RD-F-33, 12 Mar. 1965.-20187 (15,46-62) Pigeon Key, old pier at N. end of island, 3% miles W. of Marathon, RD-F-36, 21 April 1965.-13041 (8, 43-58) E. side of Indian Key in 2 to 6 feet, CRR-F-357, H. Feddern and W. Starck II, 11 May 1963.- 9112 (15) Indian Key, S. side, CRR-F-140, C. R. Robins et al., 3 May 1958.-7494 (52) Bay side of Knight Key at E. end of Seven Mile Bridge, CRR-F-98, C. R. Robins et al., 25 Aug. 1957.-6631 (7, 41-58) Indian Key, E. side, CRR-F-203, W. Starck II et al., 19 Sept. 1959.-3136 (2, 34-38) Barnes Sound, 1.5 miles N. of Jewfish Creek bridge, CRR-F-82, C. R. Robins et at., 22 July 1958.-2126 (5) Long Key, 3.5 miles S. of Greyhound Key, bay side, CRR-F-106, C. R. Robins and R. Manning, 12 Oct. 1957.-1183 (4) Grassy Key, 1 mile S. of Tom's Harbor viaduct no. 4, CRR-F-34, C. R. Robins et al., 27 Oct. 1956.-393 (1,48) Long Key Bight, viaduct at Greyhound Key just NE. of Long Key, CRR-F-33, C. R. Robins et at., 27 Oct. 1956. MONROECOUNTY,EVERGLADESNATIONAl..PARK: 20193 (1, 56) Fla- mingo, Buttonwood Canal at highway bridge, E. S. Iversen 934, 17 March 1965.-20192 (1, 50) Flamingo, Buttonwood Canal at highway bridge, E. S. Iversen 933, 17 March 1965. PALMBEACHCOUNTY: 20770 (15, 43-58 + 77 additional specimens) Jupiter, S. side of Inland Waterway on U.S. 1, RD-F-47, 19 Sept. 1965.- 20508 (2) Lake Worth, Riviera Beach bridge, RD-F-40, 27 May 1965.- 20507 (7) Intracoastal Waterway bridge on U.S. 1 at Jupiter, RD-F-39, 27 May 1965. DRYTORTUGAS:20676 (4, 49-65) Loggerhead Key, S. side, along shore, WAS-Tortugas-2, W. Starck II, 14 July 1961. VENEZUELA:13034 (15, 50-57 + 13 additional specimens) Los Roques, S. island of Dos Mosquises, DdeS 465, D. de Sylva et at., 5 Aug. 1963. PUERTORICO: 1945 (2) Beach pond at Rio Afiasco, CRR-PR-4, D. Erdman and Sr. Zalduondo, 18 Dec. 1957. Anchoa hepsetus.-MoNROE COUNTY, EVERGLADESNATIONALPARK: 20191 (30,83-106) Flamingo, Buttonwood Canal at highway bridge, E. S. Iversen 905, 16 Feb. 1965.-20190 (20, 51-66) Flamingo, Buttonwood Canal at highway bridge, E. S. Iversen 685, 26 June 1964.-20189 (5) Flamingo, Buttonwood Canal at highway bridge, E. S. Iversen 906, 16 Feb. 1965.-11772 (1) Flamingo, Buttonwood Canal at highway bridge, D. Tabb, 11 Nov. 1962.-11326 (1) Florida Bay, marker 2, FB-51, D. Tabb and R. Manning, 13 Nov. 1958.-8496 (1) Joe Kemp Channell and 2, CRR-F-291, 10 Oct. 1959.-8458 (1) Florida Bay, between E. Cape and 98 Bulletin of Marine Science [20(1) marker 3, drag No.1, FB-121, D. Tabb, 10 Nov. 1958.-8456 (1) Florida Bay marker T-6, FB-115, D. Tabb, 10 Nov. 1958.-6211 (1) Coot Bay end of Buttonwood Canal, FB-134, D. Tabb, 13 April 1959. COLLIERCOUNTY: 3073 (1, 65) 14 mile S. of Everglades (City) on road to Chokoloskee, CRR-F-284, S. and R. Cissel, 4 Aug. 1957.-4573 (8,58-65) Bonita Springs Beach, R. Cissel, 4 Aug. 1957.-4554 (1,98) Bonita Springs Beach, CRR-F-187, S. and R. Cissel, 2 March 1958. ST. LUCIE COUNTY: 20188 (15, 58-72 + 28 additional specimens) Fort Pierce, S. bridge over Indian R., RD-F-38, 27 May 1965. ST. JOHNSCOUNTY: 13461 (2, 95-117) St. Augustine, off Light in 56 ft, CRR-F-367, Andrew Jones, 1 Feb. 1961.-8113 (3, 95-114) off St. Augustine in 8 to 9 fms., CRR-F-270, Andrew Jones, 29 Nov. 1960.- 6940 (6,81-97) off St. Augustine in 8 to 9 fms., CRR-F-232, J. Higman. NORTHERNGULF OF MEXICO: 16638 (2) lat. 30012'N, long. 88°08'W, in 5 fms., OREGONsta. 4042, 28 Nov. 1962.-10403 (11, 92-101) lat. 29°37'N, long. 84°54.5'W, in 3 fms., OREGON sta. 2711, 21 Jan. 1960. GEORGIA: 7608 (1) lat. 32°59'N, long. 79°14'W, in 5-6 fms., SILVER BAYsta. 1365,21 Oct. 1959. Anchoa nasuta.-DADE COUNTY: 20198 (15, 54-64 + 17 additional speci- mens) Bear Cut at Virginia Key, CRR-F-391, C. R. Robins and class, 8 Oct. 1965.-9962 (1) Key Biscayne, shoreline along Cape Florida, CRR- F-297, C. R. Robins, 15 May 1961.-8842 (2) Virginia Key, NE. shore at ocean, DRP-F-235, D. Paulson et ai., 13 Jan. 1961.-691 (50, 55-61 + 11 additional specimens) Bear Cut at Virginia Key, J. McNulty and D. de Sylva, 5 Apr. 1955. PALM BEACHCOUNTY: 20180 (58) Inland Waterway and U.S. 1 at Jupiter, RD-F-47, 19 Sept. 1965.-20179 (15, 56-64 + 65 additional specimens) Lake Worth, Ph miles N. of Riviera Beach bridge on Route AlA, RD-F-46, 19 Sept. 1965. ST. LUCIE COUNTY: 3326 (15, 39-49 + 8 additional specimens) Fort Pierce, N. bridge over Indian R., D. Tabb, 19 July 1956. BREVARDCOUNTY: 100352 (18, 30-38) lat. 27°50'N, long. 80004.5'W, SILVERBAYsta. 5098, 26 Sept. 1963.-100355 (2, 50-52) lat. 28°17'N, long. 80028'W, SILVERBAYsta. 2620, 20 Jan. 1961. PINELLASCOUNTY: 100612 (1,57) lat. 28°04'N, long. 83°42'W, ORE- GONsta. 2157, 8 Apr. 1958. SOUTHCAROLINA:100358 (5, 39-45) lat. 32°26'N, long. 79°50'W, GILL cruise 9, regular sta. 44, 30 Nov. 1954. NORTHCAROLINA:100327 (7, 41-52) lat. 33°50.5'N, long. 78°06'W, SILVERBAY sta. 4521, 2 Nov. 1962.-100336 (2, 50-52) lat. 34°06'N, long. 77°46'W, SILVERBAYsta. 1213,2 Sept. 1959.-100370 (1,62) lat. 1970] Daly: Southern Florida Anchovies 99 35°13'N, long. 7Y32'W, GILL cruise 2, 11 May 1953.-100605 (1, 53) lat. 34°53'N, long. 76°09'W, GILL cruise 5, regular sta. 76,23 Feb. 1954. Anchoa mitchilli.-DADE COUNTY: 20503 (48) Snake Creek canal, salinity dam S-29 near Greynolds Park, RD-F-23, 12 Dec. 1964.-20199 (50, 38- 51) Bear Cut at Virginia Key, RD-F-7, 22 Sept. 1964.-9940 (1) Key Biscayne, shoreline along Cape Florida, CRR-F-297, C. R. Robins and class, 15 May 1961. MONROECOUNTY,FLORIDAKEYS: 8233 (2, 36-45) Canal at the home of W. A. Starck on Lower Matecumbe Key, CRR-F-283, W. Starck II and E. Belcher III, 4 June 1960.-3303 (1) Long Key, 3.3 miles SW. of Grey- hound Key, Florida Bay side, W. Courtenay et al., 28 June 1958.-247 (15,35-48 + 29 additional specimens) Key Largo, 1.8 miles from junction of U.S. 1 and N. Key Largo Rd., CRR-F-37, C. R. Robins et al., 17 Nov. 1956. MONROECOUNTY,EVERGLADESNATIONALPARK: 20196 (8, 52-65) Flamingo, Buttonwood Canal at highway bridge, E. S. Iversen 920,3 March 1965.-20195 (8, 36-57 + 17 additional specimens) Flamingo, Button- wood Canal at highway bridge, E. S. Iversen 765, 2 Sept. 1964.-20194 (5,50-56 + 8 additional specimens) Flamingo, Buttonwood Canal at high- way bridge, E. S. Iversen 921, 3 March 1965.-15824 (4) Buttonwood Canal, CRR-F-381, D. Tabb and R. Manning, 21 Nov. 1959.-8460 (25) Florida Bay, between E. Cape and marker No.3, FB-121, D. Tabb, 10 Nov. 1958.-6274 (20) Buttonwood Canal at Coot Bay end of Canal, D. Tabb, 13 Apr. 1959.-4867 (3, 37-40 + 10 additional specimens) Con- chie Channel off Cape Sable in grass beds, CRR-F-191, D. Tabb and D. Dubrow, 23-24 Apr. 1958. COLLIERCOUNTY: 6889 (777) MarcO Island, Gulf of Mexico N. of junction of Florida Route 92 and beach, DRP-SF-136, D. R. Paulson et al., 6 Sept. 1958.-3067 (26) ],4 mile S. of Everglades (City) on road to Chokoloskee, CRR-F-73, C. R. Robins and A. Moffett, 15 May 1957.- 2051 (6,33-37) freshwater canal 8.2 miles W. of Carnestown, CRR-F-87, A. Moffett and R. Cissel, 8 July 1957.-1891 (3,32-43) freshwater canal near town of Everglades, CRR-F-124, A. Moffett and R. Cissel, 16 July 1957. LEE COUNTY: 11692 (1, 49) Bonita Springs Beach, CRR-F-285, S. and R. Cissel, 13 Aug. 1957.-8220 (1) Bonita Springs Beach N. of public beach, CRR-F-284, S. and R. Cissel, 4 Aug. 1957.-5567 (6, 31-43) Bonita Springs Beach, CRR-F-208, S. and R. Cissel, 18 July 1958.-4553 (8,41-52 + 22 additional specimens) Bonita Springs Beach, CRR-F-187, S. and R. Cissel, 2 March 1958. MANATEECOUNTY: 3937 (12) inlet above Ellentown on Manatee R., CRR-F-56, J. Murdock and R. Volpe, 25 Feb. 1957. 100 Bulletin of Marine Science [20(1 )

NORTHERNGULF OF MEXICO: 16639 (1,62) lat. 30012'N, long. 88°08' Win 5 fms., OREGONsta. 4042, 28 Nov. 1962.-10402 (4,51-60) lat. 29° 37'N, long. 84°54.5'W in 3 fms., OREGONsta. 2711, 21 Jan. 1960. PALM BEACHCOUNTY: 20178 (3, 40-48 + 65 additional specimens) Lake Worth, 1% miles N. of Riviera Beach bridge on Route AlA, RD-F- 46, 19 Sept. 1965.-3148 (12, 23-45 + 1 additional specimen) Lake Worth at junction of Lake Worth Canal and Inland Waterway, CRR-F-86, W. Courtenay et al., 2 July 1957. MARTINCOUNTY: 20205 (15, 32-40) St. Lucie R. at Route 76 bridge, E. of Stuart, RD-F-48, 19 Sept. 1965. ST. JOHNSCOUNTY: 8454 (10,33-39 + 47 additional specimens) Ana- stasia State Park E. of St. Augustine, CRR-F-292, Andrew Jones, 1 Dec. 1960. Anchoviella perfasciata.-DADE COUNTY: 20501 (3) Bear Cut at Virginia Key, RD-F-ll, 19 Oct. 1964.-20203 (12,44-51) Biscayne Bay, pass be- tween two southernmost Ragged Keys, RD-F-8, 24 Sept. 1964.-20202 (20,52-77) Bear Cut at Virginia Key, RD-F-12, 20 Oct. 1964.-20185 (30,59-73) Bear Cut at Virginia Key, RD-F-29, 19 Feb. 1965.-8245 (8) Bear Cut at Virginia Key, CRR-F-239, C. R. Robins and B. Collette, 25 Oct. 1960. MONROECOUNTY,FLORIDAKEYS: 3890 (1,46) W. Summerland Key, ocean side, 1,4 mile beyond Bahia Honda Bridge, CRR-F-92, C. R. Robins et al., 17 Aug. 1957.-998 (5,37-44) Little Duck Key, 1,4 mile S. of Seven Mile Bridge, ocean side, CRR-F-91, C. R. Robins et ai., 17 Aug. 1957.

ADDITIONALSPECIES Five additional species mentioned only in the key were represented by only a few specimens in collections from southern Florida examined during this study. These species and the specimens collected are listed as follows: 1. Engraulis estauquae (Hildebrand): FSU 13144 (2, 51-56) Intra- coastal Waterway 4% miles N. of Jupiter Inlet, Florida, 12 Sept. 1960.- RD-F-47 (1, 91) Inland Waterway at U.S. 1, Jupiter, Florida, 19 Sept. 1965. 2. Engraulis eurystoie? (Swain & Meek): RD-F-29 (6, 55-64) Insti- tute of Marine Science pier, Miami, Florida, 19 Feb. 1965. 3. Anchoa cayorum (Fowler): UMML 20677 (2,52) Loggerhead Key, Dry Tortugas, S. side, along shore, WAS-Tortugas-2, W. Starck II, 14 July 1961. 4. Anchoa cubana (Poey): RD-F-47 (2, 53-57) Inland Waterway at U.S. 1, Jupiter, Florida, 19 Sept. 1965. 5. Anchoa Iyolepis (Evermann & Marsh): UMML 11793 (1,52) Log- 1970] Daly: Southern Florida Anchovies 101 gerhead Key, Dry Tortugas, S. side, along shore, WAS-Tortugas-2, W. Starck II, 14 July 1961. The first two species in the preceding list are placed in the genus En- graulis following Whitehead (1964). Identification of the specimens listed as Engraulis eurystole is not yet definite; if the name used here is correct, this would be a sizable extension in range, since Hildebrand (1943, 1964) found the species no farther south than Beaufort, North Carolina. The dis- tribution of Engraulis estauquae has also been extended, since this species had previously been recorded only from Venezuela (Hildebrand, 1943, 1964; Whitehead, 1964).

ACKNOWLEDGMENTS I thank Dr. C. Richard Robins of the University of Miami, Rosenstiel School of Marine and Atmospheric Sciences, for suggesting the problem of anchovy systematics and for the encouragement and advice he so generously gave me. I also appreciate the guidance and assistance given me by Dr. Donald P. de Sylva, who was the Chairman of my Committee of Graduate Study, and by the other members of my committee, Drs. C. Richard Robins, Leonard J. Greenfield, Cesare Emiliani, and Alton T. Butson. The study could not have been accomplished without all those who helped me with field collections. I am also grateful to Dr. Robins for the privilege of using the collections in the research museum at the Rosenstiel School of Marine and Atmospheric Sciences and to Dr. Thomas H. Fraser and Mr. David G. Smith for cataloging the specimens. Dr. Richard H. Chesher, Dr. William N. Eschmeyer, and Dr. Robert J. Livingston read and criticized the original manuscript. At the Bureau of Commercial Fisheries, Tropical Atlantic Biological Laboratory, Dr. William J. Richards and Mr. Frederick H. Berry helped me with further research, and Mrs. Ann Weeks aided in editing the final manu- script.

SUMMARY Meristic characters of southern Florida anchovies were enumerated and compared. Samples collected around the southern end of the Florida penin- sula were used to analyze variation in gill rakers, vertebrae, and dorsal-, anal-, and pectoral-fin rays. Most of the specimens were identified with the following five species: Anchoa lamprotaenia Hildebrand, Anchoa hepsetus (Linnaeus), Anchoa nasuta Hildebrand & Carvalho, Anchoa mitchilli (Valenciennes), and Anchoviella perfasciata (Poey). Although these five common species were easily separated on the basis of meristic characters alone, proportional characters and pigmentation were also found to be helpful in sorting and identification. 102 Bulletin of Marine Science [20(1 ) Anchoa mitchilli is distributed around the tip of Florida; Anchoa lampro- taenia and Anchoviella perfasciata were collected only on the lower east coast and in the Florida Keys; Anchoa nasuta was collected only on the lower east coast; and Anchoa hepsetus was distributed north of Jupiter on the east coast and also along the entire west coast of Florida. I conclude, on the basis of field observations and collections, that these fishes feed in the currents at night and form large schools composed of a mixture of species in protected areas during the day. Although abundantly collected at night-light stations, few specimens of Anchoviella perfasciata were taken with shore seines and cast nets during the day, which may indi- cate a tendency to remain in slightly deeper water than the other species.

SUMARIO SISTEMATICA DE LAS ANCHOAS DEL SUR DE LA FLORIDA (PISCES: ENGRAULIDAE) Se enumeran y comparan los caracteres meristicos de las anchoas del Sur de ]a F]orida. Se usaron muestras colectadas alrededor del extremo meri- diona] de ]a peninsula de ]a Florida para ana]izar variaciones en los rastrillos branquiales, las vertebras y los radios de las a]etas dorsal, anal y pectoral. La mayoria de los ejemplares fueron identificados con las cinco especies siguientes: Anchoa lamprotaenia Hildebrand, A. hepsetus (Linnaeus), A. nasuta Hildebrand & Carvalho, A. mitchilli (Valenciennes) y Anchoviella perfasciata (Poey). Aunque estas cinco especies comunes fueron facilmente separadas basandose s6lamente en caracteres meristicos, tambien resultaron Miles en la separaci6n e identificaci6n de las mismas las caracteristicas de sus pro- porciones y pigmentaci6n. Anchoa mitchilli esta distribuida alrededor del extremo sur de ]a F]orida, Anchoa lamprotaenia y Anchoviella perfasciata fueron colectadas s6lamente en la parte baja de la costa oriental y en los cayos de la Florida; Anchoa nasuta fue colectada unicamente en la parte baja de la costa orienta], y Anchoa hepsetus estaba distribuida al norte de Jupiter en la costa oriental y tambien a 10 largo de toda la costa occidental de la Florida. Yo conc1uyo, basandome en observaciones hechas en el area y en las co]ecciones, que estos peces se alimentan en la corriente durante la noche y durante el dia forman grandes cardumenes compuestos por diversas espe- cies en areas protegidas. Aunque abundantemente colectados en estaciones con iluminaci6n nocturna, pocos ejemplares de Anchoviella perfasciata fueron cogidos durante e] dia con redes de ribera y redes de lanzar, 10 que puede indicar una tendencia a permanecer en agua ligeramente mas pro- funda que las otras especies. 1970] Daly: Southern Florida Anchovies 103

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