184

Clupeiformes
Meristic characters in species belonging to the order Clupeiformes whose adults or larvae have been collected in the study area. Families and genera are listed alphabetically. Rarely observed counts in parentheses. Sources: Munroe (2000; 2002a; 2002b). Also see species accounts.
Family Species Clupeidae Alosa aestivalis Alosa mediocris Alosa pseudoharengus Alosa sapidissima Brevoortia tyrannus Clupea harengus Dorosoma cepedianum Dorosoma petenense Etrumeus teres Harengula jaguana Opisthonema oglinum Sardinella aurita Engraulidae Anchoa hepsetus Anchoa mitchilli Engraulis eurystole 4044 3844 4345 1316 1317 1316 1923 2330 1518 1315 1112 1416 7 7 7 4753 5355 4650 5557 4550 5262 4751 4344 4650 4143 4549 4549 1520 1520 1218 (14)1819(21) 1824 (16)1719(22) 1015 1115 1522 1719 1722 1619 1521 1923 1520 (18)2122(25) 1824 (15)1719(21) 2537 1727 1013 1718 2025 1620 1418 1516 1416 1318 1319 1321 1417 1217 1416 1315 1519 1516 911 9 10 810 7 610 710 (7)8 8 (7)8 89 78 Dorsal Fin Rays Anal Fin Rays Pectoral Fin Rays Pelvic Fin Rays

Vertebrae

Note: Three species of engraulids can best be distinguished by the relative positions of their dorsal and anal fins and by anal fin ray counts. These characters are most reliable in larvae >10 mm, juveniles and adults. Dorsal and anal fin origins can be almost opposite each other in adult A. mitchilli.

Early Stages of Fishes in the Western North Atlantic Ocean

185

Clupeiformes
Synopsis of characters for distinguishing larvae of the Clupeidae. "Dors-Anal" refers to number of myomeres between posterior dorsal and anterior anal fins. This number decreases at transformation in most species, as fin positions shift. Note that this number remains relatively stable through ontogeny in the genus Alosa.

Total Myomeres Alosa aestivalis Alosa mediocris Alosa pseudoharengus Alosa sapidissima Brevoortia tyrannus Clupea harengus Dorosoma cepedianum Dorosoma petenense 4753 5355 4650 5557 4550 5262 4751 4344

Preanal Myomeres 4245 3842 3943 4147 3840, then 3536 47, then 4146 3944 36

Predorsal Myomeres 3031, then 2425 33 to 25

Dors-Anal Myomeres 1113 78 79 9 to 71 5 to 1 89 to 4 10 to 71 10 to 71

Notochord Pigment Dorsal Tip No In small larvae Yes No In small larvae Varies No No

Notochord Pigment Ventral Tip Yes Yes Yes Yes Yes Yes No No

Miscellanous Slim-bodied, small-eyed Deep-bodied, large-eyed High D and A fin ray counts Late anal fin ray formation Very high anal fin ray count Scattered pigment on caudal peduncle Early teeth; low anal fin ray count Low myomere count High anal fin ray count

Etrumeus teres

4850

3940, then 36 35, then 27 4041, then 3436 3840, then 34

2732, then 26 25, then 10 2627, then 2223 2627, then 18

5 to 2

No

Yes

Harengula jaguana Opisthonema oglinum Sardinella aurita

1

3942 4549 4548

5 to 7 810 to 57 78 to 5

Yes No No

If yes, forms late Yes Yes

Determined from illustrations; need confirmation

Patterns of ventral pigment, posterior to cleithrum, in larvae of four species of Alosa. These patterns may be incomplete in many larvae. (Walsh et al., 2005, modified from Sismour, 1994.) Walsh et al., (2005) also provide a sizedependent key to larvae of these four species (plus Dorosoma cepedianum) from the southern part of the study area (Roanoke River, North Carolina).

A. pseudoharengus

A. aestivalis

A. mediocris

A. sapidissima

186

Alosa aestivalis (Mitchill, 1815) Clupeidae Blueback herring

Range: Habitat: Atlantic coast of North America from Nova Scotia to St. John's River, Florida Pelagic, schooling species, inhabiting coastal areas over the inner continental shelf (deeper during winter), moving into brackish or fresh waters to spawn Meristic Characters Myomeres: 4753 Vertebrae: 1416+3335 Dorsal fin rays: 1520 Anal fin rays: 1521 Pectoral fin rays: 1418 Pelvic fin rays: 911 Caudal fin rays: 78+10+9+67

Spawning: Adults migrate into coastal rivers during spring for spawning; begins in late Apr, continues through May in southern part of range, as late as Sep in Connecticut Eggs: Pelagic to semi-demersal, slightly adhesive Diameter: 0.871.11 mm Chorion: yellowish, semitransparent Chorion; somewhat thick, inner surface corrugated Oil globules small, unequal in size, scattered Yolk: granular Perivitelline space: wide

Yolk-sac larva

Larvae:

Hatching occurs at 3.15.0 mmTL; eye pigmented Body elongate with long, straight gut; anus always posterior to dorsal fin Sequence of fin ray formation: DCA, P1, P2 (ossification sequence based on illustrations) 1113 myomeres between posterior dorsal and anterior anal fins Discrete melanophores below pectoral fin base become a double line of pigment Pigment along upper and lower gut in distinct rows Notochord tip pigment present on ventral side only Transformation occurs between 20 and 30 mm

Note:

1. Slimmer and smaller-eyed than A. pseudoharengus 2. See comparative table in Clupeidae Introduction

Early Juvenile:

E. 45.0 mmTL

Figures: References:

Adult: H.L. Todd (Hildebrand, 1963a); Egg and yolk-sac larva: Mansueti and Hardy, 1967; AB: Chambers et al., 1976; C: Ann S. Green (Hildebrand, 1963a; D: Nancy D. Patton (Hildebrand, 1963a); E: Mansueti and Hardy, 1967 Jones et al., 1978; Able and Fahay, 1998; Munroe, 2000; 2002b

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187

Alosa aestivalis

A. 12.0 mmTL

B. 14.8 mmTL

C. 20.5 mmTL

D. 25.0 mmTL

188

Alosa mediocris (Mitchill, 1814) Clupeidae Hickory shad

Range: Habitat: Atlantic coast of North America from Bay of Fundy to Florida Inner continental shelf, estuaries, tidal rivers; oceanic distribution during winter not well known Meristic Characters Myomeres: 5355 Vertebrae: 17+37 Dorsal fin rays: 1520 Anal fin rays: 1923 Pectoral fin rays: 1516 Pelvic fin rays: 9 Caudal fin rays: 9+10+9+7

Spawning: In tidal freshwater, late Apr through early Jun Eggs: Spherical, semi-demersal, slightly adhesive Diameter: 0.961.65 mm Chorion: transparent, thick, finely corrugated Yolk: densely segmented, amber Oil globules: few, small Perivitelline space: very wide

Larvae:

Hatching occurs at 5.26.5 mm; eye pigmented Body elongate with long, straight gut; anus always posterior to dorsal fin Preanus length 8086% TL Yolk-sac larva Posterior gut with muscle-band striations Flexion occurs at 1015 mm Sequence of fin ray formation: DC, A, P2P1 (ossification sequence based on illustrations) Pigmentation includes elongate melanophores along ventral surface of gut with few spots on dorsal surface in early larvae; later larvae have melanophores on ventrum from throat area to region above anal fin Transformation occurs between 20 and 30 mm 1. See comparative table in Clupeidae Introduction

Note:

Early Juvenile:

E. 35.2 mmTL

Figures: References:

Adult: H.L. Todd (Hildebrand, 1963a); Egg, yolk-sac larva and AE: Mansueti, 1962 Jones et al. 1978; Able and Fahay, 1998; Munroe, 2000; 2002b

Early Stages of Fishes in the Western North Atlantic Ocean

189

Alosa mediocris

A. 15.8 mmTL

B. 19.5 mmTL

C. 24.3 mmTL

D. 27.0 mmTL

190

Alosa pseudoharengus (Wilson, 1811) Clupeidae Alewife

Range: Atlantic coast of North America from Newfoundland to South Carolina, most abundant between Gulf of Maine and Chesapeake Bay; also landlocked populations, Great Lakes and New York Pelagic, schooling species, inhabiting coastal areas adjacent to freshwater nursery areas; spends fall and winter in continental shelf waters 56110 m deep, primarily off Southern New England, Georges Bank and Gulf of Maine Adults migrate into coastal rivers during spring for spawning; larvae and juveniles occur in estuaries throughout range, except those with limited freshwater input. Demersal and adhesive Diameter: 0.871.11 mm Chorion: semitransparent and yellowish Yolk: segmented Oil globules: small, unequal in size, scattered Perivitelline space: wide

Habitat:

Spawning:

Meristic Characters Myomeres: 4650 Vertebrae: 4650 Dorsal fin rays: 1218 Anal fin rays: 1520 Pectoral fin rays: 1416 Pelvic fin rays: 910 Caudal fin rays: 10+9 (PrC)

Eggs:

Larvae:

Yolk-sac larva Hatching occurs at 3.15.0 mm; eyes pigmented Body elongate with long, straight gut; anus always posterior to dorsal fin Preanus length about 80% TL Posterior gut with muscle-band striations at sizes >10.0 mm Flexion occurs at about 9.0 mmSL Sequence of fin ray formation: D, C A P2 P1 (ossification sequence based on illustrations) Pigmentation includes row of spots on dorsal half of anterior gut, ventral surfaces of posterior gut; early larvae have rows of spots on dorsal and ventral surfaces of caudal peduncle; 1 large melanophore above base of pectoral fin; dorsolateral pigment appears in larger larvae Transformation occurs between 25 and 30 mm 1. Deeper-bodied and larger-eyed than Alosa aestivalis 2. See comparative table in Clupeidae Introduction

Note:

Early Juvenile: Larger juveniles have single, dark spot on side posterior to opercle Sharp scutes form along midline of belly

E. 42.0 mmTL
Figures: References: Adult: H.L. Todd (Hildebrand, 1963a); Egg, yolk-sac larva: Mansueti and Hardy, 1967; AB: Chambers et al., 1976; C: Norden, 1967; D: Ann S. Green (Hildebrand, 1963); E: Mansueti and Hardy, 1967 Jones et al., 1978; Able and Fahay, 1998; Munroe, 2000; 2002b

Early Stages of Fishes in the Western North Atlantic Ocean

191

Alosa pseudoharengus

A. 9.0 mmTL

B. 15.0 mmTL

C. 24.5 mmTL

D. 29.0 mmTL

192

Alosa sapidissima (Wilson, 1811) Clupeidae American shad

Range: Habitat: Atlantic coast of North America from Labrador to Florida; most abundant between Connecticut and North Carolina Congregate in Gulf of Maine during summer and fall, then most overwinter off Middle Atlantic Bight, at depths to 230 m Meristic Characters Myomeres: 5557 Vertebrae: 5557 Dorsal fin rays: 1421 Anal fin rays: 1825 Pectoral fin rays: 1318 Pelvic fin rays: 810 Caudal fin rays: 10+9 (PrC)

Spawning: Adults migrate into coastal rivers to spawn during spring; larvae develop in freshwater, juveniles form schools as they drift downstream summer and fall Eggs: 1. Demersal, adhesive early stages Later stages non-adhesive Diameter: 2.53.8 mm Chorion: Transparent, pale pink or amber Yolk: segmented Oil globules: none Perivitelline space: wide

Larvae:

Yolk-sac Larva Hatching occurs at 5.710.0 mm Body elongate with long, straight gut; anus always posterior to dorsal fin Air bladder obvious by 14 mm TL Preanal finfold retained throughout larval development Sequence of fin ray formation: D, C A P2 P1 (ossification sequence based on illustrations) Transformation occurs between 25 and 30 mm

Note:

See comparative table in Clupeidae Introduction

2. Ventral pigment in small A. sapidissima (center) differs from that in A. aestivalis (left) and A. pseudoharengus (right) in sizes <13.0 mmTL (after Sismour, 1994) Early Juvenile:

E. 51.8 mmTL

Figures: References:

Adult: H.L. Todd (Hildebrand, 1963a); Egg, yolk-sac larva and A: Ryder, 1887; BE: Mansueti and Hardy, 1967 Jones et al., 1978; Able and Fahay, 1998; Munroe, 2000; 2002b

Early Stages of Fishes in the Western North Atlantic Ocean

193

Alosa sapidissima

A. 14.0 mmTL

B. 22.9 mmTL

C. 21.0 mmTL

D. 31.8 mmTL

194

Brevoortia tyrannus (Latrobe, 1802) Clupeidae Atlantic menhaden

Range: Habitat: Atlantic coast of North America from the Gulf of St. Lawrence to northern Florida Pelagic, brackish estuaries and bays as well as inner continental shelf; rarely far from land

Spawning: May spawn during every month of the year, depending on location, with peaks during spring and fall; in the study area, most larvae occur in the Middle Atlantic Bight during fall into winter, with more limited occurrences in coastal waters during spring Eggs: Pelagic, spherical Diameter: 1.301.95 mm Chorion: smooth and thin Yolk: segmented Oil globules: single, 0.110.17 mm in diameter Perivitelline space: wide Late embryo has 2 rows of dorsolateral spots from snout to tail

Meristic Characters Myomeres: 4550 Vertebrae: 4550 Dorsal fin rays: 1824 Anal fin rays: 1824 Pectoral fin rays: 1319 Pelvic fin rays: 7 Caudal fin rays: 79+10+9+67

Yolk-sac larva

Larvae:

Hatching occurs at 2.44.5 mm; eyes unpigmented Body elongate, with straight gut 7080% TL; anus always posterior to dorsal fin Flexion occurs at 810 mm Posterior gut has obvious muscle-band striations Air bladder evident at about 11 mm No teeth until 20 mm Sequence of fin ray formation: C, D, A P2 P1 Pigmentation includes spots along entire dorsal surface of gut and along ventral surface of posterior half of gut in larvae larger than about 5 mm; few melanophores along dorsum of body disappear at 56 mm; pigment present on ventral side of notochord tip; dorsal notochord tip pigment only present in small larvae When spawned offshore, larvae enter estuaries after about 1 month at sizes of 10+ mm Transformation occurs at about 30 mm (in estuaries) 1. High number of dorsal and anal rays compared to other clupeids 2. Myomeres between posterior dorsal and anterior anal fins decrease from 5 to 1 3. Brevoortia smithi occurs only as far north as Cape Hatteras; 4547 myomeres; transforms at 2023 mm

Note:

Early juvenile:

F. 32.0 mm

Figures: References:

Adult: Whitehead, 1977; Egg and yolk-sac larva: Kuntz and Radcliffe, 1917 (redrawn); AD: Mansueti and Hardy, 1967 (redrawn); EF: Lewis et al., 1972 Houde and Swanson, 1975; Ditty et al. 1994; Munroe, 2000; 2002b

Early Stages of Fishes in the Western North Atlantic Ocean

195

Brevoortia tyrannus

A. 8.3 mmTL

B. 10.7 mmTL

C. 16.6 mmTL

D. 23.1 mmTL

E. 27.0 mmTL

196

Clupea harengus Linnaeus, 1758 Clupeidae Atlantic herring

Range: Both sides of North Atlantic Ocean; in the western North Atlantic from Greenland and Labrador to Cape Hatteras; separate stocks occur in Gulf of St. Lawrence, Banquereau Bank, Scotian Shelf, Gulf of Maine and Georges Bank Pelagic, schooling; mostly offshore and migrating into deeper waters during winter; some populations move into coastal waters for spawning Meristic Characters Myomeres: 5262 Vertebrae: 5262 Dorsal fin rays: 1622 Anal fin rays: 1521 Pectoral fin rays: 1321 Pelvic fin rays: 610 Caudal fin rays: 1013+10+9+89

Habitat:

Spawning: Mostly in fall, with a peak in Sep; also a spring spawn, peaking in May; occurs in bays and offshore banks, usually over substrates of rock, cobble or gravel. Georges Bank and periphery of Gulf of Maine are important centers; larvae seldom occur south of Hudson Canyon or off Nw Jersey Eggs: Demersal, adhesive, clumping, off-round Diameter: 1.01.4 mm Chorion: smooth, transparent and thick Yolk: segmented Oil globules: none Perivitelline space: wide

Yolk-sac larva

Larvae:

Hatching occurs at 410 mmTL; eyes pigmented Body elongate with long straight gut; anus always posterior to dorsal fin Preanus length 80%TL Flexion occurs at 1617 mm Posterior gut has obvious muscle-band striations Air bladder forms at 510 mm, but not obvious until 30 mm Sequence of fin ray formation: C D, A P2 P1 Pigmentation includes melanophores along the dorsal surface of the gut anteriorly, ventral surface of the gut posteriorly; larger larvae have only dorsal gut pigment; early larvae have single streak of pigment on midline of isthmus; large melanophores occur near anus, and at base of caudal fin; pigment present on ventral surface of notochord tip, variable on dorsal surface Transformation occurs at about 30 mm 1. High myomere count and late forming anal fin rays are unique for North Atlantic clupeids 2. Myomeres between posterior dorsal and anterior anal fins decrease from 89 to 4 3. See Mallotus villosus for note on similar larva

Note:

Early Juvenile:

F. 46.0 mmSL
Figures: References: Adult: Hildebrand, 1963a; Eggs: Berg et al., 1949; Yolk-sac larva, AB, D: Krevanovsky, 1956 (redrawn); C: Ehrenbaum, 1909 (redrawn); E: Fage, 1920 (redrawn); F: Susan Kaiser (Able and Fahay, 1998) Russell, 1976; Jones et al., 1978; Able and Fahay, 1998; Munroe, 2000; 2002b

Early Stages of Fishes in the Western North Atlantic Ocean

197

Clupea harengus

A. 8.2 mmTL

B. 20.2 mmTL

C. 29.0 mmTL
Pigment increases on head and dorso-lateral region

D. 29.0 mmTL (Head Region)

E. 41.0 mmTL

198

Dorosoma cepedianum (Lesueur, 1818) Clupeidae Gizzard shad

Range: North American fresh water drainages into the Atlantic Ocean and Gulf of Mexico; reaches estuarine waters of Chesapeake and Delaware bays and as far north as Hudson River estuary Near surface in open, quiet waters, deeper in fall and winter; found in submerged vegetation and over mud, sand and gravel substrates In fresh water in sloughs, ponds, lakes and larger rivers; spring through summer, with peak AprJun Demersal, adhesive Diameter: 0.75 mm Chorion: smooth, tough Yolk: finely granular Oil globule: 1 large and 15 smaller ones Perivitelline space: narrow Meristic Characters Myomeres: 4751 Vertebrae: 4751 Dorsal fin rays: 1015 Anal fin rays: 2537 Pectoral fin rays: 1417 Pelvic fin rays: 710 Caudal fin rays: 911+10+9+7

Habitat: Spawning: Eggs:

Larvae:

Hatching occurs at about 3.2 mm Body elongate with long straight gut; anus always posterior to dorsal fin Remnant of preanal finfold still present at 17 mm Yolk-sac larva Flexion occurs between 11 and 17 mm Sequence of fin ray formation: D C A, P2 P1 Pigmentation includes large melanophores near anus, and a series of spots on dorsal surface of anterior gut; larger larvae develop scattered pigment on caudal fin along base of anal fin, and dorsolaterally on body Transformation occurs at sizes larger than 24 mm; body begins to deepen at about 35 mm Scales complete at about 50 mm Large melanophore forms behind opercle in juveniles Ventral scutes form at about 35 mm Note high anal fin ray count

Early Juvenile:

F. 50.0 mmTL

Figures: References:

Adult: H.L. Todd (Goode, 1884); Egg, yolk-sac larva, and BC: Warner, 1940; A, D: Lippson and Moran, 1974; E: Fowler, 1935; F: Fowler, 1945 Lippson and Moran, 1974; Jones et al., 1978

Early Stages of Fishes in the Western North Atlantic Ocean

199

Dorosoma cepedianum

A. 11.0 mmTL

B. 17.5 mmTL

C. 22.0 mmTL

D. 24.2 mmTL

E. 35 mmTL

200

Dorosoma petenense (Gnther, 1867) Clupeidae Threadfin shad

Range: Ohio River drainage of North America to fresh waters of northern Guatemala and Belize; introduced into Chesapeake Bay drainages and occurs in Chesapeake and Delaware Canal Pelagic, schooling, in large bays, lakes, reservoirs, estuaries; in salinities 032 ppt, but mostly below 5 ppt

Habitat:

Spawning: Eggs and larvae may be restricted to fresh water, although juveniles and adults are occasionally collected in estuarine conditions Eggs: Demersal, adhesive Diameter: 0.75 mm No other data

Meristic Characters Myomeres: 4344 Vertebrae: 4344 Dorsal fin rays: 1115 Anal fin rays: 1727 Pectoral fin rays: 1217 Pelvic fin rays: (7) 8 Caudal fin rays: 10+9 (PrC)

Larvae:

Hatching occurs at 4.14.4 mm Body elongate with long, straight gut; anus always posterior to dorsal fin Flexion occurs at about 10 mmTL Yolk-sac larva About 36 preanal myomeres Sequence of fin ray formation: C, D A P2 P1 (ossification sequence based on illustrations) Adult fin ray complement present by 1820 mm Pigmentation includes a series of melanophores on dorsal surface of anterior gut, and scattered pigment over the anal fin base and caudal peduncle Transformation begins at about 20 mm

Figures: References:

Adult: Miller, 1963; Yolk-sac larva and AE: Taber, 1969 Lippson and Moran, 1974; Jones et al., 1978

Early Stages of Fishes in the Western North Atlantic Ocean

201

Dorosoma petenense

A. 8.2 mmTL

B. 11.0 mmTL

C. 15.6 mmTL

D. 17.4 mmTL

E. 20.3 mmTL

202

Etrumeus teres (DeKay, 1842) Clupeidae Round herring

Range: Habitat: Western North Atlantic Ocean from Bay of Fundy to Florida and Gulf of Mexico; also eastern Pacific and Indo-Pacific oceans Pelagic, schooling in continental shelf waters; occasionally in bays and coastal waters, but mostly over outer continental shelf depths Meristic Characters Myomeres: 4650 Vertebrae: 1517+3234 Dorsal fin rays: 1522 Anal fin rays: 1013 Pectoral fin rays: 1416 Pelvic fin rays: 8 Caudal fin rays: 1013+10+9+89

Spawning: Not well-described; possibly restricted to south of Cape Hatteras and Gulf of Mexico; larvae occasionally collected in study area Eggs: Pelagic, spherical Diameter: 1.171.37 mm Chorion: smooth, transparent and thick Yolk: segmented Oil globules: none Perivitelline space: narrow

Larvae:

Hatching occurs at 3.84.8 mm; eyes unpigmented Yolk-sac Larva Body elongate with long, straight gut; anus always posterior to dorsal fin Teeth apparent at about 6 mmTL Snout long and pointed in small larvae Flexion occurs at 810 mmSL Posterior gut has obvious muscle-band striations Sequence of fin ray formation: C, D A P2 P1 Fin rays not complete until transformation; pelvic forms late and migrates posteriorly Pigmentation includes about 8 widely spaced, large spots over anterior gut; small spots in double row form over posterior gut; a large melanophore over anus; spot at tip of lower jaw in some larvae Diagonal streaks of pigment over lower caudal fin base are distinctive Transformation occurs at 3033 mmTL 1. Early teeth formation and low anal fin ray count are unique among clupeid larvae 2. Myomeres between posterior dorsal and anterior anal fins decrease from 5 to 2

Note:

Early Juvenile:

F. 42.5 mmTL

Figures:

References:

Adult: Whitehead, 1977; Egg: O'Toole and King, 1974; Yolk-sac larva: Mito, 1961a; A: Mito, 1961; B: Houde and Fore, 1973; C: Uchida et al. 1958; D: Nancy Arthur (Moser et al. 1996); EF: Hildebrand, 1963a (Yolk-sac larva and AC redrawn) E.D. Houde, 1981 (pers. comm.); Shaw and Drullinger, 1990; Munroe, 2000; 2002b

Early Stages of Fishes in the Western North Atlantic Ocean

203

Etrumeus teres

A. 5.7 mmTL

B. 8.5 mmTL

C. 15.3 mmTL

Eye oval in early stages, becomes round at transformation

D. 17.7 mmTL

E. 27.5 mmTL

204

Harengula jaguana Poey, 1865 Clupeidae Scaled sardine

Range: Habitat: Western North Atlantic Ocean from New Jersey to southern Brazil, including Gulf of Mexico and Caribbean Sea Pelagic and demersal in coastal waters, over mud or sand substrates; often in bays, estuaries or high salinity lagoons

Spawning: JanSep (peak AprAug); most activity over inner continental shelf at night Eggs: Pelagic, spherical Diameter: 1.551.85 mm Chorion: transparent Yolk: segmented Oil globule: single, 0.070.10 mm in diameter Perivitelline space: wide

Meristic Characters Myomeres: 3942 Vertebrae: 4143 Dorsal fin rays: 1718 (19) Anal fin rays: 1718 Pectoral fin rays: (13) 1415 Pelvic fin rays: (7) 8 Caudal fin rays: 89+10+9+7

Larvae:

Hatch at about 2.5 mmTL, eyes unpigmented, mouth parts unformed Body elongate, with straight gut 8590% TL (decreases to 75% at transformation) Anus always posterior to dorsal fin Preanus myomeres 35 at 6.0 mm, 27 at 22 mm Predorsal myomeres 25 at 6.0 mm, 10 at 22 mm Flexion occurs at 911 mm Posterior gut has obvious muscle-band striations Air bladder evident at 6.0 mm Sequence of fin ray formation: C, D A P2 P1 Dorsal fin rays complete at 1416 mm; anal fin rays complete at 1315 mm Pigment sparse on head; early larvae have spots on dorsal (and ventral) tip of notochord, spots at cleithral symphysis, 2 spots near anus, ventral row of spots along posterior gut plus row of spots along dorsum of anterior gut; later larvae form row of spots along dorsum of posterior gut, few spots on bases of caudal fin rays; largest larvae add short rows of spots under dorsal fin and from posterior anal fin on venter of caudal peduncle; few spots also on hindbrain and on "cheek" area Transformation occurs at about 1422 mm 1. Low total myomere count; myomeres between dorsal and anal fins 57

Note:

Early Juvenile:

F. 21.3 mmSL

Figures: References:

Adult: Munroe and Nizinski, 2002; Egg: Gorbunova and Zvyagina, 1975; A, C, EF: Houde et al., 1974 (redrawn); B, D: Ditty et al., 2006 Houde et al., 1974; Gorbunova and Zvyagina, 1975; Fahay, 1983; Munroe and Nizinski, 2002

Early Stages of Fishes in the Western North Atlantic Ocean

205

Harengula jaguana

A. 6.0 mmSL

B. 8.6 mmSL

C. 11.5 mmSL

D. 14.4 mmSL

E. 16.1 mmSL

206

Opisthonema oglinum (Lesueur, 1817) Clupeidae Atlantic thread herring

Range: Western North Atlantic Ocean from southern New England and Bermuda, through the Gulf of Mexico, Caribbean Sea and West Indies; mostly tropical and sub-tropical waters; also eastern Pacific Ocean Pelagic, schooling in coastal waters, usually in high salinity, more rarely in bays or estuaries

Habitat:

Spawning: MayJun off North Carolina; larvae rarely collected in study area Eggs: Pelagic, spherical Diameter: 1.081.31 mm Chorion: smooth, thin and clear Yolk: lightly segmented Oil globule: single; 0.120.16 mm in diameter Perivitelline space: wide Paired dorsolateral series of melanophores on late embryo

Meristic Characters Myomeres: 4549 Vertebrae: 1213+3236 Dorsal fin rays: 1722 Anal fin rays: 2025 Pectoral fin rays: 1519 Pelvic fin rays: 89 Caudal fin rays: 9+10+9+67

Larvae:

Hatching occurs at >3.0 mm; eyes unpigmented Body elongate with long, straight gut; anus always posterior to dorsal fin Preanus length decreases from 8691% TL before flexion to 75% SL after transformation Predorsal myomeres decrease from 25 to 15 Flexion occurs at about 10 mmSL Posterior gut has obvious muscle-band striations Air bladder present, but not obvious until transformation Sequence of fin ray formation: C D A P2 P1 Pigmentation: in early larvae, melanophores on ventral midline under pectoral fins, posterior to anus, a double row along ventral surface of hindgut, and a dorsolateral row on each side of foregut Internal pigment over posterior gut well-defined; internal spots over air bladder prominent Transformation occurs at 1525 mmSL 1. Myomeres between posterior dorsal and anterior anal fins decrease from 810 (<16 mm) to 57 at transformation 2. Notochord tip pigment restricted to ventral side 3. High anal fin ray count 4. Pigment increases on dorsum of head at about 24 mmSL

Note:

Early Juvenile:

H. 30.8 mmSL

Figures: References:

Adult: A.S. Green (Munroe, 2002b); Egg and AH: Richards et al., 1974 Richards et al., 1974; Munroe, 2000; 2002b; Munroe and Nizinski, 2002

Early Stages of Fishes in the Western North Atlantic Ocean

207

Opisthonema oglinum

A. 4.0 mmTL

B. 10.7 mmSL

C. 13.7 mmSL

D. 17.1 mmSL

E. 19.7 mmSL

F. 24.4 mmSL

G . 24 .4 mmS L (Dorsal Head)

208

Sardinella aurita Valenciennes, 1847 Clupeidae Spanish sardine

Range: Eastern and western North Atlantic Ocean and Mediterranean Sea; in the western North Atlantic from Cape Cod and Bermuda to Brazil, including Gulf of Mexico and Caribbean Sea Pelagic in inner continental shelf waters Year-round with distinct peaks in different areas; fallwinter off Florida; NovJun (peak JanFeb) off Venezuela; mid-JunSep off North America; SepFeb in Gulf of Mexico; larvae uncommonly collected in study area during spring Pelagic and spherical Diameter: 1.031.12 (1.25) mm Chorion: thin, unsculptured, unpigmented Yolk: vaguely segmented Oil globule: single, 0.130.18 mm in diameter Perivitelline space: moderately wide Meristic Characters Myomeres: 4348 Vertebrae: 4549 Dorsal fin rays: 1619 Anal fin rays: 1620 Pectoral fin rays: 1516 Pelvic fin rays: 78 Caudal fin rays: 8+10+9+7

Habitat: Spawning:

Eggs:

Larvae:

Hatching occurs at 2.5 mm, eyes and body unpigmented, yolk-sac elliptical with oil globule posteriorly Preanal length ranges from 8389% SL, then decreases to 73% SL at transformation; head length ranges from1427% SL, then increases to 29% SL at transformation; body depth ranges from 812% SL, then increases to 25% SL at transformation Dorsal fin migrates forward: predorsal length 6068% SL until transformation when it decreases to 41% SL Myomeres between posterior dorsal fin and anterior anal fin decrease from 78 to 5 Flexion occurs at 7.5 to 9.5 mm Sequence of fin ray formation: C D, A P2 P1 Pigmentation: melanophores occur on nape, over brain and in area of cleithral symphysis; row of elongate melanophores along dorsal aspect of foregut; increasing number of spots along dorsal aspect of hindgut, fewer spots along ventral aspect of hindgut; other melanophores occur along dorsal fin base and lateral body after 16 mm; a few internal melanophores over the anteriormost vertebrae; pigment at notochord tip usually only ventral; pigment on caudal fin increases in larger larvae Transformation occurs at 1623 mm 1. See Clupeidae comparative table on Clupeiformes Introductory page 2. Lower anal fin ray count than in Opisthonema oglinum

Note:

Early juvenile: In early juveniles, last ray of anal fin is longer than other rays, but last ray of dorsal fin is not elongate; no dark spot on body behind opercle

Figures: References:

Adult: Hildebrand, 1963a; AF: Ditty et al., 1994 Fage, 1920; Matsuura, 1972, 1975; Houde and Fore, 1973; E. D. Houde, 1981 (pers. comm.); Munroe and Nizinski, 2002

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Sardinella aurita

A. 3.6 mmSL

B. 5.5 mmSL

C. 8.1 mmSL

D. 11.8 mmSL

E. 14.0 mmSL

F. 19.0 mmSL

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Anchoa hepsetus (Linnaeus, 1758) Engraulidae Striped anchovy

Range: Atlantic coasts of North America and South America from Nova Scotia to Uruguay; absent from Florida Keys and most of Caribbean Sea Schooling, pelagic, in estuarine and oceanic waters; mostly over inner continental shelf; oceanic occurrences not well-described Meristic Characters Myomeres: 4144 Vertebrae: 4044 Dorsal fin rays: 1316 Anal fin rays: 1923 Pectoral fin rays: 1315 Pelvic fin rays: 7 Caudal fin rays: 79+10+9+78

Habitat:

Spawning: Spring and summer; AprSep off NE United States, with peak May Jul; commences near Cape Hatteras, later in southern New England waters; mostly over inner continental shelf; sporadic occurrences until Sep Eggs: Pelagic, elliptical Diameter: long axis 1.201.66 mm; short axis 0.700.94 mm Chorion: smooth and transparent Yolk: segmented Oil globules: none Perivitelline space: narrow

Larvae:

Hatching occurs at 3.64.0 mm; yolk mass tapers posteriorly Body long and slender; anus under dorsal fin Mouth large, terminal, extends to under middle of eye; becomes sub-terminal Flexion occurs between 5 and 10 mm Gut with muscle-band striations posteriorly Sequence of fin ray formation: C, D, A P2 P1 Pigment very light; series of melanophores along dorsal surface of gut and along anal fin base

Note:

1. Amount of overlap between dorsal and anal fins and number of anal fin rays best characters for distinguishing this species from 2 other engraulid species in study area (see Clupeiformes Introduction)

Early Juvenile:

E. 34.0 mmTL

Figures: References:

Adult: Whitehead et al., 1988; Egg, AB: Hildebrand and Cable, 1930; C: Wayne LaRoche (Farooqi et al., 1995); D: Lippson and Moran, 1974; E: Nancy Arthur (Able and Fahay, 1998) (A, B, D redrawn) Able and Fahay, 1998; Berrien and Sibunka, 1999; Munroe, 2000; 2002a; Nizinski and Munroe, 2002

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211

Anchoa hepsetus

A. 3.6 mmTL

B. 5.6 mmTL

C. 10.3 mmSL

D. 15.0 mmTL

212

Anchoa mitchilli (Valenciennes, 1848) Engraulidae Bay anchovy

Range: Habitat: Atlantic coast of North America from Maine to Florida, through Gulf of Mexico to Yucatan Peninsula, Mexico Schooling, pelagic; estuarine and coastal ocean; sandy beaches, open bays, muddy coves

Spawning: Primarily in water <20 m deep in bays, estuaries, sounds and coastal ocean; begins as early as Apr off Cape Hatteras, expands rapidly northward over inner continental shelf waters during May and June, occurs as far north and east as Long Island to Narragansett Bay, Rhode Island in July; declines between September and October Eggs: Pelagic, slightly elliptical Diameter: long axis: 0.841.11 mm Chorion: smooth and transparent Yolk: segmented Oil globules: none Perivitelline space: narrow

Meristic Characters Myomeres: 3841 Vertebrae: 3844 Dorsal fin rays: 1317 Anal fin rays: 2330 Pectoral fin rays: 1112 Pelvic fin rays: 7 Caudal fin rays: 9+10+9+78

Larvae:

Hatching occurs at 1.82.7 mm (smaller than other engraulids); yolk tapers posteriorly Body long and slender, with anus under dorsal fin Mouth large and terminal, extends to middle of eye; becomes subterminal Flexion occurs at 78 mm Gut with muscleband striations posteriorly Sequence of fin ray formation: C, D, A P2 P1 Pigment very light, but somewhat more dense than in larvae of 2 other engraulids in study area; series of melanophores along dorsal surface of gut and along anal fin base Transformation occurs at about 20mm 1. Amount of overlap between dorsal and anal fins and number of anal fin rays best characters for distinguishing this species from 2 other engraulid species in study area (see Clupeiformes Introduction)

Note:

Early Juvenile:

F. 33.0 mmSL

Figures: References:

Adult: Whitehead et al. 1988; Egg, A, C: Kuntz, 1915; B: Lippson and Moran, 1974; D: Mansueti and Hardy, 1967; E: Fowler, 1945; F: Susan Kaiser (Able and Fahay, 1998) (AE redrawn) E. D. Houde, 1982 (pers. commun.); Vouglitois, et al., 1987; Able and Fahay, 1998; Berrien and Sibunka, 1999; Munroe, 2000; 2002a

Early Stages of Fishes in the Western North Atlantic Ocean

213

Anchoa mitchilli

A. 1.9 mmTL

B. 8.4 mmTL

C. 10.0 mmTL

D. 12.0 mmTL

E. 19.5 mmTL

214

Engraulis eurystole (Swain and Meek, 1885) Engraulidae Silver anchovy

Range: Atlantic coast of North America from Massachusetts to northern Gulf of Mexico; also Venezuela to northern Brazil; larvae occur as far north as Scotian Shelf Schooling, pelagic; mostly over continental shelf waters Meristic Characters Myomeres: 4345 Vertebrae: 2527+1719=4345 Dorsal fin rays: 1316 Anal fin rays: 1518 Pectoral fin rays: 1416 Pelvic fin rays: 7 Caudal fin rays: ?+10+9+?

Habitat:

Spawning: Spring into fall; not well-described; larvae rarely collected in estuaries, or as far offshore as Gulf Stream Eggs: Pelagic, elliptical Diameter: long axis: 1.021.25 mm; short axis: 0.500.80 mm Chorion: smooth and transparent Yolk: segmented Oil globules: none Perivitelline space: narrow

Larvae:

Hatching occurs at 23 mm; yolk tapers posteriorly Body long and slender; anus under dorsal fin Mouth large, terminal; extends to middle of eye; becomes subterminal Gut with muscle-band striations posteriorly Air bladder prominent in larger larvae (Fig. D and F) Sequence of fin ray formation: C, D, A P2 P1 Pigment very light, increases with development; note ventral series of melanophores along dorsal surface of gut and along anal fin base; row of elongate spots forms along midisthmus in larger larvae; larvae near transformation develop pigment on head, along lateral midline, on caudal fin rays and on dorsal peritoneum Transformation occurs between 20 and 23 mm 1. Amount of overlap between dorsal and anal fins and number of anal fin rays best characters for distinguishing this species from 2 other engraulid species in study area (see Clupeiformes Introduction)

Note:

Early Juvenile:

F. 27.4 mmSL

Figures: References:

Adult: Whitehead et al., 1988; Egg, AC: Kuntz and Radcliffe, 1917; E: Lippson and Moran, 1974; D, F: Markle et al., 1980 (AC, E redrawn) Markle et al., 1980; Able and Fahay, 1998; Berrien and Sibunka, 1999; Nizinski and Munroe, 2002

Early Stages of Fishes in the Western North Atlantic Ocean

215

Engraulis eurystole

A. 3.2 mmTL

B. 3.4 mmTL

C. 5.2 mmTL

D. 18.0 mmSL

E. 22.8 mmTL