MURAENINAE, Rafinesque, 1815

Smith, David G., Bogorodsky, Sergey V., Mal, Ahmad O. & Alpermann, Tilman J., 2019, Review of the moray eels (Anguilliformes: Muraenidae) of the Red Sea, with description of a new species, Zootaxa 4704 (1), pp. 1-87 : 7-9

Key to the species of the Red Sea morays.

1a. Dorsal and anal fins not confined to posterior end of body, origin of anal fin immediately behind anus... 2 ( MURAENINAE )

1b. Dorsal and anal fins confined to posterior end of body, origin of anal fin well behind anus........ 31 ( UROPTERYGIINAE )

2a. Anus well behind mid-length, preanal length 1.4–1.5 in TL; dorsal-fin origin behind gill opening; color dark brown to orangebrown, with numerous narrow pale yellowish or white bars on head and body; teeth molariform...... Gymnomuraena zebra

2b. Anus near mid-length, preanal length 1.8–2.9 in TL; dorsal-fin origin before or above gill opening; teeth and color variable.. ................................................................................................... 3

3a. Teeth in jaws stout, some molariform, none long and needle-like; vomerine teeth molariform in 2–6 rows...... 4 ( Echidna )

3b. Teeth in jaws moderate to slender, pointed, not molariform, some long and needle-like; vomerine teeth sharp to bluntly pointed ................................................................................................. 5

4a. Head length 8.1–11 in TL; maxillary teeth uniserial; vomerine teeth in 2 rows; color in adults light gray or white, finely flecked with black, with two longitudinal rows of snowflake-like black blotches containing yellow spots............. E. nebulosa

4b. Head length 6.8–7.8 in TL; maxillary teeth biserial; vomerine teeth in 5–6 rows in adults; head and body with alternating broad dark brown and narrow white bars, obscure with growth............................................. E. polyzona

5a. Body slender, depth at anus 37–63 in TL; vertebrae 183–212; eye small, closer to tip of snout than to rictus; grayish brown, paler ventrally, the fins darker............................................................. Strophidon sathete

5b. Body moderately elongate to moderate, depth at anus 11–37 in TL; vertebrae 106–167; eye moderate, over middle of upper jaw; color variable........................................................................................ 6

6a. Jaws strongly arched, meeting only at their tips, long fangs exposed when mouth closed; color plain brown...................................................................................................... 7 ( Enchelycore )

6b. Jaws usually not arched, closing completely and teeth not exposed when mouth is closed; if jaws arched, conspicuous markings present on body....................................................................................... 8

7a. Preanal length 2.3–2.6 in TL; predorsal length 6.0– 8.3 in TL; vomerine teeth biserial, becoming uniserial posteriorly; tip of anterior nostril not flared, posterior nostril before eye; margin of median fins yellow.......................... E. bayeri

7b. Preanal length 2.0– 2.2 in TL; predorsal length 9.5–11 in TL; vomerine teeth uniserial; tip of anterior nostril flared and funnellike, posterior nostril over eye; margin of median fins white................................... E. schismatorhynchus

8a. Posterior nostril long and tubular, its length almost half eye diameter; head anteriorly dark brown, the posterior half and anterior trunk mottled with whitish flecks with three longitudinal rows of large whitish blotches posteriorly....... Muraena helena

8b. Posterior nostril not tubular, at most with a low rim much less than half eye diameter; color not as described above............................................................................................... 9 ( Gymnothorax )

9a. Dorsal-fin origin above gill opening; vomerine teeth biserial, diverging anteriorly in large specimens; color highly variable, usually light tan or gray, usually finely speckled with small dark spots, often grouped to form irregular blotches.... G. pictus

9b. Dorsal-fin origin before gill opening; vomerine teeth in 1 or 2 rows; color not as described above..................... 10

10a. Intermaxillary teeth in 5 longitudinal rows, a median row and 2 lateral rows on each side; body mottled with pale and dark brown or pale grayish brown with about five longitudinal rows of small black spots superimposed anteriorly and with pale flecks posteriorly............................................................................ G. buroensis

10b. Intermaxillary teeth in 3 longitudinal rows, a median row and 1 lateral row on each side; color not as described above.... 11

11a. Color uniform or mainly uniform........................................................................ 12

11b. Color pattern barred, spotted, mottled or with irregular markings............................................... 18

12a. Ground color pale yellowish to whitish gray............................................................... 13

12b. Ground color brown to dark brown...................................................................... 14

13a. Body moderately stout, depth at anus 15–26 in TL; maxillary teeth biserial, vomerine teeth biserial anteriorly; color pale yellowish or whitish gray, densely covered with irregular darker markings; head pores not in pale spots; sensory papillae on head and anterior body black, forming conspicuous lines; fins with inconspicuous light blue edge................... G. griseus

13b. Body slender, depth at anus 31–37 in TL; maxillary and vomerine teeth uniserial; color uniform light tan or gray; head pores in pale spots; sensory papillae not black; fins with a conspicuous light blue edge (white in preservative)...... G. phasmatodes

14a. Predorsal length 11–13 in TL, body elongate, depth at anus 25–34 in TL; vertebrae 143–152; teeth finely serrate on posterior edge; color light brown to tan, often white ventrally, head pores in white spots, fins with narrow pale edge...................................................................................................... G. angusticauda

14b. Predorsal length 6.6–12 in TL, body not elongate, depth at anus 15–25 in TL; vertebrae 111–138; teeth smooth (except G. pindae ); color mainly uniform, head pores not in white spots.................................................... 15

15a. Maxillary teeth biserial, rows equal in length, those in outer row obtusely pointed; dentary teeth blade-like, in one row, except for several larger conical inner teeth anteriorly on each side; color brown, posterior one-fourth of body and fins yellow in smaller specimens, yellow color gradually disappears with growth.................................. G. pseudoherrei

15b. Maxillary teeth uniserial or biserial, when biserial inner row shorter than outer, teeth conical or triangular in form; dentary teeth conical, triangular or retrorse; color not as described above................................................... 16

16a. Teeth triangular, the large anterior ones in both jaws serrate; vertebrae 118–123; ground color uniform dark brown, with fine reticular pattern on body; tail and fins blackish posteriorly.............................................. G. pindae

16b. Teeth slender, conical, smooth; vertebrae 128–138; ground color uniform brown.................................. 17

17a. Preanal length usually 2.0 in TL; preanal vertebrae 58–61; vomerine teeth uniserial or slightly irregular; third median intermaxillary tooth distant from eye; third IO pore under or slightly ahead of anterior margin of eye; color uniform brown except lower jaw and ventral part of branchial area pale (pale yellow in life); head pores unmarked...................... G. hepaticus

17b. Preanal length usually 2.1 in TL; preanal vertebrae 55–57; vomerine teeth biserial in adults; third median intermaxillary tooth close to eye; third IO pore slightly behind anterior margin of eye; color uniform brown, lower jaw and ventral part of branchial area not pale; head pores with dark rim......................................................... G. cinerascens

18a. Body with dark bars or bar-like markings................................................................. 19

18b. Body with spots or irregular markings.................................................................... 21

19a. Teeth smooth; light grayish brown with about 15–20 dark brown bars about as wide as pale interspaces, without superimposed markings; top of head yellow in life............................................................ G. rueppelliae

19b. Teeth serrate; bars with superimposed small spots or irregular markings......................................... 20

20a. Preanal length 2.2–2.4 in TL; trunk with two rows of moderately large irregular brown spots superimposed with small spots on dorsal half and another row midventrally forming short bars posteriorly; markings on branchial region a series of simple spots........................................................................................... G. randalli

20b. Preanal length 2.0– 2.2 in TL; body with 16-20 dark brown bars on ventral two-thirds of body, obscured dorsally by irregular, small spots, forming reticular pattern, dorsal and anal fins with short black bars; markings on branchial region forming horizontal lines................................................................................... G. reticularis

21a. Body with irregular markings........................................................................... 22

21b. Body with discrete pale or dark spots..................................................................... 25

22a. Body yellow, densely mottled with dark brown, snout dark brown; a black blotch over gill opening; fins with a yellow or pale green margin posteriorly.................................................................. G. flavimarginatus

22b. Color with variable irregular pattern, margin of fins not yellow; no black blotch on gill opening...................... 23

23a. Body grayish with irregular dark markings, the lighter background often forming reticulations; head greenish-yellow in life.......................................................................................... G. undulatus

23b. Body brown with pale dendritic markings, never forming a reticulation; head not yellow............................ 24

24a. Preanal length 2.2–2.4 in TL; vertebrae 123–128; pale markings irregular, those on tail forming oblique streaks dorsally........................................................................................ G. pharaonis n. sp.

24b. Preanal length 2.0– 2.1 in TL; vertebrae 137–142; pale markings as rosettes anteriorly, more discrete spots on tail, not drawn out into oblique bars dorsally....................................................................... G. baranesi

25a. Body with dark spots on light background................................................................. 26

25b. Body with light spots on dark background................................................................. 27

26a. Body with small to moderate-size spots, not overlapping, sometimes forming larger blotches; gill opening in a black blotch............................................................................................ G. javanicus

26b. Head and body with moderate to large polygonal black spots separated by narrow white interspaces; gill opening not in a black blotch.................................................................................... G. favagineus

27a. Predorsal length 11–14 in TL; head and trunk with close-set pale spots as large as eye, becoming vertically elongate and bar-like posteriorly; gill opening in a black blotch.......................................................... G. elegans

27b. Predorsal length 6.6–11.0 in TL; spots somewhat uniformly round, not bar-like posteriorly; no black blotch on gill opening.. .................................................................................................. 28

28a. Body covered with white or bluish white spots, increasing in size posteriorly and becoming black-edged; gill opening and inside of mouth yellow in life.................................................................. G. nudivomer

28b. Body of adults with small round or irregular spots; gill opening and inside of mouth not yellow...................... 29

29a. Preanal length 1.8–2.0 in TL; teeth serrate; vomerine teeth biserial or staggered; space between spots greater than spot diameter..................................................................................... G. moluccensis

29b. Preanal length 2.1–2.4 in TL; teeth smooth; vomerine teeth uniserial; space between spots usually subequal to spot diameter. .................................................................................................. 30

30a. Vertebrae 138–144; spots smaller, those on tail and dorsal fin posteriorly close-set, round................... G. punctatus

30b. Vertebrae 131–140; spots larger, those on tail and dorsal fin posteriorly not close-set, those in dorsal fin elongate G. johnsoni

31a. Anus well behind midlength, preanal length 1.5–1.6 in TL; head and body with large brown spots........ Scuticaria tigrina

31b. Anus near or before midlength, preanal length 2.0– 2.4 in TL; color uniform or with irregular markings (except spotted U. polyspilus )................................................................................ 32 ( Uropterygius )

32a. Ground color tan to white, with rounded dark brown spots........................................... U. polyspilus

32b. Head and body without spots........................................................................... 33

33a. Teeth in upper jaw uniserial, wedge-shaped; vertebrae 145–148; color uniform brown........................ U. golanii

33b. Teeth in upper jaw conical, in more than one row; vertebrae 110–140; color variable............................... 34

34a. Teeth in upper jaw in 4 rows, at least anteriorly; color uniform brown...................................... U. genie

34b. Teeth in upper jaw in 2–3 rows; color variable............................................................. 35

35a. Teeth in upper jaw in 3 rows; 0–1 branchial pore; vertebrae 135–140; color tan with large, dendritic, vertically aligned dark brown markings............................................................................ U. nagoensis

35b. Teeth in upper jaw in 2 rows; 1 branchial pore; vertebrae 103–124; color variable................................. 36

36a. Pre-anal fin vertebrae 109–118; color usually uniform brown, sometimes with irregular, indistinct pale markings; head pores in white spots................................................................................. U. concolor

36b. Pre-anal fin vertebrae 92–111; body with reticular or dendritic pattern; head pores not white......................... 37

37a. Head large, the length 6.3–8.3 in TL; color dark with complexly dendritic lighter blotches, not forming reticulations; gill opening below mid-side...................................................................... U. macrocephalus

37b. Head moderate, the length 8.1–9.9 in TL; color light brown or gray with irregular dark brown lines on upper head and body, partly interconnected to form a fine reticulation; gill opening at mid-side.............................. U. micropterus

SUBFAMILY MURAENINAE

Diagnosis. Dorsal and anal fins not confined to posterior end of body; anal fin begins immediately behind anus. Most species with 2 branchial pores, some with 1 and a few with more than 2.