1

R. R. Eakin

3/6/14

KEY TO GENERA OF ARTEDIDRACONIDAE

(33 species)

1a First dorsal fin at least partly above base of pectoral fin; first dorsal-fin spines 2-7

………………………………………………………….. Artedidraco Lönnberg, 1905

(6 species)

1b First dorsal fin above or slightly behind operculum; first dorsal-fin spines 1-3 ……. 2

2a Posttemporal ridges absent (upper limb of posttemporal bone straight, not arched

upward to form a prominent curved ridge); middle lateral line without tubular

scales; anal-fin rays 13-16; pectoral-fin rays 16-18; caudal-fin rays 8-10 …………..

……………………………………………………………... DolloidracoRoule, 1913

(This is a monotypic genus; Dolloidraco longedorsalis Roule, 1913.) (“Dollo’s plunderfish”)

2b Posttemporal ridges often well developed (upper limb of posttemporal bone

arched upward to form a prominent curved ridge); middle lateral line often with

tubular scales posteriorly; anal-fin rays 15-19; pectoral-fin rays 17-22;

caudal-fin rays 9-10 ………………………………………………………………… 3

3a Interorbital region narrow (bony width 2.2-3.9% SL); first dorsal fin

usually with 3 spines, their length 15-32% SL ...... Histiodraco Regan, 1914

(This is a monotypic genus; Histiodraco velifer Regan, 1914.)

(“sailfin plunderfish”)

3b Interorbital region wide (bony width 3.8-9.0% SL); first dorsal fin

usually with 2 spines, their length 4.5-14.0% SL …...... Pogonophryne Regan, 1914

(25species)

KEY TO THE SPECIES OF THE GENUS ARTEDIDRACO LÖNNBERG, 1905

1a Upper lateral line with 6 or more tubular scales, the row of tubular scales extending

to below 2 or more rays of second dorsal fin ...... 2

1b Upper lateral line with 2-9 tubular scales, the row of tubular scales not extending

beyond first ray of second dorsal fin ...... 4

2a Second dorsal-fin rays 26-30 ...... 3

2b Second dorsal-fin rays 23-26 ...... 5

3a Mental barbel tapered to a point; tubular upper lateral-line scales extending to

below 7 or more rays of second dorsal fin; pectoral fin extending beyond origin

of anal fin ...... A. shackletoniWaite, 1911

(“Shackleton’s plunderfish”)

(includes barred and spotted morphs)

3b Mental barbel expanded distally; tubular upper lateral-line scales not extending

beyond seventh ray of second dorsal fin; pectoral fin not reaching origin of anal

fin ...... A. glareobarbatus Eastman & Eakin, 1999

(“gravelbeard plunderfish”)

(includes barred and spotted morphs)

4a Depth of caudal peduncle 19-23% HL; length of pectoral fin 60-72% HL ………...

...... A. loennbergi Roule, 1913

(“Loennberg’s plunderfish”)

4b Depth of caudal peduncle 23-30% HL; length of pectoral fin 73-88% HL ……...…

...... A. skottsbergi Lönnberg, 1905

(“Skottsberg’s plunderfish”)

5a Upper lateral line with 18-22 tubular scales ...... A. orianae Regan, 1914

(“Oriana’s plunderfish”)

5b Upper lateral line with 6-16 tubular scales ...... A. mirus Lönnberg, 1905

(“South Georgia plunderfish”)

KEY TO SPECIES GROUPS OF THE GENUS POGONOPHRYNE REGAN, 1914

1a Head and body without dark markings ……………. “P. albipinna” (unspotted) group

1b Head and body with dark markings ………………………………………………. 2

2a Top of head without conspicuous dark markings relative to sides of head;

ridges on top of head (supraorbital, parietal, supratemporal, posttemporal)

well developed; second dorsal-fin rays 23-27 (usually 24-26); vertebrae 35-37;

bony interorbital width usually greater than 7% SL; usually a dark saddle-shaped

marking extending from anterior base of second dorsal fin to upper lateral line ……

……………………………………………… “P. scotti” (dorsally-unspotted) group

(“Scott’s plunderfish”; “saddleback plunderfish”)

(The “P. scotti” group consists of a single species, P. scotti Regan, 1914.)

2b Top of head with conspicuous dark markings as on sides of head; ridges on top

of head not well developed or only posttemporal ridges well developed; second

dorsal-fin rays 25-29 (usually 26-28); vertebrae 36-39; bony interorbital width

usually less than 7% SL; no dark saddle-shaped marking between anterior base

of second dorsal fin and upper lateral line …………………….…… (spotted groups) 3

3a Eye not filling orbit anteriorly; pectoral-finrays usually fewer than 20; second dorsal-

fin rays usually 26-27; caudal fin eitherdark (with or without stripes) with light dorsal

and ventral borders or light (and striped)...... “P. marmorata” group

3b Eye filling orbitanteriorly; pectoral-fin rays usually at least 20; second dorsal-fin rays

usually 27-28; caudal fin striped ...... 4

4a Spots on top of head predominantly uniformly small, round and densely distributed

(covering more than half the surface); lower jaw not projecting or projecting slightly

beyond upper jaw; snout sloping steeply (greater than 45° from the horizontal) in

lateral view …………………………………………………… “P. barsukovi” group

4b Spots on top of head predominantly large, round or irregular, and either sparsely

distributed (covering less than half the surface) or in the form of sparsely or densely

distributed vermiculations; lower jaw projecting well beyond upper jaw; snout sloping

gradually (45° or less from the horizontal) in lateral view … “P. mentella” group

KEY TO “P. MENTELLA” GROUP OF POGONOPHRYNE

Mental barbel 20-30% SL ...... “long-barbeled” species

Mental barbel 9-13% SL ...... “short-barbeled” species

Mental barbel 15-20% SL)...... “medium-barbeled” species

KEY TO “MEDIUM-BARBELED” SPECIES OF

“P. MENTELLA”GROUP

1a Terminal expansion of mental barbel with bluntly palmate

processes arranged in a densely packed, scale-like pattern ......

...... P. squamibarbata Eakin and Balushkin, 2000

(“scalebeard plunderfish”)

1b Terminal expansion of mental barbel with flattened, jagged

processes arranged mainly in rows ...... P. fusca Balushkin and Eakin, 1998

(“dusky plunderfish”)

1c Terminal expansion of mental barbel weakly developed; unspotted patch on dorsal

surface of head . . . . .P. bellingshausenensis Eakin, Eastman and Matallanas, 2008

(“spotlessnape plunderfish”)

KEY TO “SHORT-BARBELED SPECIES” OF “P. MENTELLA”

GROUP

1a Ventral surface of abdomen with spots . . P. ventrimaculata Eakin, 1987

(“short-barbeled spotbelly plunderfish”)

1b Ventral surface of abdomen without spots ...... 2

2a Terminal expansion of mental barbel relatively long (50-67% of barbel length)

and pointed, with densely distributed folds, serrated or scalloped

ridges and rosettes of leaf-like processes; second dorsal fin in mature males

relatively high (about 21% SL), with straight (not sinuous distally) rays and a

prominent elevated anterior lobe ......

...... P. brevibarbata Balushkin, Petrov and Prutko, 2010

(“shortbeard plunderfish”)

2b Terminal expansion of mental barbel short (less than 33% of barbel length) and

ovoid, with scale-like, mostly bluntly palmate processes; second

dorsal fin in mature males very high (about 25% SL), with anterior rays

sinuous distally and lacking a prominent anterior lobe ......

...... P. neyelovi Shandikov and Eakin, 2013

(“hopbeard plunderfish”)

2c Mental barbel somewhat tapered and lacking a conspicuous terminal

expansion (less than 33% of barbel length), its flattened tip with

slightly tapered, finger-like processes basally and scale- and leaf-like

serrated processes distally; second dorsal fin in mature males relatively

low (about 15-18% SL), with straight (not sinuous distally) rays and

lacking an anterior lobe ...... P. tronio Shandikov, Eakin and Usachev, 2013

(“turquoise plunderfish”)

KEY TO “LONG-BARBELED” SPECIES OF “P. MENTELLA”

GROUP

1a Ventral surface of abdomen with dark spots; terminal expansion of mental

barbel composed of folds in a honeycomb pattern ......

...... P. favosa Balushkin and Korolkova, 2013

(“honeycombbeard plunderfish”)

1b Ventral surface of abdomen without dark spots; terminal expansion of mental

barbel not composed of folds in a honeycomb pattern ...... 2

2a Terminal expansion of mental barbel orange-colored in fresh (unpreserved)

condition; top of head with relatively few large, rounded and sparsely

distributed spots; vertebrae 36-38 . . . . . P. orangiensis Eakin and Balushkin, 1998

(“orangebeard plunderfish”)

2b Terminal expansion of mental barbel not orange in color; top of head

with many densely distributed spots; vertebrae 38-39 ...... 3

3a Stalk of mental barbel relatively slender; longest spine of first dorsal fin

usually more than 60% as long as longest ray of second dorsal fin ...... 4

3b Stalk of mental barbel relatively stout; longest spine of first dorsal fin

usually less than 60% as long as longest ray of second dorsal fin ...... 5

4a Terminal expansion of mental barbel usually less than 33% of barbel length . . . . 6

4b Terminal expansion of mental barbel usually greater than 33% of barbel

length ...... P. lanceobarbata Eakin, 1987

(“lancebeard plunderfish”)

5a Terminal expansion of mental barbel with irregular, tapered

processes (convolutions sometimes present), its length greater than 60%

of barbel length ...... P.macropogon Eakin, 1981

(“greatbeard plunderfish”)

5b Terminal expansion of mental barbel with brain-like convolutions, its

length less than 60% of barbel length . P. cerebropogon Eakin and Eastman, 1998

(“brainbeard plunderfish”)

6a Terminal expansion of mental barbel with unconnected, irregular flattened

folds oriented longitudinally ...... P. eakini Balushkin, 1999

(“Eakin’s plunderfish”)

6b Terminal expansion of mental barbel with unconnected, bluntly

palmate (resembling a cat’s paw) processes ...... P. mentella Andriashev, 1967

(“longbeard plunderfish”)

KEY TO “P. MARMORATA” GROUP OF POGONOPHRYNE

1a Ventral surface of body without dark spots ...... 2

2a Terminal expansion of mental barbel paddle-shaped, smooth or with

membranous folds; posttemporal ridges not well developed; dorsal

surface of head sparsely spotted ...... P. platypogonEakin, 1988

(“flatbeard plunderfish”)

2b Terminal expansion of mental barbel club-shaped, usually with

brainlike convolutions; posttemporal ridges well developed; dorsal surface

of head densely spotted ...... P. marmorataNorman, 1938

(“marbled plunderfish”)

(includes “dwarf plunderfish” P. minor Balushkin and Spodareva, 2013)

2c Terminal expansion of mental barbel weakly developed, with thin folds;

posttemporal ridges well developed; dorsal surface of head densely spotted

...... P. skorai Balushkin and Spodareva, 2013

(“Skora’s plunderfish”)

1b Ventral surface of body with dark spots ......

...... P. maculiventrata Spodareva and Baluhkin, 2014

(“spotbelly plunderfish”)

KEY TO “P. BARSUKOVI” GROUP OF POGONOPHRYNE

1a Mental barbel usually tapered and without a prominent terminal expansion;

processes, if present, not branched ...... P. barsukoviAndriashev, 1967

(“Barsukov’s plunderfish”; “stubbeard plunderfish”)

(D2 of mature males with dark elevated anterior lobe)

1b Mental barbel with a terminal expansion composed of fingerlike processes,

some of which may be branched ...... P. permitiniAndriashev, 1967

(“Permitin’s plunderfish”; “finespotted plunderfish”)

(D2 of mature males usually without dark elevated anterior lobe???)

KEY TO “P. ALBIPINNA” GROUP OF POGONOPHRYNE

This is a generally deep-water, unspotted group of four species with relatively high numbers of second dorsal-fin rays (27-28) and vertebrae (37-38).

1a Fins light in color; terminal expansion of mental barbel about 40% of barbel

length, composed of rounded, unbranched processes ……………………………… 2

1b Fins dark basally, light distally; mental barbel lacking a well-developed terminal

expansion ………………………………………………………………………….. 3

2a Fins largely white; gill rakers on first gill arch 21; upper lateral-line pores

(tubular scales) 12; mental barbel about 17% SL, its terminal expansion about three

times as wide as stalk ……………………………………… P. albipinna Eakin, 1981

(“whitefin plunderfish”)

2b Fins not white; gill rakers on first gill arch 18; upper lateral-line pores (tubular scales)

19; mental barbel about 22 %SL, its terminal expansion about twice as wide as stalk

…………………………………………………………………. P. dewitti Eakin, 1988

(“DeWitt’s plunderfish”)

3a Mental barbel relatively long (15-20% SL), slender and whip-like; depth of head at

posttemporals less than 20% SL; head concave in dorsal profile; middle lateral-line

pores 8-13 ………………………………P. stewarti Eakin, Eastman and Near, 2009

(“Stewart’s plunderfish”; “whipbeard plunderfish”)

3b Mental barbel short (less than 10% SL) and blunt; depth of head at posttemporals

at least 20% SL; head convex in dorsal profile; middle lateral-line pores 12-29

(usually 16-29) ………………………………………….P. immaculata Eakin, 1981

(“spotless plunderfish”)

REFERENCES

The basis for the keys was Eakin’s (1990) chapter on the Artedidraconidae in Fishes of the Southern Ocean (Gon and Heemstra, 1990; see references therein). The continuing discovery of new species (mostly Pogonophryne) since publication of FSO has added much-needed information about inter- and intraspecific variation within the family. When attempting to key artedidraconids, especially species of Pogonophryne, one should be aware that while every species studied in detail exhibits considerable intraspecific variation in mental barbel structure, this organ is still considered a valid diagnostic character exhibiting consistent morphological differences among most species. Intraspecific barbel variation is more a matter of degree than kind. For example, in P. scotti barbels range from tapered to expanded and the processes covering them vary from small nubs to elaborate convoluted folds. Histological examination, however, reveals a similar basic morphology regardless of degree of development. In contrast, scale-like or filamentous processes versus convoluted folds represent interspecific rather than intraspecific variation. Recent molecular analyses continue to shed light on the complex relationships among artedidraconids.

Balushkin AV. 1999. Pogonophryne eakini sp. nova (Artedidraconidae, Notothenioidei, Perciformes): A new species of plunderfish from the Antarctic. J Ichthyol 39:799- 802.

Balushkin AV, Eakin RR. 1998. A new toad plunderfish Pogonophryne fusca sp. nova (Fam. Artedidraconidae: Notothenioidei) with notes on species composition and species groups in the genus Pogonophryne Regan. J Ichthyol 38:574-579.

Balushkin AV, Korolkova ED. 2013. New species of plunderfish Pogonophryne favosa

Sp. n. (Artedidraconidae, Notothenioidei, Perciformes) from the Cosmonauts Sea

(Antarctica) with description in artedidraconids of unusual anatomical

structures – convexitas superaxillaris. J Ichthyo 53(8): 562-574 (Vopro

Ikhtiol 53(5): 563-576)

Balushkin AV, Petrov AF, Prutko VG. 2010. Pogonophryne brevibarbata sp. nov.

(Artedidraconidae, Notothenioidei, Perciformes) – A new species of toadlike

plunderfish from the Ross Sea, Antarctica. Proc. Zool. Inst. Acad. Sci. USSR

314(4):381-386.

Balushkin AV, Spodareva VV. 2013. Pogonophryne skorai sp. n. (Perciformes:

Artedidraconidae), a new species of toadlike plunderfish from the Bransfield

Strait and coastal waters of the South Shetland Islands, Antarctica. Russian

J of MarBiol 39(3): 190-196, Pleiades Publishing, Ltd. (Biologiya

Morya).

Balushkin AV, Spodareva VV. 2013. Dwarf toad plunderfish Pogonophryne minor sp.

n. (Artedidraconidae; Notothenioidei; Perciformes) – a new species and one of

the smallest species of autocthonous ichthyofauna of marginal seas of the

Antarctic continent. J. Ichthyol 53(1):1-6 (Vopro Ikhtiol 53(1):16-21).

Eakin RR. 1990. Artedidraconidae. In: Gon O, Heemstra PC, editors. Fishes of the Southern Ocean. Grahamstown, South Africa: J.L.B. Smith Institute of Ichthyology. p 332-356.

Eakin RR, Balushkin AV. 1998. A new species of toadlike plunderfish Pogonophryne orangiensis sp. nova (Artedidraconidae, Notothenioidei) from the Weddell Sea, Antarctica. J Ichthyol 38:800-803.

Eakin RR, Balushkin AV. 2000. A new species of Pogonophryne (Pisces: Perciformes: Artedidraconidae) from East Antarctica. Proc Biol Soc Wash 113:264-268.

Eakin RR, Eastman JT. 1998. New species of Pogonophryne (Pisces, Artedidraconidae) from the Ross Sea, Antarctica. Copeia 1998(4):1005-1009.

Eakin RR, Eastman JT, Jones CD. 2001. Mental barbel variation in Pogonophryne scotti Regan (Pisces: Perciformes: Artedidraconidae). Antarct Sci 13:363-370.

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Artedidraconidae) from the Bellingshausen Sea, Antarctica. Polar Biol 31:1175-

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Eakin RR, Eastman JT, Near TJ. 2009. A new species and a molecular phylogenetic

analysis of the Antarctic fish genus Pogonophryne (Notothenioidei:

Artedidraconidae). Copeia 2009(4):705-713.

Eakin RR, Eastman JT, Vacchi M. 2006. Sexual dimorphism and mental barbel structure in the South Georgia plunderfish Artedidraco mirus (Perciformes: Notothenioidei: Artedidraconidae). Polar Biol 30:45-52.

Eastman JT, Eakin RR. 1999. Fishes of the genus Artedidraco (Pisces, Artedidraconidae) from the Ross Sea, Antarctica, with the description of a new species and a colour morph. Antarct Sci 11:13-22.

Eastman JT, Eakin RR. 2001. Mental barbel and meristic variation in the Antarctic notothenioid fish Dolloidraco longedorsalis (Perciformes: Artedidraconidae) from the Ross Sea. Polar Biol 24:729-734.

Fischer W, Hureau, J-C, editors. 1985. FAO species identification sheets for fishery

purposes. Southern Ocean (Fishing Areas 48, 58, and 88) (CCAMLR Convention

Area). FAO, Rome, 2: 471p.

Gon O, Heemstra PC, editors. 1990. Fishes of the Southern Ocean. Grahamstown, South Africa: J.L.B. Smith Institute of Ichthyology. 462 p.

La Mesa M, Vacchi M. 2005. On the second record of the Antarctic plunderfish Artedidraco glareobarbatus (Artedidraconidae) from the Ross Sea. Polar Biol 29:40-43.

Shandikov GA, Eakin RR, Usachev S. 2013. Pogonophryne tronio, a new species of

Antarctic short-barbeled plunderfish (Perciformes: Notothenioidei:

Artedidraconidae) from the deep Ross Sea with new data on Pogonophryne

brevibarbata. Polar Biol 36(2):273-289.

Shandikov GA, Eakin RR. 2013 Pogonophryne neyelovi, a new species of Antarctic

short-barbeled plunderfish (Perciformes: Notothenioidei: Artedidraconidae)

from the deep Ross Sea. ZooKeys 296:59-77.

Spodareva VV, Balushkin AV. 2014. Description of a new species of plunderfish of

genus Pogonophryne (Perciformes: Artedidraconidae) from the Bransfield

Strait (Antarctica) with a key for the identification of species of the group

“marmorata.” J Ichthyol 54(1): 1-6.