Six Species of the Lernanthropidae - Zoological Studies

Zoological Studies 50(5): 611-635 (2011)Six Species of the Lernanthropidae (Crustacea: Copepoda) Parasitic onMarine Fishes of Taiwan, with a Key to 18 Species of the Family Knownfrom TaiwanJu-shey Ho 1 , Wei-Cheng Liu 2 , and Ching-Long Lin 2, *1Department of Biological Sciences, California State University, Long Beach, CA 90840-3702, USA2Department of Aquatic Sciences, National Chiayi University, Chiayi 600, Taiwan(Accepted April 12, 2010)Ju-shey Ho, Wei-Cheng Liu, and Ching-Long Lin (2011) Six species of the Lernanthropidae (Crustacea:Copepoda) parasitic on marine fishes of Taiwan, with a key to 18 species of the family known from Taiwan.Zoological Studies 50(5): 611-635. Six species of copepods belonging to the Lernanthropidae Kabata, 1979were found parasitic on the gill filaments of 6 species of marine fishes of Taiwan. They are: Lernanthropodeschorinemi Pillai, 1962 on Scomberoides commersonnianus Lacepède (Carangidae); Lernanthropodes trachinotiPillai, 1962 on Trachinotus blochii (Lacepède) (Carangidae); Lernanthropus incilis sp. nov. on Evoxymetoponpoeyi Günther (Trichiuridae); Mitrapus heteropodus (Yü 1933) on Nematalosa nasus (Bloch) (Clupeidae);Sagum epinepheli (Yamaguti et Yamasu, 1960) on Epinephelus awoara (Temminck et Schlegel) (Serranidae);and Sagum folium sp. nov. on Paracaesio caerulea (Katayama) (Lutjanidae). Aside from the 2 new species, theother 4 known species were recorded for the 1st time from Taiwan. A key to the 18 species of lernanthropidsoccurring on marine fishes of Taiwan is provided. In this paper we propose treating Mitrapus rubiginosus (Redkar,Rangnekar et Murti 1949) as a junior synonym of M. heteropodus.http://zoolstud.sinica.edu.tw/Journals/50.5/611.pdfKey words: Lernanthropidae, Parasitic copepods, Taiwan, Marine fish.Lernanthropidae Kabata, 1979 is a largefamily of siphonostomatoid copepods comprisingover 150 species. They are exclusively parasiticon gill filaments of marine teleosts. They usetheir prehensile antennae and maxillipeds toattach tenaciously to a host’s gill filaments. Theattachment is assisted, in the case of the female,by leg 3 which is modified into a pair of large,folded lamellae designed for clamping onto a host’sgill filaments. Thus, lernanthropids can often causepathological effects like desquamation, erosion,and necrosis of the host’s gill filaments (Maneraand Dezfuli 2003) and, in cases of heavy infection,may lead to asphyxiation, anemia, and secondarybacterial infections (Tokşen et al. 2006).Lernanthropids are largely parasites ofwarm-water fishes. Thus, while 44 species areknown from India (Pillai 1985), only 9 species areknown from Japan (Ho and Do 1985). So far,we have discovered and reported 12 species oflernanthropids from Taiwan (Ho et al. 2008, Liu etal. 2009a b). In this paper we add 6 more species,which means that 18 species of lernanthropidsin 7 genera are known from Taiwan. Since wehave examined about 20% of the marine fishesof Taiwan, we firmly believe there are morespecies of lernanthropids waiting to be discoveredfrom Taiwan. Therefore, a key to the species oflernanthropids from Taiwan is provided at the endof this report to facilitate species identification ofthis group.*To whom correspondence and reprint requests should be addressed. Tel and Fax: 886-5-2783065. E-mail:cllin@mail.ncyu.edu.tw611

612Zoological Studies 50(5): 611-635 (2011)MATERIALS AND METHODSFish caught and landed at various fishingports in Taiwan in the last 10 yr were purchasedand transferred in an icebox to the laboratory forexamination of parasitic copepods. Parasites werecarefully removed from the host’s gill filamentsunder a dissection microscope, and preservedin 70% ethanol. Selected specimens werelater cleared in 85% lactic acid overnight beforedissection of the appendages and examinationunder a compound microscope with a series ofmagnifications up to 1500x. All drawings weremade with the aid of a drawing tube attached to acompound microscope. Measurements of bodyparts were taken after specimens were soaked inlactic acid overnight. The mean value is given withthe range following in parentheses.Below, a full description is given of thefemale, and if the male is available, only sexuallydimorphic characters are mentioned for the male.RESULTSOrder Siphonostomatoida Thorell, 1859Family Lernanthropidae Kabata, 1979Genus Lernanthropodes Bere, 1936Lernanthropodes chorinemi Pillai, 1962(Figs. 1, 2)Material examined: 2on 1 (of 9) talangqueenfish, Scomberoides commersonnianusLacepsède, 1801, landed at Dong-shi Fishing Porton 13 Jan. 2000.Female: Body (Fig. 1A-C) cylindrical, 3.56(3.50-3.62) mm long (from anterior rim of head totip of caudal ramus), comprising a subtriangularhead, cylindrical trunk, and small urosome. Head0.98 (0.88-1.08) mm long and 0.95 (0.92-0.98) mmwide, with broadly protruding posterolateral cornersand narrowed anterior end. Trunk cylindrical,narrower than head, only 0.82 (0.80-0.84) mmwide, and without a dorsal plate. Posterior partof trunk with ventrally fused lamellae of leg 3appearing wider than head, 1.28 (1.24-1.32) mmwide. Genital complex (Fig. 1D) longer than wide,401 (324-478) × 324 (316-332) μm, with a laterallyprotruding egg sac attachment area. Abdomen(Fig. 1D) also with laterally protruding sides, longerthan wide, 223 (211-235) × 215 (194-235) µm,with distinct anal slit. Caudal ramus (Fig. 1D) along distally attenuated process, 324 (284-365) ×97 (89-105) µm, carrying 2 dorsal setae in basalregion and 2 setae at distal end. Egg sac long andstraight.Antennule (Fig. 1E) filiform and 7-segmented;armature formula: 0, 2, 1, 3, 1, 4, and 4.Parabasal process (Fig. 1E) present. Antenna(Fig. 1F) 2-segmented; corpus about 2.5-timeslonger than claw, bearing 1 broad basal seta onmedial surface; claw bearing 2 similar basal setaeand terminal striations. Mandible comprising 2sections; with 8 teeth on terminal blade. Maxillule(Fig. 1G) bilobate, smaller outer lobe tipped with1 spiniform element and larger inner lobe with 3unequal elements. Maxilla (Fig. 1H) 2-segmented,lacertus unarmed; brachium with denticlesscattered on medial surface and bearing 1 bifidspiniform element subterminally and 1 elementdistally; terminal claw fringed with row of largerdenticles along both edges. Maxilliped (Fig. 2A)2-segmented; corpus unarmed; shaft longer thanclaw, with broad seta on medial margin close todistal end; claw with striations as in antenna.Ventral surface of leg 1 (Fig. 2B) ornamentedwith denticles; both outer and inner setaeof protopod with large basal papilla; exopod1-segmented, large, and tipped with 5 robustspines; endopod a smaller lobe with long terminal,blunt process. Leg 2 (Fig. 2C) with inconspicuousprotopod carrying a short, blunt inner element andwithout outer seta; exopod tipped with 5 spiniformelements; endopod with 1 long, setiform element.Leg 3 with lamelliform rami completely fused toform a broad plate entirely covering urosomeventrally (Fig. 1B) and leaving narrow gap dorsally(Fig. 1A). Leg 4 a pair of long bilobate processesprotruding out of ventral lamella formed by leg 3(Fig. 1A-C). Leg 5 missing.Male: Not collected.Remarks: This is the 1st report of Les.chorinemi outside of India. The 1st report ofthis species was made by Pillai (1962) from gillsof a doublespotted queenfish, Scomberoideslysan (Forsskål, 1775) [named Chorinemuslysan (Forsskål) in the original report], caught offTrivandrum, India. Although both specimens fromTaiwan generally fit the description given by Pillai(1962 1985), some differences in fine structureswere noticed. For instance, the brachium ofthe maxilla in the Indian specimen is equippedsubterminally with a single (instead of double)seta, and leg 2 has an outer (instead of inner) seta.There is a remarkable difference in the size of thespecimens from the 2 places; as the specimenfrom India is 8.2 mm long, while the one from

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 613(A)(C)(F)(H)(G)p4(B)(E)(D)p4crFig. 1. Lernanthropodes chorinemi Pillai, 1962, female. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) habitus, lateral view;(D) urosome with basal part of leg 4, dorsal view; (E) antennule and parabasal process, ventral view; (F) antenna, medial view; (G)maxillule, lateral view; (H) maxilla, medial view. Scale bars: A-C = 0.5 mm; D = 0.2 mm; E and G = 20 μm; F and H = 30 μm. p4: leg 4;cr: caudal ramus.

614Zoological Studies 50(5): 611-635 (2011)Taiwan is only 3.56 mm long on average.Four species of Lernanthropodes arecurrently known, namely Les. cucullus Bere, 1936,Les. natalensis Kensley et Grindley, 1973, and Les.trachinoti Pillai, 1962, in addition to Les. chorinemi.Of these Les. chorinemi is most similar to Les.natalensis in having a triangular-shaped headand without a deep, central notch on the posteriormargin of the fused leg 3 lamellae. Nevertheless,Les. chorinemi differs from this closest congenerby possessing a well-developed parabasal processat the base of the antennules and the structuresof the maxillule (inner lobe with 3, instead of 2,terminal setae) and maxilla (with denticles and bifidspiniform element on the brachium).When Pillai (1985) gave the 2nd report ofLes. chorinemi from India, the hosts were listed asChorinemus sanctipetri and C. lysan. However,according to Froese and Pauly (2011), both ofthem are synonyms of Scomberoides lysan. InPillai’s (1985) 2nd report on Les. chorinemi fromIndia, the male was described. It looks like themale of Lernanthropinus sphyraenae (Yamagutiet Yamasu 1959) previously reported by us fromTaiwan (see Ho et al. 2008) in having both legs 3and 4 comprising a single process.Lernanthropodes trachinoti Pillai, 1962(Figs. 3, 4)Material examined: 1 on 1 (of 4) snubnosepompano, Trachinotus blochii (Lacepède, 1801),landed at Dong-shi Fishing Port on 19 Jan. 2008.Female: Body (Fig. 3A-C) cylindrical, 5.46mm long (from anterior rim of head to posteriormargin of fused leg 3 lamellae), comprising head,trunk, and small urosome. Head squarish, 1.36× 1.18 mm, with anterolateral corners protrudingventrally into rounded knob (Fig. 3C). Trunkcylindrical, slightly wider (1.26 mm wide) thanhead, without dorsal plate. Posterior part of trunkwith ventrally fused lamellae of leg 3 appearingwider (2.02 mm wide) than head. Genital complex(Fig. 3D) slightly wider than long, 478 × 494 µm,with laterally protruding egg sac attachment area.Abdomen (Fig. 3D) also with laterally protrudingsides and wider than long, 267 × 308 µm. Caudalramus (Fig. 3D) elongate, 356 × 162 µm, carrying2 dorsal setae in basal region and 2 setae at distalend. Egg sac long and straight (not illustrated).Antennule (Fig. 3E, F) filiform and indistinctly7-segmented; armature formula: 1, 2, 1, 2, 1, 3 +(A)(B)(C)Fig. 2. Lernanthropodes chorinemi Pillai, 1962, female. (A) Maxilliped, medial view; (B) leg 1, ventral view; (C) leg 2, ventral view.Scale bars: A = 40 μm; B = 20 μm; C = 10 μm.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 615(A)(B)(C)(H)(I)p4(G)(E)(D)p4(F)crFig. 3. Lernanthropodes trachinoti Pillai, 1962, female. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) habitus, lateral view; (D)urosome and leg 4, dorsal view; (E) antennule, ventral view; (F) tip of antennule, ventral view; (G) antenna, medial view; (H) mandible; (I)maxillule, lateral view. Scale bars: A-C = 1 mm; D = 0.5 mm; E = 50 μm; F = 10 μm; G = 0.1 mm; H and I = 30 μm. p4: leg 4; cr: caudalramus.

616Zoological Studies 50(5): 611-635 (2011)1 aesthetasc, and 8 + 1 aesthetasc. Parabasalprocess absent. Antenna (Fig. 3G) 2-segmented;corpus about twice as long as claw, former bearing1 broad basal seta on medial surface; claw bearingsimilar basal seta and terminal striations. Mandible(Fig. 3H) and maxillule (Fig. 3I) essentiallyas in previous species. Maxilla (Fig. 4A, B)2-segmented; lacertus unarmed; brachium bearing1 subterminal seta on medial margin, 1 blunt,terminal element, and large patch of denticles onouter surface; terminal claw fringed with rows ofdenticles along both edges. Maxilliped (Fig. 4C)2-segmented; corpus with fine denticles scatteredin myxal region; shaft longer than claw, with 1subterminal seta on medial margin; claw withstriations as in antenna.Ventral surface of leg 1 (Fig. 4D) with denticlesscattered on protopod and endopod;outer protopodal seta simple and thin, but innerprotopodal seta spiniform and arising from largepapilla; exopod 1-segmented and large, tipped with5 robust spines, inner 2 of which bear denticles onboth sides; endopod smaller than exopod, carrying1 long seta terminally. Leg 2 (Fig. 4E) protopodinconspicuous, without inner and outer setae;exopod armed as in leg 1, but seta on endopodbilaterally denticulate. Leg 3 with lamelliform ramicompletely fused to form a long plate completelycovering urosome ventrally (Fig. 3B) and leavinglarge gap dorsally (Fig. 3A); posterior edge ofventral lamella with 3 indentations (Fig. 3B). Leg 4a pair of long bilobate processes (Fig. 3A) arising(A)(B)(D)(C)(E)Fig. 4. Lernanthropodes trachinoti Pillai, 1962, female. (A) Maxilla, medial view; (B) tip of maxilla, medial view; (C) maxilliped, medialview; (D) leg 1, ventral view; (E) leg 2, ventral view. Scale bars: A = 50 μm; B = 10 μm; C = 0.1 mm; D = 30 μm; E = 20 μm.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 617from basal region of urosome (Fig. 3D). Leg 5absent.Male: Not collected.Remarks: Lernanthropodes trachinoti is sofar known to occur on pompanos (Trachinotus)from India and Australia. In India, like in Taiwan, itwas taken from a snubnose pompano (Pillai 19621985), but in Australia, it was reported from anotherspecies of pompano, Tra. botla (Shaw, 1803)(see Kabata 1979b). It was intriguing to note thespecimen of Les. trachinoti in Pillai’s (1985) 2ndreport differed from his original report (Pillai 1962)in having a triangular head (cephalothorax) and alarge fused lamellae of leg 3 completely concealingthe bifid leg 4 and urosome in ventral view of theanimal. In other words, it may represent a differentspecies of Lernanthropodes. In fact, Pillai (1985)remarked in his 2nd report of Les. trachinoti that“This species closely resembles L. cuculus Bereand distinguished by only minor differences. Theymay turn out to be the same.” Inasmuch as theoriginal description of Les. cuculus is sketchy, nofurther comment can be made at this point. Bere’s(1936) specimens of Les. cuculus were found onTra. carolinus (Linnaeus, 1766) and Tra. falcatus(Linnaeus, 1758) from the Gulf of Mexico.Specimens of Les. trachinoti from Taiwan fitwell with the original report of the species givenby Pillai (1962). The male of this species is notknown from India nor Taiwan, but Kabata (1979b)found it on Tra. botla from Australia.Genus Lernanthropus de Blainville, 1822Lernanthropus incilis sp. nov.(Figs. 5-7)Material examined: 4 and 2 found on gill filaments of Poey’s scabbardfish,Evoxymetopon poeyi Günther, 1887, landed atCheng-gong Fishing Port: 3 and 1 from 3(of 3) E. poeyi on 11 Feb. 2009, and 1 and 1 from 1 (of 1) E. poeyi on 25 Mar. 2009. Femaleholotype (USNM 1131890) and male allotype(USNM 1131891) were deposited in the NationalMuseum of Natural History, Smithsonian Institution,Washington, DC.Female: Body (Fig. 5A-C) large, 7.63 (7.50-7.76) mm long (from anterior rim of head to end ofcaudal ramus), divisible into head, neck, trunk, andurosome. Head nearly squarish, 1.95 (1.92-1.98)× 2.01 (1.80-2.22) mm, with narrowed antennalarea. Neck (1st pediger) short and wide, bearinglarge dorsal lobe. Remaining pedigers fused intotrunk, with pedigers 2 and 3 protruding out to forma lateral lobe and pediger 4 expanded posteriorlyinto a large subcircular dorsal plate that is deeplyemarginated in center. Genital complex andabdomen (Fig. 5D) wider than long, 0.40 (0.38-0.42) × 0.89 (0.84-0.94) and 0.48 (0.46-0.50) × 0.62(0.62-0.62) mm, respectively. Caudal ramus (Fig.5D) transformed into a long process, 2.21 (2.04-2.38) × 0.42 (0.40-0.44) mm, bearing 2 basal setaeon ventral surface (Fig. 5E), 1 subterminal seta onouter margin, and 2 small setae at tip. Egg saclong and straight.Antennule (Fig. 5F, G) stocky, indistinctly5-segmented; armature formula: 0, 0, 0, 0, and 9 +2 aesthetascs. Parabasal process (Fig. 5F) short.Antenna (Fig. 5H) robust, 2-segmented; corpusunarmed; claw armed with basal seta. Mandible(Fig. 6A) and maxillule (Fig. 6B) essentially as inprevious species. Maxilla (Fig. 6C) 2-segmented,with unarmed lacertus larger and longer thanbrachium; latter subterminally bearing 1 short,spiniform process and patch of denticles on medialsurface (usual terminal seta missing); terminalclaw (Fig. 6D) fringed with row of denticles onmedial surface. Maxilliped (Fig. 6E) 2-segmented;corpus robust and unarmed; subchela comprisingsmall, seta-bearing shaft and striated claw.Leg 1 (Fig. 6F) with protopod protruding outinto a process which carries an outer seta at itsbase; protopod also with inner conical process;exopod tipped with 5 stocky spines and endopodwith 1 blunt seta (Fig. 6G). Leg 2 (Fig. 6H) morereduced than leg 1, with inconspicuous protopodand weakly armed exopod (Fig. 6I). Leg 3 (Fig.5B) greatly modified, comprising large fleshy,folded lamella splayed ventrally at posterolateralcorners of trunk (Fig. 5C). Leg 4 (Fig. 5B) a pairof long, bifid processes with round, blunt tip. Leg5 (Fig. 5D) modified into a unilobate, long, obtuseprocess.Male: Body (Fig. 7A, B) smaller than femaleand without dorsal plate on trunk, measuring4.58 mm long (from tip of head to end of caudalramus). Head (cephalosome) wider than long, 1.64× 1.88 mm, with antennal region set apart fromrest of head. First 2 pedigers identifiable by theirlateral swellings, wider than long, measuring 0.24× 1.00 and 0.40 × 1.08 mm, respectively. Genitalcomplex indistinguishably fused to trunk. Caudalramus (Fig. 7A, B) long, slender, 745 × 186 μm,and armed as in female.Antennule (Fig. 7C) stocky as in female, butunsegmented and terminally armed with 3 moresetae (Fig. 7D). Parabasal process with basal

618Zoological Studies 50(5): 611-635 (2011)papilla arising near base of antennule (Fig. 7C).Leg 1 (Fig. 7E) with rows of denticles on outermargin of protopod and medial margin of endopod.Leg 2 (Fig. 7F) carrying a process lateral toexopod, subterminally bearing 1 seta-bearingpapilla and terminally 1 smaller seta-bearingpapilla; exopod with dense patch of denticlesterminally in addition to bearing 4 spiniformelements; endopod with dense patch of denticleson medial surface. Leg 3 (Fig. 7A) modified intopair of long, thin, bifid processes. Leg 4 (Fig. 7A)constructed as in leg 3, but longer and armedwith bifid denticles on distal 1/2 of exopod. Leg 5absent.Etymology: The species name incilis means“cut in” in Latin. It alludes to the possession of a(A)(B)(E)(D)p3p5crp4(C)(F)(H)(G)Fig. 5. Lernanthropus incilis sp. nov., female paratype. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) habitus, lateral view;(D) urosome and leg 5, ventral view; (E) basal part of caudal ramus circled in D; (F) antennule and parabasal process, dorsal view;(G) tip of antennule, ventral view; (H) antenna, medial view. Scale bars: A-C = 1 mm; D = 0.5 mm; E = 25 μm; F = 0.1 mm; G = 20 μm;H = 0.2 mm. p3: leg 3; p4: leg 4; p5: leg 5; cr: caudal ramus.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 619(A)(B)(C)(D)(E)(F)(G)(H)(I)Fig. 6. Lernanthropus incilis sp. nov., female paratype. (A) Mandible; (B) maxillule, lateral view; (C) maxilla, medial view; (D) tipof maxilla, medial view; (E) maxilliped, medial view; (F) leg 1, ventral view; (G) rami of leg 1, ventral view; (H) leg 2, ventral view;(I) exopod of leg 2, ventral view. Scale bars: A and D = 20 μm; B = 50 μm; C, F, and H = 0.1 mm; E = 0.2 mm; G and I = 30 μm.

620Zoological Studies 50(5): 611-635 (2011)deep, central incision on the posterior rim of thedorsal plate.Remarks: More than 100 species are currentlyclassified under Lernanthropus. All of themare characteristic in having a large dorsal platecoming off the posterior border of the trunk. Theposterior margin of this dorsal plate is entire (witheven margin) in most cases. It is only in thefollowing 3 species that we see, like Lus. incilis sp.nov., the presence of a deep, central incision inthe posterior rim of the dorsal plate: Lus. barnardiCapart, 1959; Lus. monodi Delamare-Debouttevilleet Nunes-Ruivo, 1954; and Lus. obscurus Wilson,1913. Nevertheless, the new species from(A)(E)p3p4cr(F)(C)(B)(D)Fig. 7. Lernanthropus incilis sp. nov., male paratype. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) antennule and parabasalprocess, dorsal view; (D) tip of antennule, dorsal view; (E) leg 1, ventral view; (F) leg 2, dorsal view. Scale bars: A and B = 1 mm;C = 0.1 mm; D = 20 μm; E and F = 40 μm. p3: leg 3; p4: leg 4; cr: caudal ramus.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 621Taiwan is distinguished from Lus. barnardi andLus. monodi by the structure of the trunk (with 2lateral indentations). It further differs from Lus.barnardi by the shape of the dorsal plate (largeand comprising 2 subcircular plates) and from Lus.monodi by the short neck and structure of legs 3and 4 (without pointed tip). Both Lus. barnardi andLus. monodi were reported off the west coast ofAfrica (Delamare-Deboutteville and Nunes-Ruivo1954, Capart 1959).As far as the general appearance is concerned,Lus. incilis sp. nov. resembles Lus.obscurus the most. Both of them are also unusualfor Lernanthropus in carrying a pair of plumpantennules. However, Lus. incilis sp. nov. differsfrom Lus. obscurus in the structure of leg 5 (notfoliaceous) and in bearing a large dorsal outgrowthin the neck region. Lernanthropus obscurus is sofar known only from the West Indies (Wilson 1913).Genus Mitrapus Song et Chen, 1976Mitrapus heteropodus (Yü, 1933)(Figs. 8-10)Material examined: 4 and 1 on 3 (of45) Bloch’s gizzard shad, Nematalosa nasus(Bloch, 1795), landed at Sheng-dah Fishing Porton 14 Jan. 1999.Female: Body (Fig. 8A-C) small, but shortand broad, 1.87 (1.62-2.12) mm long (from anteriorrim of head to end of caudal ramus), divisible intohead, short neck, broad trunk, and small urosome;posterior part of trunk (4th pediger) carrying alarge, semicircular dorsal plate. Head slightlylonger than wide, 0.63 (0.58-0.68) × 0.61 (0.60-0.62) mm, with rounded sides. Trunk widestpart of body, with forward protruding shouldersand another forward protrusion found in front ofleg 2 (Fig. 8B). Genital complex and abdomenindistinguishably fused into 1 unit (Fig. 8D).Caudal ramus (Fig. 8D, E) longer than wide, 69(65-73) × 45 (41-49) μm, bearing 3 subterminaland 2 terminal setae. Egg sac straight (notillustrated).Antennule (Fig. 8F) filiform, indistinctly6-segmented; armature formula: 3, 0, 2, 1, 4 +1 aesthetasc, and 8 + 1 aesthetasc. Parabasalprocess absent. Antenna (Fig. 8G) robust,2-segmented; corpus unarmed; claw long andsharply pointed, armed with 2 medial setae inbasal region. Mandible (Fig. 8H) and maxillule (Fig.8I) generally as in previous species. Maxilla (Fig.9A) 2-segmented; lacertus larger than brachiumbut unarmed; latter armed subterminally with 2short setae on medial surface in addition to 1larger terminal seta; terminal claw fringed withrow of denticles around medial margin. Maxilliped(Fig. 9B) 2-segmented; corpus robust, with 1small papilla on myxal surface followed by row ofdenticles; subchela comprising small seta-bearingshaft and long, curved claw.Leg 1 (Fig. 9C) protopod with simple outerseta and apically blunt, spiniform inner seta;1-segmented exopod with 5 robust, denticulatedterminal spines; endopod an inflated lobe bearingdenticles on ventral surface and tipped with ashort, spiniform seta. Leg 2 (Fig. 9D) protopodlacking outer and inner seta; 1-segmented exopodwith 3 denticles scattered on ventral surface andarmed with 4 terminal spines, of which only inner2 are fringed with denticles; endopod armedgenerally as in leg 1. Leg 3 (Fig. 8A-C) greatlymodified, comprising large fleshy, folded lamellasplayed outward at posterolateral corners oftrunk. Leg 4 (Fig. 8A, B) a pair of greatly unequal,bilobate processes, with exopod about 6 times aslong as endopod. Leg 5 missing.Male: Attached to basal region of female leg4 exopod (see Fig. 8B, C). Body (Fig. 10A, B)653 μm long (from tip of head to end of caudalramus). Head (cephalosome) oblong, 338 ×207 μm, with narrowed antennal region. Trunksubrectangular, narrower than head; each of4 pedigers identifiable by its lateral swellings.Genital complex wider than long, 72 × 80 μm,indistinguishably fused to trunk anteriorly and toabdomen posteriorly. Caudal ramus longer thanwide, 34 × 19 μm, shaped and armed as in female.Antennule (Fig. 10C, D) filiform, indistinctly6-segmented; armature formula: 1, 3, 2, 1, 2 + 1aesthetasc, and 9 + 1 aesthetasc. Legs 1 (Fig.10E) and 2 (Fig. 10F) similar to those in female,except for carrying denticles on ventral surface ofprotopod and equipped with longer terminal setaon endopod. Legs 3 and 4 (Fig. 10A, B) eachrepresented by 1 short outer seta in basal regionof small knob on posterolateral corners of trunk.Leg 5 absent.Remarks: When Song and Chen (1976)created Mitrapus to accommodate Lus.heteropodus Yü, 1933, 3 other species ofLernanthropus were included, namely Lus.rubiginosus Redkar, Rangnekar et Murti, 1949,Lus. engraulis Tripathi, 1962, and Lus. oblongusPillai, 1964. Our specimens from Taiwan areidentifiable with either Lus. heteropodus or Lus.rubiginosus. However, we formally consider Lus.

622Zoological Studies 50(5): 611-635 (2011)(A)(G)(B)p3(D)(E)p4(I)cr(C)(F)(H)Fig. 8. Mitrapus heteropodus (Yü, 1933), female. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) habitus, lateral view; (D)urosome, dorsal view; (E) caudal ramus, ventral view; (F) antennule, dorsal view; (G) antenna, ventral view; (H) mandible; (I) maxillule;lateral view. Scale bars: A-C = 0.5 mm; D and G = 50 μm; E, H, and I = 20 μm; F = 30 μm. p3: leg 3; p4: leg 4; cr: caudal ramus.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 623rubiginosus reported by Redkar et al. (1949) tobe conspecific with Lus. heteropodus reported byYü (1933), because of the resemblance betweenthem in the gross morphology of both sexes andthe general structure of the female appendages.Furthermore, according to Redkar et al. (1949)their specimens of Lus. rubiginosus (6 females and6 males) were obtained from Chatoessus nasusDay [= Nematalosa nasus (Bloch)], the samespecies of gizzard shad from which we found ourspecimens of M. heteropodus.The 3 species of Mitrapus show differencesin the ratio of the endopod to the exopod in thefemale leg 4. It is about 1: 16 in M. engraulis,1: 6 in M. heteropodus, and 1: 2 in M. oblongus.Another distinguishing point of these 3 speciesis seen in the structure of their shoulders. Theshoulder of M. engraulis is rounded without ananterior protrusion, with a small anterior protrusionin M. oblongus, and bearing a large anteriorprotrusion in M. heteropodus. It is interesting tonote that the species with the rounded shoulder(M. engraulis) is a parasite of the anchovy(Engraulidae), and the 2 other species withprotruding shoulders (M. heteropodus and M.oblongus) are parasitic on herrings (Clupeidae).Genus Sagum C. B. Wilson, 1913Sagum epinepheli (Yamaguti et Yamasu 1960)(Figs. 11, 12)Material examined: 1 on 1 (of 27) yellowgrouper, Epinephelus awoara (Temminck etSchlegel, 1842), landed at Dong-gang on 27 Dec.2003.Female: Body (Fig. 11A, B) globular, coveredwith denticles on dorsal surface, 3.86 mm long(A)(B)(C)(D)Fig. 9. Mitrapus heteropodus (Yü, 1933), female. (A) Maxilla, medial view; (B) maxilliped, medial view; (C) leg 1, ventral view; (D) leg 2,dorsal view. Scale bars: A = 40 μm; B = 50 μm; C and D = 20 μm.

624Zoological Studies 50(5): 611-635 (2011)(from tip of head to posterior margin of dorsalplate), comprising large head, short neck (1stpediger), rectangular trunk with a large subcirculardorsal plate, and minute, concealed urosome.Head bearing beak-like lateral protrusions, 1.24× 1.82 mm, with both sides turned ventrally (Fig.11C). Neck carrying globular swellings (Fig. 11B)on ventral surface lateral to leg 1 (Fig. 12D).Trunk (Fig. 11A, C) with smooth, shoulder-likeanterolateral corners and posterolateral cornersprotruding to rear along lateral sides of dorsalplate. Components of urosome fused into 1 shortunit (Fig. 11D) and entirely concealed under dorsalplate in dorsal view. Genital complex wider thanlong, 340 × 486 µm. Abdomen also wider thanlong, 146 × 186 µm. Caudal ramus (Fig. 11E) along attenuated process carrying 1 seta and 2knobs in swollen, basal region and 2 setae in distalregion. Egg sac (not shown in Fig. 11) long andcoiled underneath dorsal plate.Antennule (Fig. 11F, G) indistinctly 7-segmented,with armature of 0, 0, 1, 1, 0, 2 and10 + 2 aesthetascs. Antenna broken (see Fig.11B). Mandible (Fig. 11H) as in previous species.Maxillule broken and lost during dissection.Maxilla (Fig. 12A) 2-segmented, with unarmedlacertus; brachium distally bearing 1 patchof denticles and 1 small, blunt element (Fig.12B); terminal claw armed with row of denticlesaround margin. Maxilliped (Fig. 12C) indistinctly3-segmented; corpus unarmed; subchela with 1small subterminal seta on shaft; terminal claw withstriations.Leg 1 (Fig. 12D) protopod missing outer setaand with inner element appearing as a spiniformseta; exopod 1-segmented, tipped with 5 stockyspines; endopod reduced to a simple lobe. Leg2 (Fig. 12E) protopod protruding laterally into(A)(B)(E)p3(D)(F)p4cr(C)Fig. 10. Mitrapus heteropodus (Yü, 1933), male. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) antennule, dorsal view; (D)tip of antennule, dorsal view; (E) leg 1, ventral view; (F) leg 2, ventral view. Scale bars: A and B = 0.1 mm; C, E, and F = 20 μm;D = 10 μm. p3: leg 3; p4: leg 4; cr: caudal ramus.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 625(A)(D)(E)cr(G)p4(F)(C)(B)(H)p3p4Fig. 11. Sagum epinepheli (Yamaguti et Yamasu, 1960), female. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) habitus, lateralview; (D) urosome and leg 4, ventral view; (E) caudal ramus, dorsal view; (F) antennule, dorsal view; (G) tip of antennule, dorsal view; (H)mandible. Scale bars: A-C = 1 mm; D = 0.5 mm; E = 50 μm; F = 0.1 mm; G = 20 μm; H = 30 μm. p3: leg 3; p4: leg 4; cr: caudal ramus.

626Zoological Studies 50(5): 611-635 (2011)large, setulate process; exopod a lobe tipped with3 small spiniform elements; endopod reduced toseta-bearing papilla. Leg 3 (Fig. 11B, C) greatlymodified into fleshy, bent lamella; protopod foldedand protruding ventrally (see Fig. 11C); exopodlarger than endopod, expanded posteriorlyinto a large lamella with dorsal side fused toposteroventral protrusion of trunk (see Fig. 11C);endopod a long lamella concealing urosome inventral view of animal (see Fig. 11B). Leg 4 (Fig.11D) protopod with outer seta; exopod larger thanendopod, but both rami with foliaceous basal partand long, filiform distal part. Leg 5 missing.Male: Not collected.Remarks: The present species was reportedfrom Japan (Yamaguti and Yamasu 1960) andIndia (Pillai and Sebastian 1967). In all instances,just like from Taiwan, the parasites were foundparasitic on gill filaments of groupers belonging tothe genus Epinephelus.Although 11 species of Sagum are listed inthe World of Copepods by Walter (2010), manyof them are so poorly known that a meaningfulcomparison of the morphology between congenersis impossible. Exceptions to this fact are thefollowing 4 species: S. flagellatum Wilson, 1913;S. foliaceum (Richiardi, 1880); S. petersi (vanBeneden, 1852); and S. vespertilio Kabata, 1979.Sagum epinepheli can easily be separated fromS. foliaceum and S. petersi by the presence of(A)(B)(C)(D)(E)Fig. 12. Sagum epinepheli (Yamaguti et Yamasu, 1960), female. (A) Maxilla, medial view; (B) tip of maxilla, medial view; (C)maxilliped, medial view; (D) leg 1, ventral view; (E) leg 2, ventral view. Scale bars: A and C = 50 μm; B = 20 μm; D and E = 30 μm.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 627a pair of lateral horns on the head, and fromS. flagellatum by not having posteroventralprotrusions of the trunk “prolonged backward andoutward like the skirts of a long military cloak”(Wilson 1913).Ho et al. (2008) reported the occurrence ofS. vespertilio on Lethrinus nebulosus (Forsskål)collected from Penghu, Taiwan. In that reportS. tuberculatum Pillai, 1985 was proposed tobe relegated to the synonym of S. vespertilio.Sagum epinepheli can be distinguished from S.vespertilio by having a pair of smaller lateral hornson the head, lacking a lateral process on the neck,carrying a relatively longer, terminal filament oneach ramus of leg 4, and the absence of leg 5.Sagum folium sp. nov.(Figs. 13-15)Material examined: 15 and 3 foundon gill filaments of Japanese snapper, Paracaesiocaerulea (Katayama 1934): 2 on 1 (of 4)P. caerulea, landed at Dong-gang Fishing Porton 10 Oct. 2003; 13 and 3 on 5 (of 6)P. caerulea landed at Cheng-gong Fishing Porton 23 Sept. 2004. Female holotype (USNM1131888) and male allotype (USNM 1131889)were deposited in the National Museum of NaturalHistory, Smithsonian Institution, Washington, DC.Female: Body (Fig. 13A-C) globular, 4.13(4.04-4.22) mm long (from tip of head to posteriormargin of dorsal plate), comprising large head,short neck (1st pediger), semi-rectangular trunkwith a large dorsal plate, and minute urosome.Head slightly longer than wide, 1.25 (1.06-1.44)× 1.19 (1.16-1.22) mm, with both sides turnedventrally. Trunk with sclerites on dorsal andlateral sides and anteriorly protruding shoulders.Urosomal somites fused into 1 short unit (Fig. 13D)and entirely concealed under dorsal plate in dorsalview. Genital complex wider than long, 316 (292-340) × 458 (446-470) µm. Abdomen also widerthan long, 174 (162-186) × 279 (251-308) µm.Caudal ramus (Fig. 13E) leaf-like, inserted intoposterolateral corner of abdomen, carrying 3 short,naked setae in distal 1/2 of dorsal surface andanother 2 setae at distal end. Egg sac (not shownin Fig. 13) long and coiled underneath dorsal plate.Antennule (Fig. 13F, G) indistinctly 7-segmented,with armature formula of 4, 1, 1, 0, 1,2 and 8 + 2 aesthetascs. Antenna (Fig. 14A)2-segmented; corpus carrying 1 small, basalpapilliform element on medial surface; terminalclaw stocky, also carrying similar basal elementon medial surface and apical surface striations.Mandible (Fig. 14B) composed of 2 sections; with8 teeth on terminal blade. Maxillule (Fig. 14C)bilobate; smaller outer lobe tipped with 1 element;larger inner lobe fringed with spinules on distal1/2 of medial margin in addition to carrying 3unequal, terminal elements. Maxilla (Fig. 14D, E)2-segmented, with unarmed lacertus; brachiumbearing 1 subterminal and 1 terminal blunt elementand row of denticles around margin of terminalclaw. Maxilliped (Fig. 14F) 2-segmented; corpuscarrying 1 papilliform element in myxal area;subchela with 1 small subterminal seta on shaftand 1 basal blunt element, median row of minutedenticles and apical striations on terminal claw.Leg 1 (Fig. 14G) with inconspicuous protopodcarrying 1 slender outer seta and 1 spiniform,pinnate inner element; exopod 1-segmented,fringed with setules on outer margin and tippedwith 5 stocky spines; endopod reduced to a lobetipped with a small, blunt element. Leg 2 (Fig.14H) more reduced than leg 1, without protopod;exopod a lobe tipped with 5 blunt elements andendopod with1 blunt element. Leg 3 (Fig. 13B,C) with both rami greatly modified into foliaceousstructure; exopod larger than endopod, occupyingmajor portion of lateral part of trunk (see Fig.13C). Leg 4 (Fig. 13B, D) rami subcylindrical,with setulate basal papilla on outer surface ofprotopod. Leg 5 (Fig. 14I) represented by a bent,blunt process near posterolateral corner of genitalcomplex; carrying 1 setulate papilla subterminallyon medial surface.Male: Body (Fig. 15A, B) smaller than female,1.81 (1.68-1.94) mm long (from tip of head to endof caudal ramus), without dorsal plate on trunk.Head (cephalosome) shaped like a piece of toast,slightly wider than long, 0.89 (0.86-0.92) × 0.91(0.74-1.08) mm. First pediger forming a short neckand remaining pedigers fused to form a rectangulartrunk with conical posterolateral protrusion.Genital complex and abdomen indistinguishablyfused to each other. Caudal ramus (Fig. 15C) alobe measuring 89 (81-97) µm long and 49 (41-57) µm wide, armed as in female.Antennule (Fig. 15D, E) filiform and indistinctly6-segmented, with armature formula of 1, 3,2, 0, 1, and 11 + 2 aesthetascs. Leg 2 (Fig. 15F)protopod with simple outer seta; exopod armed indistal region with 4 stocky spines and 3 patches ofspinules; endopod with single subterminal setuleand tuft of terminal setules. Legs 3 and 4 (Fig.15A, B) represented by pair of bifid cylindrical

628Zoological Studies 50(5): 611-635 (2011)processes coming off both sides of trunk. Leg 5absent.Etymology: The species name foliummeans “leaf” in Latin. It alludes to the unusualtransformation of the caudal rami appearing like apair of leaves at the end of the urosome.Remarks: Without the presence of a pairof lateral horns on the head, the new speciesis closer to S. foliaceum and S. petersi than tothe remaining 3 well-described congeners of S.epinepheli, S. flagellatum, and S. vespertilio.Nevertheless, S. folium sp. nov. can be easilyseparated from those 2 similar species by thestructure of the caudal ramus (being leaf-like) andleg 5 (a bent, blunt, short process subterminallycarrying a medial setulate papilla).Among the 5 well-described species ofSagum, the male is known for 2 species, S.epinepheli and S. foliaceum. The male of thenew species can be easily distinguished from thatof S. foliaceum in having its leg 3 constructed ofbilobate (vs. unilobate) cylindrical processes andfrom that of S. epinepheli in having its large head(cephalosome) shaped like a piece of toast anddistinctly wider than the trunk (fused pedigeroussomites and urosome).(A)(B)(D)p4p5crp3p4(F)(C)(E)(G)Fig. 13. Sagum folium sp. nov., female paratype. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) habitus, lateral view; (D)urosome, showing leg 4, leg 5, and caudal rami, dorsal view; (E) caudal ramus, dorsal view; (F) antennule, ventral view; (G) tip ofantennule, ventral view. Scale bars: A-C = 1 mm; D = 0.5 mm; E = 0.1 mm; F = 50 μm; G = 20 μm. p3: leg 3; p4: leg 4; p5: leg 5; cr:caudal ramus.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 629(D)(A)(B)(E)(G)(F)(C)(H)(I)Fig. 14. Sagum folium sp. nov., female paratype. (A) Antenna, medial view; (B) mandible; (C) maxillule, lateral view; (D) maxilla,medial view; (E) tip of maxilla, medial view; (F) maxilliped, medial view; (G) leg 1, dorsal view; (H) leg 2, dorsal view; (I) leg 5. Scalebars: A = 0.2 mm; B and G = 40 μm; C and I = 50 μm; D and F = 0.1 mm; E = 20 μm; H = 30 μm.

630Zoological Studies 50(5): 611-635 (2011)DISCUSSIONT h e L e r n a n t h r o p i d a e , f o l l o w i n g t h eLernaeopodidae and Caligidae, is the 3rd-largestfamily of fish-parasitizing Siphonostomatoida. Thefamily contains over 150 species, with a greatmajority of them occurring in tropical waters.Thus, in the 3 countries of India, Japan, and theUK, where the fauna of fish-parasitizing copepodsare best known, we see that India stands outas having the most lernanthropid species, at 44species (Pillai 1985), whereas Japan is surroundedby colder water and has only 8 species (Hoand Do 1985) as is the UK with only 5 species(Kabata 1979a). However, in consideration of thenumber of genera, Taiwan contains 7 genera ofthe Lernanthropidae (Lernanthropinus Do in Hoand Do, 1985, Lernanthropodes, LernanthropsisDo in Ho and Do, 1985, Lernanthropus, Mitrapus,Norion von Nordmann, 1864, and Sagum). Indiais known to have representatives from 6 genera(Aethon, Lernanthropinus, Lernanthropsis,Lernanthropodes, Lernanthropus, and Sagum)and the other 2 aforementioned countries have5 genera in each, namely Lernanthropinus,Lernanthropsis, Lernanthropus, Mitrapus, andSagum in Japan (Ho and Do 1985) and Aethon,Lernanthropodes, Lernanthropus, Norion, andSagum in the UK (Kabata 1979a).A c c o r d i n g t o t h e c o p e p o d d a t a b a s eproduced by Boxshall (2011), Lernanthropus isthe largest genus of lernanthropids comprising111 species. As a matter of fact, more than 3/4(75.5% or 111/147) of lernanthropids belong toLernanthropus. This striking uneven compositionof species number is also evident in the(A)(D)(E)(F)(B)(C)p4crp3Fig. 15. Sagum folium sp. nov., male paratype. (A) Habitus, dorsal view; (B) habitus, ventral view; (C) caudal ramus, dorsal view; (D)antennule, dorsal view; (E) tip of antennule, dorsal view; (F) leg 2, dorsal view. Scale bars: A and B = 0.5 mm; C = 25 μm; D = 50 μm;E = 20 μm; F = 30 μm. p3: leg 3; p4: leg 4; cr: caudal ramus.

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 631Table 1. Hosts and localities of 18 species of the Lernanthropidae known from Taiwan. Data in this tableare taken from Bassett-Smith (1898), Boxshall and Montú (1997), Byrnes (1988), Cressey and Collette(1970), Delamare-Deboutteville and Nunes-Ruivo (1954), Gusev (1951), Ho and Do (1985), Ho and Kim(2004), Ho and Sey (1996), Ho et al. (2008), Kabata (1962 1979b), Kensley and Grindley (1973), Kim (1998),Kirtisinghe (1937 1964), Krøyer (1863), Leong (1986), Liu et al. (2009a b), Pillai (1962 1963 1985), Pillaiand Sebastian (1967), Shiino (1955), Shishido (1898), Tripathi (1962), Yamaguti (1936 1954), Yamaguti andYamasu (1959 1960), and Yü (1933). The host names are valid ones following Froese and Pauly (2011).For those articles where a synonym was used for the host, they are identified with a number and noted atthe bottom of the tableParasite Host LocalityLernanthropinus sphyraenae (Yamaguti et Yamasu, 1959) Mene maculata (Bloch et Schneider) TaiwanSphyraena obtusata CuvierSri LankaSphyraena pinguis GüntherJapanLernanthropodes chorinemi Pillai, 1962 Scomberoides commersonnianus Lacepéde TaiwanScomberoides lysan (Forsskål)IndiaLernanthropodes trachinoti Pillai, 1962 Trachinotus blochii (Lacepéde) India, TaiwanTrachinotus botla (Shaw)AustraliaLernanthropsis mugilii (Shishido, 1898) Acanthopagrus schlegelii schlegelii (Bleeker) 2) IndiaMugil cephalus LinnaeusJapan, India, Korea,Taiwan, AustraliaMugil soiuy BasilewskyChina, Sri Lanka, RussiaPagrus major (Temminck et Schlegel) 3) IndiaLernanthropus cadenati Delamare-Deboutteville et Acanthopagrus berda (Forsskål)KuwaitNunes-Ruivo, 1954Elops machnata (Forsskål)TaiwanElops senegalensis ReganSenegalMegalops cyprinoides (Broussonet)India, TaiwanSparidentex hasta (Valenciennes) 4)KuwaitLernanthropus chrysophrys Shishido, 1898 Acanthopatrus australis (Günther) AustraliaAcanthopagrus berda (Forsskål) 5)Japan, India, Australia,TaiwanAcanthopagrus latus (Houttuyn)TaiwanAcanthopagrus schlegelii schlegelii (Bleeker) 2) Japan, TaiwanLernanthropus corniger Yamaguti, 1954 Alepes djedaba (Forsskål) 6) South AfricaMegalaspis cordyla (Linnaeus)Indonesia, India, China,Malaysia, Thailand,TaiwanMene maculata (Bloch et Schneider)TaiwanMyripristis vittata ValenciennesTaiwanRastrelliger kanagurta (Cuvier)IndiaLernanthropus cornutus Kirtisinghe, 1937 Ablennes hians (Valenciennes) US east coast, Haiti,Mexico, Panama, Brazil,Peru, Hawaii, Taiwan,Mauritius,the PhilippinesPlatybelone argalus argalus (Lesueur) Gulf of GuineaStrongylura anastomella (Valenciennes) 7) Japan, KoreaStrongylura exilis (Girard)PeruStrongylura incisa (Valenciennes) 8)Gilbert Is., AustraliaStrongylura leiura (Bleeker) 9)Sri Lanka, Taiwan,the PhilippinesStrongylura marina (Walbaum)British HondurasStrongylura strongylura (Hasselt)Malay Peninsula

632Zoological Studies 50(5): 611-635 (2011)Table 1. (continued)Parasite Host LocalityStrongylura timucu (Walbaum)Strongylura urvillii (Valenciennes)Tylosurus acus acus (Lacepède)Tylosurus acus melanotus (Bleeker)Tylosurus choram (Rüppell)Tylosurus crocodilus crocodilus (Péron etLesueur)Tylosurus punctatus (Günther)Brazilthe PhilippinesPuerto Rico, Angola,Panama, Mexico,Java, the PhilippinesJapan, TaiwanRed SeaUS east coast,Venezuela, Mexico,Panama, Kenya,Zanzibar, Senegal,Madagascar, Red Sea,Mauritius, Seychelles,Gulf of Aden, Senegal,Java, Arabian Gulf,Gulf of Thailand,Borneo, Taiwan,the Philippines, Japan,Hawaiithe PhilippinesLernanthropus giganteus Krøyer, 1863 Carangoides ferdau (Forsskål) IndiaCarangoides praecustus (Bennett)ChinaCaranx crysos (Mitchill)JamaicaCaranx hippos (Linnaeus) 11)Jamaica, BrazilCaranx ignobilis (Forsskål)Sri Lanka, TaiwanCaranx leptolepis CuvierKuwaitCaranx melampygus CuvierAdenCaranx sansun (Forsskål)Sri LankaCaranx senegallus CuvierSenegalCaranx sexfasciatus Quoy et Gaimard TaiwanLernanthropus incilis sp. nov. Evoxymetopon poeyi Günther TaiwanLernanthropus otolithi Pillai, 1963 Otolithes ruber (Bloch et Schneider) 12) IndiaPennahia pawak (Lin)TaiwanPterotolithus maculatus (Cuvier) 13)IndiaLernanthropus pomadasysis Rangnekar and Murti, 1961 Pomadasys kaakan (Cuvier) TaiwanPomadasys maculatus (Bloch)IndiaLernanthropus pristipomoides Kirtisinghe, 1937 Kyphosus vaigiensis (Quoy et Gaimard) TaiwanMitrapus heteropodus (Yü, 1933) Konosirus punctatus (Temminck et Schlegel) China, JapanNematalosa nasus (Bloch)TaiwanNorion priacanthi (Kirtisinghe, 1956) Priacanthus macracanthus Cuvier TaiwanSagum epinepheli (Yamaguti et Yamasu, 1960) Epinephelus akaara (Temminck et Schlegel) JapanEpinephelus awoara (Temminck et Schlegel) TaiwanEpinephelus sp.IndiaSagum folium sp. nov. Paracaesio caerulea (Katayama) TaiwanSagum vespertilio Kabata, 1979 Lethrinus laticaudis (Alleyne et Macleay) 14) AustraliaLethrinus nebulosus (Forsskål)TaiwanSynonyms used in the original publications are: 1) Chorinemus lysan in Pillai (1962); 2) Sparus macrocephalus in Shishido (1898),Shiino (1955), Song and Chen (1976), and Pillai (1985); 3) Pagrosomus major in Pillai (1985); 4) Acanthopagrus cuvieri in Ho and Sey(1996); 5) Sparus longispinis in Yamaguti (1936); 6) Caranx djedaba in Kensley and Grindley (1973); 7) Ablennes anastomella in Hoand Do (1985); 8) Tylosurus incisus in Kabata (1962); 9) Tylosurus leisurus in Kirtisinghe (1964); 10) Strongylura crocodile in Delamare-Deboutteville and Nunes-Ruivo (1954); 11) Caranx carangus in Krøyer (1863); 12) Otolithus argenteus in Pillai (1963); 13) Otolithusmaculates in Pillai (1963); and 14) Lethrinus fletus in Kabata (1979b).

Ho et al. – Copepods Parasitic on Marine Fishes of Taiwan 633lernanthropids of Taiwan, where nine of the 18known species are in Lernanthropus (see Table 1).As stated in the “Introduction”, we believe thatmany more species of lernanthropids are yet tobe discovered from the marine fishes of Taiwan,because so far fewer than 20% of the availablespecies of marine fish from Taiwan have beenexamined for parasitic copepods. Anotherreason why we speculate that more species oflernanthropids will be found from Taiwan is the factthat a large number of lernanthropids reported fromthe Indo-West Pacific region are parasitic on fishesthat are also known to occur in waters of Taiwan.As shown in table 2, as far as we are aware, asmany as 25 species of such lernanthropids in4 genera are yet to be found from the fishes ofTaiwan.To encourage future work on lernanthropidsof Taiwan, a key to the 18 known species of thefamily in Taiwan is provided below. Since the maleof many species of Lernanthropidae are unknownand the taxonomy of the family is largely basedon the morphology of the female, the followingkey is, consequently, applicable only to femalelernanthropids. The publication containing the besttaxonomic description for each species is providedin parentheses following each species name inthis key to facilitate a more-rapid verification of thespecies identification.1. Fourth pediger without dorsal plate ................................... 2- Fourth pediger with a single, large dorsal plate ................ 4- Fourth pediger with a pair of long, widely separated dorsalplates .......................................................................................... Lernanthropinus sphyraenae (Ho et al. 2008: 252-257)2. Fourth pediger with a pair of round, dorsal knobs ..................................Lernanthropsis mugilii (Ho et al. 2008: 257-261)- Fourth pediger without outgrowth; rami of leg 3 fused toform a ventral plate (Lernanthropodes) ..............................33. Head triangular; parabasal process present ................................. Lernanthropodes chorinemi (present report: 612-614)- Head rectangular; parabasal process absent ................................ Lernanthropodes trachinoti (present report: 614-617)4. Egg strings coile..................................................................5Table 2. Species of lernanthropids reported from the Indo-West Pacific region with hosts that also occur inwaters of TaiwanParasite Fish host Locality, reported byLernanthropinus decapteri Decapterus russelli (Carangidae) India, Pillai (1985)Lernanthropinus forficatus Lepturacanthus savala (Trichiuridae) a India, Pillai (1985)Lernanthropinus gibbosus Saurida tumbil (Synodontidae) India, Pillai (1985)Lernanthropinus sauridae Saurida elongata (Synodontidae) Japan, Ho and Do (1985)Lernanthropus abitocephalus Pomadasys maculatus (Haemulidae) India, Pillai (1985)Lernanthropus atrox Acanthopagrus schelegelii (Sparidae) Japan, Ho and Do (1985)Lernanthropus brevicephalus Lutjanus malabaricus (Lutjanidae) India, Pillai (1985)Lernanthropus breviculus Cheilinus chlorourus (Labridae) Australia, Kabata (1979b)Lernanthropus chlamydotus Strongylura strongylura (Belonidae) India, Cressey and Collette (1970)Lernanthropus chirocentrosus Chirocentrus dorab (Chirocentridae) India, Pillai (1985)Lernanthropus dussumieri Dussumieria acuta (Clupeidae) India, Pillai (1985)Lernanthropus gisleri Argyrosomus japonicus (Sciaenidae) Australia, Kabata (1979b)Lernanthropus koenigii Parastromateus niger (Carangidae) India, Pillai (1985)Lernanthropus latis Lates calcarifer (Latida) India, Pillai (1985)Lernanthropus lappaceus Eleutheronema tetrad actylum (Polynemiae) b India, Pillai (1985)Lernanthropus lativentris Lethrinus harak (Lethrinidae) c India, Pillai (1985)Lernanthropus leiognathi Secutor ruconius (Leiognathidae) India, Pillai (1985)Lernanthropus nemipteri Nemipterus furosus (Nemipteridae) Thailand, Ho and Kim (2004)Lernanthropus opisthopteri Opisthopterus tardoore (Clupeidae) India, Pillai (1985)Lernanthropus secutoris Secutor insidiator (Leiognathidae) India, Pillai (1985)Lernanthropus sillaginis Sillago sihama (Sillaginidae) India, Pillai (1985)Lernanthropus triangularis Gerres filamentosus (Gerridae) India, Pillai (1985)Lernanthropus trifoliatus Polydactylus sextarius (Polynemidae) d Sri Lanka, Kirtisinghe (1964)Mitrapus oblongus Sardinella fimbriata (Clupeidae) India, Pillai (1985)Norion tayenus Priacanthus tayenus (Priacanthidae) Thailand, Ho and Kim (2004)aHost was named Trichiurus savala in Pillai’s report (1985: 550). b Host was named Polynemus tetradactylum in Pillai’s report (1985:565). c Host was named Lethrinus rhodopterus in Pillai’s report (1985: 572). d Host was named Polynemus sextarius in Pillai’s report(1985: 603).

634Zoological Studies 50(5): 611-635 (2011)- Egg strings linear................................................................85. Leg 2 present (Sagum) ......................................................6- Leg 2 absent.....Norion priacanthi (Ho et al. 2008: 270-274)6. Rami of leg 4 with lamelliform basal part and filiform distalpart......................................................................................7- Rami of leg 4 subcylindrical.................................................................................Sagum folium (present report: 627-630)7. Neck region (1st pediger) smooth, without outgrowth; leg 5missing............ Sagum epinepheli (present report: 623-627)- Neck region (1st pediger) with a pair of small lateralprocesses; leg 5 present .............................................................................. Sagum vespertilio (Ho et al. 2008: 274-278)8. Endopod of leg 4 as long as, or longer than, exopod(Lernanthropus)...................................................................9- Endopod of leg 4 shorter than exopod.................................................... Mitrapus heteropodus (present report: 621-624)9. Posterolateral corner of head protruding out into a process...........................................................................................10- Head without such a process............................................ 1110. Leg 4 concealed by dorsal plate in dorsal habitus view;caudal ramus a swollen process............................................................Lernanthropus cornutus (Liu et al. 2009a: 40-45)- Leg 4 exposed in dorsal habitus view; caudal ramus acylindrical process............................................................................ Lernanthropus chrysophrys (Liu et al. 2009a: 34-40)11. Head with a pair of large, forwardly protruding horns ....................... Lernanthropus corniger (Liu et al. 2009b: 120-124)- Head without such outgrowths .........................................1212. Dorsal plate on 4th pediger with a deep, central incision .........................Lernanthropus incilis (present report: 617-621)- Dorsal plate on 4th pediger entire, without such an incision ...........................................................................................1313. Head wider than long, triangular or trapezoidal ...............14- Head longer than wide, not shaped as above ..................1514. Antennule with parabasal process; caudal ramus cylindrical.........Lernanthropus pomadasysis (Ho et al. 2008: 266-270)- Antennule without parabasal process; caudal ramus withswollen base............................................................................ Lernanthropus pristipomoides (Liu et al. 2009b: 128-131)15. Dorsal plate large, concealing urosome entirely in dorsalhabitus view.....................................................................................Lernanthropus giganteus (Liu et al. 2009b: 124-128)- Dorsal plate not entirely covering urosome, at leastexposing caudal rami in dorsal habitus view ....................1616. Antennule with parabasal process; leg 5 long, reaching tipof caudal ramus....................................................................................... Lernanthropus otolithi (Ho et al. 2008: 261-266)- Antennule without parabasal process; leg 5 absent ..............................Lernanthropus cadenati (Liu et al. 2009a: 30-34)Acknowledgments: We thank J.H. Hwang ofthe Mi-Tuo Fishing Port, F.Y. Lee of the Sheng-Dah Fishing Port, Ching-Kuo Chang of the Dong-Shi Fishing Port, Dr. S.H. Cheng of the TaiwanFisheries Research Institute in Dong-Gang, andDr. W.C. Chiang of the Eastern Marine BiologyResearch Center of Taiwan Fisheries ResearchInstitute in Cheng-Gong Fishing Port for theirkindness in making the necessary arrangementsfor us to purchase fish landed at those fishingports. We would also like to thank Dr. W.Y. Chen,Director of the Eastern Marine Biology ResearchCenter of Taiwan Fisheries Research Institute,for allowing us to use the laboratory at his centerfor fish examinations. Thanks are also due to 2anonymous reviewers for their comments andsuggestions to improve the quality of this paper.Our appreciation is also given to Y.F. Bai, J.L.Hwang, S.W. Yang, S.C. Hwang, I.C. Ho, M.J.Shih, Y.R. Cheng, Y.Y. Chiang, Y.J. Lin, H.Y. Lin,and M.D. Yü of National Chiayi Univ. for theirassistance in transportation and examination offishes. 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