Pseudanthias pillai Heemstra & Akhilesh, 2012

Akhilesh, K. V., Kishore, T. G., Muktha, M., Lisher, M. W., Ambarish, Gop P. & Anulekshmi, C., 2020, Pseudanthias vizagensis, a junior synonym of Pseudanthias pillai Heemstra & Akhilesh, 2012 (Perciformes: Serranidae), Zootaxa 4890 (1), pp. 135-147 : 137-145

Pseudanthias pillai Heemstra & Akhilesh, 2012

English Name: Pillai’s anthias

Figures 2–11 , Table 1

Pseudanthias pillai Heemstra & Akhilesh 2012: 152 , figs. 39 & 40 (type locality: off Chavakkadu, Kerala, India); Psomadakis et al. 2019: 397, pl. XXVIII, fig. 215 ( Myanmar coast).

Pseudanthias sp. Murugan & Namboothri 2012: 107, fig. 2. (Gulf of Mannar, India)

Pseudanthias vizagensis Krishna, Rao & Venu 2017: 215 , fig. 1 & 2. (type locality: Visakhapatnam, India)

Holotype. SAIAB 86517 , male, 119 mm SL, off Chavakkadu , Kerala, India depth 150–200 m, Trawl.

Paratype. SAIAB 97562 , female, 100 mm SL, trawled off Kochi, Kerala, India .

Examined material: Pseudanthias pillai . Arabian Sea: CMFRI- GB 31.1 .139.44.36.1, 84 mm SL; BNHS MF 91 , 94.5 mm SL; CMFRI- GB 31.1 .139.44.36.2, 103.6 mm SL; BNHS MF 92, 102.6 mm SL; BNHS MF 93 , 93.5 mm SL; all trawled at 120-130 m depth off Kollam, India, south-eastern Arabian Sea, collected at Neendakara harbour, 09 May 2018 . Bay of Bengal : CMFRI- GB 31.1 .139.44.36.3, 113.2 mm SL; CMFRI- GB 31.1 .139.44.36.4, 118.8 mm SL; CMFRI - GB 31.1 .139.44.36.5, 112.7 mm SL; BNHS MF 90 , 99.6 mm SL; all trawled at 100 m depth off Chennai, India, Bay of Bengal, collected at Chennai fisheries harbour, 30 December 2019 . Andaman Sea : SAIAB 208471 , 94.8 mm SL, off Ayeyarwady Delta, Myanmar, Andaman Sea, R/V Dr. Fridtjof Nansen, stn.51, trawl, 125 m depth, 03 September 2018 .

Pseudanthias vizagensis : (only syntypes designated) 89–91 mm TL, 44 specimens, no registration numbers (not deposited in MLR, Andhra University , Andhra Pradesh, India as stated by authors), trawled off Visakhapatnam , depth 180 m, Bay of Bengal, Indian Ocean, February 2013 (details from Krishna et al. 2017) .

Diagnosis. The following combination of characters distinguishes Pseudanthias pillai from congeners: dorsalfin rays continuous X, 16–17; third dorsal-fin spine longest; lateral-line scales 36–38; no auxiliary scales on body; no papillae on rear edge of orbit; basal part of dorsal and anal-fins with low scaly sheath; pectoral-fin rays 18–19; posterior margin of pectoral-fin rounded, reaching vertical at base of third anal-fin spine; scale rows above lateralline to fifth dorsal-fin spine 2–2½; circumpeduncular scales 19–22; total gill-rakers 36–39; interopercle and subopercle serrated; interorbital and pre–dorsal scales small; females reddish orange to reddish-pink in colour, scales on dorsum usually with a dark green/ brown spot; caudal-fin mostly yellowish (sometimes with pinkish blotches/ specks medially); narrow purple streak below eye, originating from the anterior margin of maxilla and extending to the pectoral-fin base; males reddish pink above lateral-line with yellow and purple horizontal stripes below lateralline; inverted heart-shaped pink spot on the interorbital region; caudal-fin bright pink with narrow yellow band on posterior margin.

Description. Detailed measurements are presented in Table 1 . Dorsal-fin rays X, 16–17; third dorsal-fin spine longest, slightly longer than fourth, its length0.8–1.0 in 4 th spine length and 2.1–2.8 in HL; fleshy cirrus sometimes present at tips of first five dorsal-fin spines in males; anal-fin rays III, 7; pectoral-fin rays 18–19; pectoral-fin posterior margin rounded, reaching vertical at anal-fin origin to 3 rd anal-fin spine; pelvic-fin pointed or filamentous, reaching base of third anal-fin spine in large males, shorter in females and intersex; lateral-line scales 36–38; scale rows above lateral-line to origin of dorsal-fin 5; scales above lateral-line to fifth dorsal-fin spine 2–2½; gill-rakers on first gill arch 10–11+28–29=38–39.

Body slender and laterally compressed, body depth at dorsal-fin origin 2.6–2.9 in SL; body width at gill opening 2.1–2.5 in HL; body width at anal-fin origin 2.7–3.4 in HL and 0.6–0.8 in orbit length; snout shorter than or subequal to orbit diameter (mostly shorter), snout length 4.8–5.7 in HL; orbit diameter 3.5–4.5 in HL; interorbital area convex in females, intersex and small males, and slightly concave in large males; interorbital width sub-equal to or slightly shorter than orbit diameter, its width 3.9–4.6 in HL.

Head length 2.5–2.7 in SL; body, head, snout, maxilla, suborbital and interorbital scaly (those in the interorbital region small and closely arranged); dorsal profile of head slightly convex; opercle with 3 flattened spines, middle spine acute, upper spine hidden by scales; preopercle vertical edge with fine serrae, slightly larger serrae at angle, 0–3 minute serrae on lower edge; sub and interopercles smooth.

Mouth slightly oblique; tip of lower jaw protrudes slightly beyond tip of upper jaw; maxilla reaching vertical at posterior margin of orbit; lower jaw with band of minute conical teeth; premaxilla with 1–2 pairs of large posteriorly curved teeth anteriorly; posterior edge of maxilla truncate with rounded corners; vomer and palatines with triangular patch of teeth.

Dorsal-fin long, spinous dorsal-fin almost naked; basal region of dorsal and anal-fin rays with low and thin scaly membrane sheath (few scales); pre-dorsal length 2.9–3.0 in SL; dorsal-fin base length 1.6–1.7 in SL; dorsalfin origin slightly anterior to pectoral-fin origin; pre-pelvic length 2.4–2.8 in SL; pelvic-fin spine length 2.1–2.7 in HL; pre-anal length 1.5–1.6 in SL; anal-fin insertion anterior to dorsal-fin insertion; first anal-fin spine short, 4.8–5.3 in HL; second anal-fin spine 2.1–2.6 in HL; anal-fin base length 6.1–6.8 in SL; caudal peduncle slender, caudal peduncle depth 2.7–3.1 in HL and 7.0–8.0 in SL; caudal peduncle length 1.6–2.0 in HL and 3.2-4.1 in SL; shape of caudal-fin varies depending on size and life stage, with posterior margin truncate, deeply emarginate, or convex, sometimes with filamentous extensions. Maximum reported size 151.5 mm TL, 119 mm SL.

Colouration (when freshly collected at port)

Females. Body reddish-orange dorsally becoming reddish-pink ventrally; stomach light pink to whitish; scales on the dorsal half of body usually with dark brownish or olive green spot formed by a cluster of minute melanophores; purple stripe along centre of the posterior half of body and caudal peduncle or several inconspicuous thin purple stripes ( Fig. 2 ); head varies from yellowish ( Fig. 2B ) to reddish ( Fig. 2A ); purple stripe on the cheek, originating at ventral margin of eye and running posteriorly to pectoral-fin base; interorbital and pre-dorsal region reddish-pink; eye yellow with black pupil, narrow inconspicuous purple dorsal margin and pinkish ventral margin; spiny dorsal-fin and anterior dorsal-fin soft rays reddish-orange, intermembrane occasionally with olive green or yellowish patches, posterior dorsal-fin soft rays orange with a yellowish tinge; pectoral and pelvic fins pink; basal half of anal fin pinkish, with yellow distal half; caudal-fin yellow, sometimes with small light pinkish spots medially ( Figs. 2 , 8–11 ). After preservation pale in color and scales above pectorals with melanophores ( Fig. 4 ).

Males. Colours more vivid and beautiful than that of females. Body reddish pink above lateral-line, with yellow and purple horizontal wavy stripes below lateral-line (sometimes broken or scattered); broad yellow saddle originating between head and dorsal-fin origin, extending ventrally behind head to posterior of pectoral-fin base; scales on the dorsal half of body usually with dark brown spot (mostly above lateral-line); stomach whitish to light orange; head with two yellow bands on ventral half, the first from tip of snout running posteriorly through ventral half of eye to pectoral-fin base; the second originating just above posterior half of maxilla, extending posteriorly to opercle margin and slightly onto body; head bands separated by a purplish to reddish pink streak; anterior half of maxilla and snout reddish pink, except for tip of upper jaw which is yellow; interorbital region with an inverted heart-shaped spot bordered by yellow bands; eye yellow with black pupil; pectoral-fin reddish orange, sometimes suffused with yellow; pectoral-fin base yellow with horizontal pinkish stripe on lower third; dorsal-fin yellow with pinkish red basal half on inter-ray membranes; dorsal-fin last two soft rays light orange to reddish; dorsal-fin spines with red filamentous membranes at tip; pelvic-fins yellowish between spinous and soft rays, sometimes pinkish between soft rays one to three, with soft rays three to five whitish; anal-fin yellowish anteriorly, becoming reddish pink posteriorly (sometimes with yellow on inter-ray membranes giving the appearance of spots); caudal-fin bright pink with narrow yellow or orange band on posterior margin ( Figs. 5 , 8–11 ).

Distribution. Pseudanthias pillai has been recorded off the west coast of India, from Ratnagiri (Arabian Sea) to the central east coast of India, Visakhapatnam (Bay of Bengal) and Myanmar (Andaman Sea) at depths of 100–200 m ( Fig. 6 ). The type locality of Pseudanthias pillai in the original description by Heemstra & Akhilesh (2012) is incorrect and should read as “Off Chavakkadu, 10°30’N, 75°24’E, Kerala, India, Arabian Sea, depth 150–200 m ”.

Remarks. Morphometric and meristic data provided by Krishna et al. (2017) appear to be inaccurate, and thus incompatible for comparison (see Gill & Psomadakis 2018). Other distinguishing characters used by Krishna et al. (2017) in table 1 were: 1) Two orange with yellowish diamond shaped bands are present interorbital region; 2) caudal-fin colour; 3) pelvic-fin colour; 4) caudal-fin shape. However, these are characteristic of P. pillai , at least at certain sizes and life stages.

Fresh colour pattern is very important for identification of Pseudanthias species and the comparison of type images ( Fig. 7 ) shows that P. pillai and P. vizagensis are conspecific (also see table 2 in Gill & Psomadakis 2018). However, Heemstra & Akhilesh (2012) did not mention the yellow bands delimiting an inverted heart-shaped pink spot on the interorbital region in P. pillai . This pattern on the anterior region of the head mentioned as a distinguishing character for P. vizagensis by Krishna et al. (2017), is present in most male/intersex specimens of P. pillai examined; in females/smaller specimens the inverted heart-shaped pink spot is absent or less distinct ( Figs. 8–10 ).

The colour pattern of specimens described and depicted by Krishna et al. (2017) are those of large-sized males. As demonstrated in the present study, P. pillai is subject to ontogenetic variation with juveniles, females, and intersex having different body colouration and caudal-fin shape with respect to large males ( Figs. 2–4 , 5 , 8–11 ). The similarities in the male and female colour patterns in some material may indicate that the female has begun to change sex. It is well known that fishes of the genus Pseudanthias are sequential hermaphrodites with females changing their sex to male as they grow, exhibiting ontogenetic variability in colour pattern, sexual dimorphism and morphometry ( Katayama 1978; Erisman & Hastings 2011; Heemstra & Akhilesh 2012). In the present study most specimens of 80 mm TL were observed to be female, with those exceeding 100 mm TL representing either male or intersex individuals ( Figs. 2–4 , 5 , 8–11 ). Contrary to conclusions by Krishna et al. (2017), caudal-fin shape cannot be considered a distinguishing character between species as it varies with sex and size. Male caudal-fin shape of P. pillai varies from lunate to truncate (sometimes with a filamentous upper lobe), or rounded posterior margin with extended upper and lower lobes. Female caudal-fin shape varies from being deeply emarginate or lunate, with or without filamentous extensions on upper and lower lobes ( Fig. 11 ). Species of Pseudanthias often occur in aggregations, with males typically attending large harems of females and immature fish ( Anderson et al. 1990; Anderson & Heemstra 2012). The unusual ( Fig. 1 ) catch mentioned by Kishore et al. (2018) might represent an accidental capture of such an aggregation.

Pseudanthias pillai closely resembles P. emma , a species described from a single specimen collected off Tanintharyi, Andaman Sea ( Myanmar). However, these species can be differentiated based on the description provided by Gill & Psomadakis (2018), with P. pillai having a more slender caudal peduncle (12–14.3 vs. 16.4% SL), larger head (36.7–39.7 vs. 34.3% SL), shorter 3 rd dorsal-fin spine (14.3–18.5 vs. 23.9% SL), and lower number of lower gill rakers (22 vs. 28–29). Furthermore, P. emma differs from P. pillai in colouration by having the sides of the body mostly dominated by orange (vs. orange with bright pink stripes), and the presence of an orange-red blotch on the breast between the bases of pectoral and pelvic-fins (vs. blotch absent).

Meristic (pectoral-fin and lateral-line counts) and morphometric (HL, pre-dorsal fin length, greatest body depth) differences between P. vizagensis and P. pillai were reported by Krishna et al. (2017). However, results from the present study further supports the supposition of Gill & Psomadakis (2018) that the claimed differences most probably reflect diverse and or inaccurate counting methodology by Krishna et al. (2017).

In the Arabian Sea, P. pillai was trawled along with Nemipterus japonicus (Bloch, 1791) , Decapterus russelli (Rüppell, 1830) and Nemipterus randalli Russell, 1986 at 100–120 m depths, with specimens ranging between 85–135 mm TL and 10–45 g in weight ( Kishore et al. 2018). In the Andaman Sea, P.pillai was trawled from a depth of 125 m along with Decapterus smithvanizi Kimura, Katahira & Kuriiwa, 2013 , Priacanthus hamrur (Fabricius, 1775) , Nemipterus sp., Lepidotrigla sp., Pterygotrigla arabica (Boulenger, 1888) , Sacura sanguinea Motomura, Yoshida & Vilasri, 2017 , Saurida sp. etc. In the Bay of Bengal, P. pillai was trawled from 100 m depth along with Chelidoperca investigatoris (Alcock, 1890) , Nemipterus spp., and Parapriacanthus sp.

Figures

FIGURE 1. Pseudanthias pillai landed at Neendakara harbour, Kerala, south-eastern Arabian Sea (photograph from Kishore et al. 2018).

FIGURE 2. Pseudanthias pillai (female): A. collected at Calicut harbour, Kerala (photograph by Said Koya); B. BNHS MF 90 - 99.6 mm SL collected at Chennai fisheries harbour, Tamil Nadu (after transportation).

FIGURE 3. Pseudanthias pillai (female) fresh colouration with an extended caudal-fin upper lobe, collected at Neendakara harbor, Kerala.

FIGURE 4. Pseudanthias pillai: BNHS MF 90, 99.6 mm SL, collected at Chennai fisheries harbour, Tamil Nadu (after preservation).

FIGURE 5. Pseudanthias pillai (male): A. BNHS MF 96, 118 mm SL, collected at Chennai fisheries harbour, Tamil Nadu; B. CMFRI-GB 31.1.139.44.36.2, 103.6 mm SL collected at Neendakara harbor, Kerala.

FIGURE 6. Current geographical distribution of Pseudanthias pillai (green star refers to type locality; red circles refer to confirmed distribution records, black squares current study specimen locations).

FIGURE 7. A. Lateral view of Pseudanthias vizagensis, syntype, 97 mm SL (photograph from Krishna et al. 2017); B. Lateral view of Pseudanthias pillai, holotype, 119 mm SL (photograph from Heemstra & Akhilesh 2012).

FIGURE 8. Antero-dorsal (A & B) and dorsal (C) views of head region: A. Pseudanthias pillai (present study); B. Pseudanthias vizagensis (photograph from Krishna et al. 2017); C. Pseudanthias pillai (SAIAB 208471), collected by the R/V Dr. Fridtjof Nansen off Myanmar, Andaman Sea.

FIGURE 9. Dorsal view of head at different developmental stages of Pseudanthias pillai: A. intersex; B. female. All other individuals represent males.

FIGURE 10. Dorsolateral view of Pseudanthias pillai: A. male; B. intersex/female.

FIGURE 11. Caudal-fin shapes of Pseudanthias pillai at different sizes/life stages (from A–I, females changing to males).

Tables

TABLE 1. Morphometric values (expressed as % SL, except SL and TL which are presented in mm) of Pseudanthias pillai specimens examined. Holotype and paratype data from Heemstra & Akhilesh (2012).

	Holotype SAIAB	Paratype SAIAB	BNHS MF 91	BNHS MF 92	BNHS MF 93	CMFRI- GB 31.1.139.44.36.1	CMFRI- GB 31.1.139.44.36.2	SAIAB	BNHS MF 90	CMFRI- GB 31.1.139.44.36.3	CMFRI- GB 31.1.139.44.36.4	CMFRI- GB 31.1.139.44.36.5
	86517 Arabian Sea	97562 Arabian Sea	Arabian Sea	Arabian Sea	Arabian Sea	Arabian Sea	Arabian Sea	208471 Andaman Sea	Bay of Bengal	Bay of Bengal	Bay of Bengal	Bay of Bengal	min	max
Total length (mm)			123	130.6	119	110.8	129.8	125.3	126.5	141.5	151.5	141.8	110.8	151.5
Standard length (mm) Body depth at dorsal origin Body width at dorsal origin	119 39 16.8	100 35 16	94.5 35.4 17.4	102.6 35.9 17.3	93.5 34.3 18.5	84.2 35.1 17	103.6 38.5 17.8	94.8 36.6 17.5	99.6 38.8 18.6	113.2 36.3 17	118.8 36.5 16.1	112.7 39.5 19.3	84.2 34.3 16.1	118.8 39.5 19.3
Head length	37	38	37.6	36.7	37.3	38.8	39.1	38.2	39.7	38.2	37.4	39.2	36.7	39.7
Snout length	7.3	9.1	6.8	7.1	6.8	7.1	6.9	7.6	8.4	8	7.5	8	6.8	8.4
Orbit diameter	8.7	9.1	10.5	9.5	10.4	10.2	9.7	10.8	8.9	9.1	8.4	8.6	8.4	10.8
Interorbital width	8.1	8.2	9.2	9.6	9	9.5	9.6	8.6	9.3	9.4	9.1	10.1	8.6	10.1
Upper jaw length	15	15	15.6	14.9	16	16	15.5	16.5	15.5	16	15.4	16	14.9	16.5
Maxilla greatest width Caudal peduncle depth Caudal peduncle length lower	5.9 13 22	6.3 12 19	6.6 13.8 18.7	6.2 13 19	7.4 13.6 21.1	6.6 13.4 19.1	7.2 14.3 20.1	7.3 12.6 19	7 13 21.9	6.1 12.8 21.7	6.8 12.6 23.3	7 13.6 22.3	6.1 12.6 18.7	7.4 14.3 23.3
Predorsal length	33	33	33.2	34.4	33.9	34.2	33.9	35	34.1	33.1	33.5	33.4	33.1	35
Preanal length	65	64	65.4	64.2	62.8	68	68.3	62	65.5	66.6	65.6	66.8	62	68.3
Dorsal-fin base length First dorsal-fin spine Second dorsal- fin spine Third dorsal-fin spine Fourth dorsal-fin spine Tenth dorsal-fin spine	62 6.6 12 18 15 13	63 5.6 10 16 14 14	58.4 6.2 10.2 15.9 15.1 13.8	62.3 5.6 8.6 14.5 13 12.7	59.8 6.3 10.2 16.1 13.4 11.9	59.3 6.7 10.6 15.4 14.9 13.5	60.5 6.3 10.4 18.5 14.6 13.6	63.3 7.1 11 16.7 13.8 12.3	61.9 6.5 9.6 14.3 13.7 12.6	59.2 4.5 9.7 15.7 13.1 12.9	59.9 6.5 10.1 14.7 12.9 11.8	59.8 4.7 9.6 15.3 12.5 12.9	58.4 4.5 8.6 14.3 12.5 11.8	63.3 7.1 11 18.5 15.1 13.8
First dorsal-fin ray			15	13.4	12.5	15	16.3	12.7	13.4	15.6	15.1	14.2	12.5	16.3
Longest dorsal- fin ray	25	24	16.8	17.1	23.6	18.2	25.5	20.1	16.3	21.3	23	21.4	16.3	25.5
Anal-fin base length			16.2	16.5	15	14.8	15.4	16.1	16.5	16.4	16.6	16.3	14.8	16.6
Anal-fin length	34	48	33.8	36.8	25.8	33.1	40.5	36.1	37.3	38.2	41	42.1	25.8	42.1

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