A review of the rudderfish genus Tubbia (Stromateoidei: Centrolophidae) with the description of a new species from the Southern Hemisphere

Abstract

A combination of morphological and molecular techniques was used to confirm the existence of a second species of the monotypic centrolophid genus Tubbia. Adults of the seamount rudderfish, T. stewarti sp. nov., which reaches about 56 cm SL, is mesopelagic at depths of 525-1438 m in the temperate waters of the Southern Hemisphere. It has a confirmed dis-tribution off Australia and New Zealand where it occurs sympatrically with the wider ranging T. tasmanica Whitley. Like most other members of the group, juveniles live in the epipelagic zone where they have been taken at 30-50 m depth. The new species has a more robust head, more slender body, more flattened interorbit, longer jaws, denser head pores, rela-tively larger eyes and nostrils, narrower caudal peduncle and more vertebral centra than T. tasmanica, and also differs sub-tly in some morphometric ratios. A rediagnosis of T. tasmanica is also provided.

Full text

Accepted by J. Sparks: 24 Dec. 2012; published: 22 Feb. 2013
ZOOTAXA
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ISSN 1175-5334 (online edition)
Copyright © 2013 Magnolia Press
Zootaxa 3616 (5): 461–477
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http://dx.doi.org/10.11646/zootaxa.3616.5.3
http://zoobank.org/urn:lsid:zoobank.org:pub:C7368437-2498-4ABB-B31A-664CA844A8EC
A review of the rudderfish genus Tubbia (Stromateoidei: Centrolophidae) with
the description of a new species from the Southern Hemisphere
PETER R. LAST1, ROSS K. DALEY1 & GUY DUHAMEL2
1Wealth from Oceans Flagship, CSIRO Marine and Atmospheric Research, GPO Box 1538, Hobart, Tas., 7001, AUSTRALIA .
E-mail: peter.last@csiro.au
2Museum National Histoire Naturelle, Département Milieux et Peuplements Aquatiques, UMR BOREA, CP 26, 43 rue Cuvier 75005
Paris France
Abstract
A combination of morphological and molecular techniques was used to confirm the existence of a second species of the
monotypic centrolophid genus Tubbia. Adults of the seamount rudderfish, T. stewarti sp. nov., which reaches about 56 cm
SL, is mesopelagic at depths of 525–1438 m in the temperate waters of the Southern Hemisphere. It has a confirmed dis-
tribution off Australia and New Zealand where it occurs sympatrically with the wider ranging T. tasmanica Whitley. Like
most other members of the group, juveniles live in the epipelagic zone where they have been taken at 30–50 m depth. The
new species has a more robust head, more slender body, more flattened interorbit, longer jaws, denser head pores, rela-
tively larger eyes and nostrils, narrower caudal peduncle and more vertebral centra than T. tasmanica, and also differs sub-
tly in some morphometric ratios. A rediagnosis of T. tasmanica is also provided.
Key words: Centrolophidae, Tubbia, new species, temperate seas, Southern Hemisphere, Australia, New Zealand
Introduction
The monotypic centrolophid genus Tubbia Whitley, represented by a single nominal species Tubbia tasmanica
Whitley 1943, can be distinguished from all other centrolophid genera by a high vertebral count (i.e. 43–44 centra,
McDowall, 1979) and supposedly by the arrangement of predorsal bones (Ahlstrom et al., 1976); Schedophilus
Cocco, the most morphologically similar centrolophid to Tubbia, has only 25–30 vertebral centra (McDowall,
1979). Tubbia is confined to temperate waters of the Southern Hemisphere, near seamounts and along continental
slopes, where it has been recorded from off southern Australia (Tasmania and nearby seamounts), New Zealand
(southern Campbell Plateau to the South Island) and South Africa (off Natal), to a depth of 850 m (McDowall,
1979; Daley et al., 1997).
Whitley’s (1943) description of T. tasmanica was based on the single juvenile holotype (as 72.5 mm SL by
McDowall, 1979, as well as now) and no other documented records existed until 1979. Until then the validity of the
genus Tubbia remained unclear. Haedrich (1967), in his groundbreaking treatment of the group, had earlier
suggested that T. tasmanica was a likely junior synonym of Schedophilus huttoni (Waite 1910), which also has a
temperate distribution in the Southern Hemisphere. He also stated “unfortunately, the species from the Australian
region are very poorly known. Because of their isolated geographic distribution, critical examination of the species
will doubtless provide much insight into the evolution of the soft-spined centrolophids”. Haedrich and Horn (1972)
later included T. tasmanica in a key to Icichthys Jordan & Gilbert. In 1979, McDowall provided a full redescription
of T. tasmanica based on the holotype, three specimens from New Zealand, and a specimen previously identified
by Smith (1934) as Schedophilus medusophagus (Cocco 1839) from southern Africa.
In November 1992, a substantial collection of adult Tubbia (265–405 mm SL) were taken during a survey of
the fishes of the Pedra Branca seamount, off southeastern Tasmania. The catch was represented by two species of
similar size that differed subtly in the morphology of the head, notably in relative eye and nostril sizes and the

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length of the jaws. They were subsequently found to vary in vertebral counts and exhibited different fixed muscle
enzyme signatures (Daley et al., 1997). An investigation of this material, as well as other newly acquired adult and
juvenile specimens, helped discriminate these forms and resolve nomenclatural issues. In this paper, we provide a
description of the new species and its ontogeny, and a rediagnosis of T. tasmanica.
Material and methods
All specimens were measured after preservation. Measurements and counts are consistent with those of Hubbs and
Lagler (1958) and McDowall (1979) and were taken directly (point to point). They are expressed as percentiles of
standard length (SL) unless otherwise stated (Table 1). All meristic data, other than pectoral-fin counts, were
obtained from x-rays; data from Australian material was compiled by John Pogonoski and incorporated in the
Australian National Fish Collection’s x-ray database; data from New Zealand was compiled by the senior author.
Vertebral counts include the occipital condyle and the hypural centra; the first centrum is often difficult to
distinguish but can be inferred from its link to the first neural spine. The junction between precaudal and caudal
centra is often difficult to distinguish in stromateoid fishes (Ahlstrom et al., 1976). In this study the last precaudal
centra is defined (somewhat artificially) as the centrum giving rise to the last haemal spine preceding the first anal
pterygiophore. The lateral line is usually totally or partly damaged, so no attempt was made to count pored scales.
Counts of pectoral-fin radials, which were difficult to take accurately in most specimens as the ventral-most
elements are obscured by a scaly sheath, are approximate; indicative counts were made for 9 specimens, including
the holotype, but some of these may be erroneous. Similarly, rakers at the base of the gill arch are difficult to access
without dissection; hence, indicative gill-raker counts were extracted from 5 partially dissected individuals.
Four Southern Hemisphere species of the genus Schedophilus, which was found to be consistently
plesiomorphic to Tubbia in a biosystematic analysis of the family Centrolophidae by Bolch et al. (1994), were used
as outgroups to discriminate T. tasmanica from the new species. Muscle tissues represented each taxon (15
specimens of Tubbia tasmanica from the Tasman Sea and New Zealand; 6 T. stewarti new species from Australia
and New Zealand; 3 Schedophilus huttoni from New Zealand; 2 S. maculatus Günther 1860 from New Zealand; 11
S. labyrinthicus McAllister & Randall 1975 from Australia and New Zealand; and 5 S. velaini (Sauvage 1879)
from the Indian Ocean) were selected for genetic analysis from whole specimens and stored at -800C. Following
protocols outlined in Ward et al. (2005), COI sequences were obtained and aligned using ClustalX software
program and analysed in MEGA 4.0 (Tamura et al., 2007). Using the Kimura two parameter (K2P) distance model
(Kimura, 1980), within and between group distance values were calculated and a neighbour-joining tree was
constructed to give a graphical representation of divergence patterns between species (Fig. 8). Bootstrapping was
performed with 1000 replications.
Type specimens of the new species are held in the collections of the Australian National Fish Collection,
Hobart (CSIRO), Museum of New Zealand Te Papa Tongarewa (NMNZ), Museum of Victoria, Melbourne (NMV),
and the Australian Museum, Sydney (AMS); their registration numbers are prefixed with these abbreviations.
Other institutional abbreviations follow Leviton et al. (1985).
Genus Tubbia Whitley
Tubbia Whitley 1943: 178. Type: Tubbia tasmanica Whitley 1943, by original designation.
Definition. Centrolophid fishes with a soft, flabby, elongate and strongly compressed body, its shape varying
ontogenetically, with adults much more elongate than young; dorsal fin single, long-based, low, not notched, with
46–53 elements; anal fin long based, with 32–38 elements; fin spines (4–6 in dorsal and 2–3 in anal fin) soft,
indistinct from and grading into flexible rays; dorsal-fin origin above pectoral-fin base; pectoral and pelvic fins
reduced in adults; bases of unpaired fins covered in thick, scaly skin with deep cutaneous furrows; longitudinal
rows of small pores present on body, most obvious on head and adjacent dorsal and anal fins; skin supported by
short papillose fibres (evident on body as papillae when skin removed); vertebral count relatively high, with 40–45
centra (including hypural).

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Remarks. Ahlstrom et al. (1976) believed the pattern of predorsal bones was so distinctive that Tubbia needed
to be considered as valid. McDowall (1979) followed suit by noting its similarity to some Schedophilus species and
defining Tubbia by its unique vertebral count (i.e. 43–44 total centra including the hypural; a count from Haedrich
and Horn (1972) of 46 centra was confirmed by McDowall as being erroneous). The counts obtained herein of
40–45 extend this range but are still well outside Schedophilus (i.e. 25–30; McDowall, 1979).
Tubbia stewarti sp. nov.
Seamount Rudderfish.
Figs 1A, 2A, 3, 5A; Tables 1–3
Tubbia tasmanica (not Whitley): McDowall, 1979, fig. 4 (misidentification).
Tubbia sp. Daley et al., 1997, p 70, fig.
Holotype. NMNZ P. 033110, 434 mm SL, off Bay of Plenty, New Zealand, 37º 09.6′ S, 177º 07.98′ E, caught by
commercial long line, depth not recorded, collected Gavin James, 1995.
Paratypes. 29 specimens. AMS I 45440–001, 393 mm SL, Pedra Branca seamount, Tasmania, 44º12′ S,
147º10′ E, 650–775 m, 12 Nov 1992; AMS I 45441–001, 368 mm SL, south Tasman Sea, Tasman Sea, 40º58′ S,
164º24′ E, 590–763 m, 19 Jun 2009; CSIRO A 4223, 61 mm SL, Storm Bay, Tasmania, 43º33′ S, 147º27′ E, 40–50
m, 8 Apr 1976; CSIRO H 2653–02, 110 mm SL, off King Island, Tasmania, 40º39′ S, 143º27′ E, 940–965 m, 7 Mar
1989; CSIRO H 2681–01, 88 mm SL, off Maatsuyker Island, Tasmania, Nov 1990; CSIRO H 3967–02, 405 mm
SL, Pedra Branca seamount, Tasmania, 44º12′ S, 147º18′ E, 650–775 m, 12 Nov 1992; CSIRO H 3968–01, 401
mm SL, Pedra Branca, Tasmania, 44º20′ S, 147º22′ E, 705–725 m, 24 Nov 1992; CSIRO H 4372–02, 417 mm SL,
Pedra Branca seamount, Tasmania, 44º11′ S, 147º10′ E, 525–650 m, 11 Apr 1993; CSIRO H 6880–02, 508 mm SL,
south Tasman Sea, Tasman Sea, 40º36′ S, 162º08′ E, ca. 1014 m, 14 Jun 2008; CSIRO H 6966–03, 561 mm SL, St
Helens seamount, Tasmania, 41º15′ S, 148º45′ E, 730–925 m, 18 Jul 2009; CSIRO H 6978–04, 490 mm SL, south
Tasman Sea, Tasman Sea, 40º58′ S, 164º24′ E, 590–763 m, 19 Jun 2009; CSIRO H 7129–01, 400 mm SL, Pedra
Branca seamount, Tasmania, 44º12′ S, 147º10′ E, 650–775 m, 12 Nov 1992; CSIRO T 42, 451 mm SL, off
Bicheno, Tasmania, ca. 42º S, ca. 148º E, 1000 m, 26 Jul 1982; CSIRO T 74, 406 mm SL, off Bicheno, Tasmania,
ca. 42º S, ca. 148º E, 1000 m, 26 Jul 1982; NMNZ P. 007612, 53 mm SL, Dusky Sound, New Zealand, 45º 45′ S,
166º 37′ E, 10 Feb 1973; NMNZ P. 009860, 364 mm SL, Chatham Rise, New Zealand, 42º 51′ S, 176º 44′ W,
880–894 m, 29 Jul 1980; NMNZ P, 011078, 99 mm SL, off North Cape, New Zealand, 34º 20′ S, 173º 16′ E, 30 m,
18 Nov 1978; NMNZ P, 011564, 499 mm SL, off Poor Knights Islands, New Zealand, 35º 14′ S, 175º 19’ E,
875–900 m, 22 Nov 1981; NMNZ P. 012893, 350 mm SL, Chatham Slope, New Zealand, 42º 37′ S, 176º 17′ E,
1065–1070 m, 28 Aug 1982; NMNZ P. 013381, 194 mm SL, Challenger Plateau, New Zealand, 41º 02′ S, 169º 30′
E, 908–910 m, 16 Feb 1983; NMNZ P. 025954, 367 mm SL, Challenger Plateau, New Zealand, 39º 40′ S, 167º 37′
E, 1080–1112 m, 27 Jul 1990; NMNZ P. 028727, 313 mm SL, East Chatham Rise, New Zealand, 43º 36′ S, 173º
52′ W, 1433–1438 m, 1 Jul 1992; NMNZ P. 031996, 110 mm SL, Chatham Rise, New Zealand, 42º 59′ S, 175º 49′
W, 778–780 m, 29 Feb 1992; NMNZ P. 037860, 182 mm SL, Solender Trough, New Zealand, 46º 48′ S, 166º 55′ E,
843–866 m, 13 Dec 2000; NMNZ P. 038753, 315 mm SL, Chatham Rise, New Zealand, 43º 10′ S, 173º 49′ W,
1060–1305 m, 9 Feb 2003; NMNZ P. 040877, 320 mm SL, Chatham Rise, New Zealand, 43º 36′ S, 174º 16′ W,
890–896 m, 23 Jul 2004; NMNZ P. 045146, 502 mm SL, Challenger Plateau, Tasman Sea, 39º 50′ S, 168º 03′ E,
864–886 m, 27 Jun 2005; NMV A 26357–001, 408 mm SL, Pedra Branca seamount, Tasmania, 44º12′ S, 147º13′
E, 525–650 m, 12 Nov 1992; NMV A 26358–001, 429 mm SL, south Tasman Sea, Tasman Sea, 40º57′ S, 164º25′
E, 763–912 m, 19 Jun 2009.
Diagnosis. A species of Tubbia with the following combination of characters: head of adult robust with a
relatively broad and moderately arched interorbit (its width 7.8–8.9% SL in specimens exceeding 300 mm SL);
nostrils large, width of posterior nostril 10–12 times in postorbital head length in adults (exceeding 300 mm SL);
eye relatively large in adults (diameter 5.9–7.0% SL) and close to dorsal margin of head (closest horizontal
distance 2.0–3.2 times in diameter of orbit); jaws relatively large, head length 2.2–2.5 times length of lower jaw;
head pores very dense; caudal peduncle relatively narrow at insertion of anal fin, width 2.4–3.0% SL in juveniles
(less than 110 mm SL), 3.3–5.2% SL in adults (exceeding 300 mm SL); vertebral centra 44–45.

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Description. Proportional measurements of the types are given in Table 1 with specific ratios provided below.
Data for the holotype are presented first; ranges for paratypes are given in parentheses (juveniles 61–110 mm SL
firstly, followed by adults 313–499 mm SL).
TABLE 1. Comparative morphometrics of adult Tubbia stewarti sp. nov., based on the holotype (NMNZ P. 033110, 434 mm
SL) and ranges for five paratypes (313–499 mm SL), and seven adult specimens of T. tasmanica (302–407 mm SL).
Morphometric characters are expressed as percentages of standard length (SL).
Tubbia stewarti Tubbia tasmanica
Holotype Min Max Min Max
Standard length (mm) 434 313 499 302 407
Caudal fork length 114.7 109.2 113.7 111.2 114.9
Predorsal length 28.5 25.7 28.8 22.3 27.3
Preanal length 53.0 50.0 51.1 45.1 51.7
Prepectoral length 27.1 23.7 26.3 23.3 26.3
Prepelvic length 28.3 25.8 27.1 25.5 28.1
Pectoral-anal length 22.7 22.3 23.8 20.3 25.0
Pectoral-pelvic length 7.6 7.6 10.9 6.3 8.2
Pelvic-anal length 24.9 23.8 26.5 20.6 25.5
Head length 27.6 23.7 28.3 23.5 28.7
Snout length (horizontal) 5.2 3.8 5.7 3.4 5.3
Snout length (direct) 6.4 4.5 6.5 5.3 6.4
Postorbital head length 15.1 12.9 14.6 13.4 17.4
Length of naked part of dorsal head 16.1 13.1 16.4 11.0 14.3
Interorbital width 8.1 7.8 8.9 6.4 7.9
Eye diameter 6.6 5.9 7.0 5.2 6.3
Distance eye-top of head (horizontal) 2.1 2.3 3.1 2.8 3.8
Anterior nostril width 0.6 0.7 0.8 0.3 0.8
Posterior nostril width 1.3 1.1 1.3 0.5 1.0
Upper-jaw length 12.8 11.4 12.6 9.6 11.3
Lower-jaw length 11.5 10.7 11.5 8.9 10.3
Gape width 8.6 7.7 9.3 7.6 9.1
Dorsal-fin base length 67.8 63.1 68.0 65.8 70.7
Anal-fin base length 40.3 40.0 43.5 40.8 47.1
Pectoral-fin length 10.5 9.7 12.3 9.5 11.4
Pelvic-fin length 4.8 5.0 6.8 4.8 6.4
Head depth (at isthmus) 14.9 17.4 17.8 15.1 19.1
Head width 11.8 11.0 13.9 10.7 14.0
Body depth (greatest) 30.1 31.3 34.3 33.3 39.3
Body depth (at vent) 29.4 30.1 34.2 32.3 38.1
Body width (across trunk) 11.6 8.8 13.4 10.2 14.3
Caudal peduncle depth (minimum) 9.3 8.8 9.7 9.1 10.5
Caudal peduncle length 6.2 6.1 8.7 5.6 8.1
Caudal peduncle depth (at anal insertion) 10.2 10.1 10.9 10.0 10.9
Caudal peduncle width (at anal insertion) 5.1 3.3 5.2 5.3 6.1

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TABLE 2. Comparative morphometrics of juvenile Tubbia stewarti sp. nov., based on the ranges for four paratypes (60.7–110 mm
SL), and the juvenile holotype (AMS IB 1148, 72.5 mm SL) and two non-type specimens of T. tasmanica (83.7–110 mm SL).
Morphometric characters are expressed as percentages of standard length (SL).
Dorsal-fin elements 50 (48–53); anal-fin elements 34 (32–38); pectoral-fin rays 20 (21–23; caudal-fin
principal rays 9 (9) + 8 (8) = 17 (17), upper procurrent rays 9–13, lower procurrent rays 9–13; total vertebral centra
44–45.
Body elongate oval-shaped; strongly compressed, more so posteriorly; greatest depth from mid-abdomen to
vent; anterior head at snout tip almost vertical in lateral view, its margin moderately convex (usually elevated to
Tubbia stewarti Tubbia tasmanica
Min Max Holotype Min Max
Standard length (mm) 60.7 110 72.5 83.7 110
Caudal fork length 116.2 120.6 114.4 117.2 122.4
Predorsal length 33.5 36.3 34.2 33.9 37.4
Preanal length 57.3 62.3 51.9 49.1 50.3
Prepectoral length 29.0 35.5 31.6 28.3 29.5
Prepelvic length 30.2 33.8 35.4 30.2 32.7
Pectoral-anal length 20.3 20.9 13.7 18.9 20.1
Pectoral-pelvic length 10.5 13.6 8.9 8.2 9.7
Pelvic-anal length 23.7 29.6 15.5 17.7 23.3
Head length 29.7 35.5 32.2 30.2 30.4
Snout length (horizontal) 5.0 7.6 5.5 5.4 5.9
Snout length (direct) 7.2 8.0 7.6 7.2 7.4
Postorbital head length 15.7 17.6 16.5 16.7 17.2
Length of naked part of dorsal head 17.1 21.0 21.7 17.8 18.8
Interorbital width 8.2 12.1 10.5 10.4 11.5
Eye diameter 7.8 11.1 9.0 6.3 7.8
Distance eye-top of head (horizontal) 2.4 4.4 3.9 4.9 6.1
Anterior nostril width 1.1 1.8 0.9 0.7 0.7
Posterior nostril width 2.2 3.0 1.3 1.0 1.4
Upper-jaw length 14.4 16.6 15.3 12.2 13.4
Lower-jaw length 13.9 15.7 13.0 11.4 12.1
Gape width 7.5 10.8 13.2 10.5 10.8
Dorsal-fin base length 64.9 67.1 65.8 67.3 69.0
Anal-fin base length 39.0 43.2 42.4 43.7 45.2
Pectoral-fin length 17.3 20.6 17.7 19.1
Pelvic-fin length 14.5 20.3 16.4 12.8 13.7
Head depth (at isthmus) 19.7 21.6 23.6 22.1 23.8
Head width 12.1 14.2 17.1 14.2 14.3
Body depth (greatest) 37.8 45.8 40.5 38.5 43.5
Body depth (at vent) 37.6 43.9 39.7 39.9 44.5
Body width (across trunk) 7.0 12.3 10.0 9.9 12.5
Caudal peduncle depth (minimum) 8.9 13.4 11.1 11.2 11.6
Caudal peduncle length 1.9 4.7 4.3 5.1 5.6
Caudal peduncle depth (at anal insertion) 8.1 11.1 11.3 11.1 11.4
Caudal peduncle width (at anal insertion) 2.4 3.0 3.5 4.6 4.7

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hind orbit, then becoming less so to dorsal-fin origin); abdomen weakly convex (usually extended slightly); profile
at bases of dorsal and anal fins convex, tapering on tail at caudal peduncle; caudal peduncle relatively deep (almost
uniform in depth anteriorly and posteriorly), strongly compressed; procurrent rays of caudal fin forming keel-like
expansions on dorsal and ventral edges of tail.
FIGURE 1. Lateral views of adult: A, Tubbia stewarti sp. nov., holotype NMNZ P. 033110, 434 mm SL (fresh); B, T.
tasmanica, CSIRO H 6853–12, 306 mm TL (fresh).
Head relatively large, deep, length 27.6% (29.7–35.5% and 23.7–28.3% SL in juveniles and adults
respectively); moderately compressed and robust, width 43% (34–45%, 44–52%) of length, slightly to much
broader than trunk. Snout short, blunt, its horizontal length 19% (16–22%, 16–21%) of head length. Nostrils
relatively enlarged, openings closely adjacent, positioned slightly forward of or at midpoint of snout and mostly
below level of mid eye; anterior opening simple, subcircular, tubular; posterior opening narrow, more slit-like,
subvertical, its length usually exceeding aperture of anterior nostril. Eye relatively large, 1.3 (1.2–2.0, 1.1–1.7)
times horizontal snout length, 4.2 (3.0–4.0, 3.6–4.2) in head length; lateral on head, not protruding but marginally
visible when viewed front on; relatively close to dorsal surface, shortest horizontal distance from eye to dorsal
margin of head 3.2 (1.7–4.4, 2.0–2.9) in eye diameter; interorbital space relatively broad, width 1.2 (1.0–1.2,
1.2–1.4) in eye diameter; not strongly arched; weak to well-developed supraorbital crest present.
Mouth large, slightly oblique, length of lower jaw 2.4 (2.1–2.3, 2.2–2.5) in head length; not protrusible,
subterminal to slightly inferior; lips narrow, firm; maxilla elongate, slender, extending almost to hind margin of
eye; gape narrow, 1.5 (1.5–2.2, 1.2–1.5) in length of maxilla; lower jaw partly enclosed within upper jaw when
mouth closed; a tall, bony, chisel-edged ridge along each jaw bearing small, villiform teeth. Teeth similar in size
and shape; uniserial around entire length of both jaws (including premaxilla and dentary); absent from vomer and
palatines; roof of mouth and tongue covered with numerous fine, tooth-like papillae. Inner anterior margins of
mandibles not united, separated by triangular fleshy anterior portion of branchial arch.

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FIGURE 2. View of the head of adult: A, Tubbia stewarti sp. nov., paratype, CSIRO H 3968–01, 401 mm SL (preserved); B,
T. tasmanica, CSIRO H 3967–03, 392 mm TL (preserved).

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FIGURE 3. Lateral views of juveniles of Tubbia stewarti sp. nov.: A, NMNZ P. 007612, 57 mm SL (illustration by McDowall,
1979); B, CSIRO H 2681–01, 87.8 mm TL (fresh); C, NMNZ P. 011078, 98.9 mm TL (preserved).

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Gill openings extensive, opercular flap well developed, postorbital length 1.5 (1.5–2.1, 2.0–2.3) times eye
diameter. Opercular margin irregular with fine soft spines and papillae; variably developed, soft, flexible, thallate
spine at posterior extremity, sometimes with an obscure smaller spine above. Preopercular margin irregular, finely
spinose and papillate; anteriormost part posteroventral to angle of jaw; posteroventral extremity narrowly rounded
(following contour of operculum); upper part oblique, directed anterodorsally, terminating just above horizontal
level of mid eye. Opercular membranes not united, free from isthmus. Gill rakers on first arch relatively large,
elongate, 4–7+1+13 (n=6); on lower limb decreasing in length anteriorly (ca. 13 mm to 4 mm in length) from
angle; sickle shaped, bases of longest rakers stout, tapering strongly to a point distally; posterior inner margins with
fine hair-like spines.
Dorsal fin with 48–53 total elements; single, long, low, continuous; anteriorly with 4–6 short, soft spines
followed by 43–47 rays, usually obscured, origin indistinct, over hind margin of operculum; insertion more or less
over anal-fin insertion; base indistinct, embedded and merging with body; soft spines grading anteriorly into longer
rays; all elements raked posteriorly, even when fin raised; tips of posterior rays well short of caudal fin when
depressed; anterior rays simple or weakly branched (sometimes narrowly divided distally); rays 16–21 longest,
subequal to or shorter than pectoral fin. Anal fin similar in shape to dorsal fin, with 32–38 total elements; base
obscured with 2–3 short, soft spines followed by 31–36 rays; posterior rays falling well short of caudal-fin base
when depressed; rays 1–2 reduced; rays 7–11 longest, subequal to longest rays of dorsal fin. Pectoral fin short,
with 20–21 rays; paddle-shaped to rounded, base relatively broad, subvertical to weakly oblique, situated below
level of eye; rays mostly weakly branched, lowermost rays fully or partly concealed under a scaly sheath of skin.
Pelvic fins greatly reduced in adult, with 1 flexible spine and 5 rays; bases closely adjacent and situated below
pectoral-fin bases, depressible into shallow groove on anteroventral midline of abdomen. Caudal fin with 17
primary elements (9 upper, 8 lower), 19–26 shorter procurrent rays; fin well developed, broad based, forked
(appearing emarginate when damaged), with narrowly rounded tips; caudal peduncle deeper than long, depth of
body at vent 3.1 (3.3–4.6, 3.2–3.9) times minimum depth of caudal peduncle.
Body and fins sheathed in thick skin covered with small deciduous scales (intact in holotype but variably
abraded in most paratypes). Scales cycloid, weakly imbricated, variable in shape; absent on orbital membrane,
nostrils, lips and on sensory pores (scales adjacent sensory pores usually modified greatly in shape to fit around
pore); their size variable, largest on flanks, expanded longitudinally on dorsal and anal fins, very small on caudal,
pectoral and pelvic fins. Skin on flanks and over unpaired fins thick; densely covered with indistinct longitudinal
rows of small, weakly or non-emergent pores; pores closely and almost equidistantly spaced; externally, more
prominent beside bases of dorsal and anal fins than on central flank; pores more evident on undersurface of skin than
on external surface; papillose fibres connect skin to musculature on sides, distal edges broken when skin removed
(Fig. 6). Pale pores most prominent on head, slightly elevated; particularly prominent on snout, around orbit, and on
lower jaw; pores on anterior nape and upper snout somewhat reticulate in appearance, merging with regular
predorsal scaled patch on midline of nape above hind margin of eye. Lateral line usually distinct, slightly undulatory,
with ca. 130 pored scales in holotype (ca. 139 in CSIRO H 3968–01); arched weakly anteriorly, following contour of
dorsal surface to caudal-fin base; scales modified slightly but barely distinct from those adjacent.
Ontogenetic variation. Like most other members of the family, juveniles differ markedly from adults in
morphology and colour. Also, as adults and juveniles are soft-bodied, considerable intraspecific variability in
morphometrics might be expected. Ontogenetic differences were most evident in measurements about the head,
from the snout to the dorsal and anal-fin origins, distance between fin bases, and pectoral and pelvic-fin lengths. In
summary, juveniles (less than 110 mm SL) have a relatively shorter pectoral-anal length (20.3–20.9% vs.
22.3–23.8% SL), caudal peduncle width at anal-fin insertion (2.8–3.0% vs. 3.3–5.2% SL) and caudal peduncle
(1.9–4.7% vs. 6.1–8.7% SL), and a larger predorsal length (33.5–36.3% vs. 25.7–28.8% SL), preanal length
(57.3–62.3% vs. 50.0–51.1% SL), head length (29.7–35.5% vs. 23.7–28.3% SL) and scaleless part of head
(17.7–21.0% vs. 13.1–16.4% SL), postorbital head (15.7–17.6% vs. 12.9–14.6% SL), snout length (7.2–8.0% vs.
4.5–6.5% SL), eye diameter (8.1–12.4% vs. 5.9–7.0% SL), anterior (1.4–1.8% vs. 0.7–0.8% SL) and posterior
nostril (2.2–3.0% vs. 1.1–1.3% SL) widths, upper-jaw (14.4–16.6% vs. 11.4–12.6% SL) and lower-jaw
(13.9–15.7% vs. 10.7–11.5% SL) lengths, prepectoral (30.7–35.5% vs. 23.7–26.3% SL) and prepelvic
(30.2–33.8% vs. 25.8–27.1% SL) lengths, pectoral (17.3–20.6% vs. 9.7–12.3% SL) and pelvic-fin lengths
(14.5–20.3% vs. 5.0–6.8% SL), and a deeper head (at isthmus 19.7–21.6% vs. 17.4–17.8% SL) and body (at vent
37.6–41.2% vs. 30.1–34.2% SL), than adults (exceeding 313 mm SL).

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Coloration. When fresh (holotype): Almost uniformly dark chocolate brown, eyes bluish; similar in
preservative with pale eyes. In preservative (paratypes): similar to holotype but with extensive irregular paler
patches on sides, particularly on the head (scale pockets demarcated when scales removed); orbital rim,
branchiostegal membrane, lips, and nostrils often noticeably darker than rest of head; eye mostly pale, with dark
outer margin coincident with rim of orbit.
Etymology. Named in honour of New Zealand ichthyologist, Andrew Stewart, whose efforts in building a
substantial collection of stromateoid fishes from the region has contributed so significantly to our understanding of
the life histories and composition of this poorly known group of fishes in the Southern Hemisphere.
Size. Reaches at least 561 mm SL; smallest available specimen 53 mm SL.
Distribution. Confirmed from off Australia and New Zealand in the Southern Hemisphere (Fig. 7) with most
material collected from deep offshore plateaus and seamounts. Possibly more widespread in the southern Indian
Ocean but this needs confirmation. Juveniles (61–99 mm SL) collected from the epipelagic zone at 30–50 m depth;
larger individuals (110–561 mm SL) occur much deeper in the mesopelagic zone at 525–1438 m depth.
Comparisons. Tubbia stewarti differs subtly from T. tasmanica in morphology and meristics and the two
species have been distinguished by their muscle protein characteristics as well as barcoding using the CO1 gene
(Fig. 8). Adult T. stewarti have a relatively broader head with a flatter, less well arched interorbit (width 7.8–8.9%
vs. 6.4–7.9% SL in specimens exceeding 300 mm SL), larger nostrils (width of posterior nostril 5–7 vs. 12–17
times in postorbital head length in juveniles less than 110 mm SL, 10–12 vs. 14–26 times in adults exceeding 300
mm SL), larger eye (diameter 5.9–7.0% vs. 5.2–6.3% SL), longer jaws (head length 2.2–2.5 vs. 2.5–2.8 times
length of lower jaw), narrower caudal peduncle (width at insertion of anal fin 2.4–3.0% vs. 3.5–4.7% SL in
juveniles less than 110 mm SL, 3.3–5.2% vs. 5.3–6.1% SL in adults exceeding 300 mm SL), and more vertebral
centra (44–45 vs. 40–43). Tubbia stewarti appears to reach a slightly larger size than T. tasmanica (known to reach
561 mm SL vs. 407 mm SL) and is typically more elongate as an adult (depth at vent 29.4–34.2% vs. 32.3–38.1%
SL). The eye of T. stewarti is also closer to the dorsal surface of the head (closest horizontal distance from eye to
head margin 1.7–4.4 times vs. 1.0–1.6 times eye diameter in juveniles, 2.0–3.2 times vs. 1.5–1.9 times eye diameter
in adults of T. tasmanica). Head pores are much more densely distributed in T. stewarti than T. tasmanica and this
state is most pronounced around the eye and on the lower jaw (sees Fig. 5). Tubbia stewarti has typically more
dorsal-fin elements but a similar anal-fin count (Table 3).
TABLE 3. Comparison of primary meristic data for Tubbia stewarti sp. nov. and T. tasmanica: A, dorsal-fin; B, anal-fin; and
C, total vertebrae counts.
Remarks. Published data for T. tasmanica also inadvertently contains data for T. stewarti. A 57 (now 53) mm
SL specimen (NMNZ P.007612) figured by McDowall (1979, p 737, Fig. 4) as T. tasmanica was re-examined in
this study and found to be a juvenile of T. stewarti. McDowall’s vertebral count and selected morphometrics for
this specimen are consistent with the diagnosis of T. stewarti. Also, a poorly preserved specimen (RUSI 7423)
A. Dorsal-fin elements
46 47 48 49 50 51 52 53 N
Tubbia stewarti 658522 28
Tubbia tasmanica 3 3985 28
B. Anal-fin elements
32 33 34 35 36 37 38 N
Tubbia stewarti 1 295432 26
Tubbia tasmanica 1 4871 21
C. Vertebral counts
40 41 42 43 44 45 N
Tubbia stewarti 13 15 28
Tubbia tasmanica 1 3 12 12 28

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reported as T. tasmanica in the same paper from off Natal (western Indian Ocean) but not examined in this study,
has a documented vertebral count typical of T. stewarti (i.e. of 44). This specimen might constitute the first record
of the species from the Indian Ocean.
The natural histories of Tubbia species are not well understood. The young appear to live mainly in the
epipelagic zone, probably in association with jellyfish. A collection of 11 juvenile Tubbia tasmanica (CSIRO H
5994–01, 35.6–85.6 mm SL), obtained from the gut content of a shortsnout lancetfish (Alepisaurus brevirostris
Gibbs) caught in the open ocean off southwestern Tasmania in December 1992, suggest these fishes may be locally
abundant at some periods. Adult fish, consistent with many other centrolophids, appear to descend to the meso- and
bathypelagic zones where they have been caught at almost 1500 m depth. Tubbia are oily compared to most other
Australian fishes, and also extremely unusual in having high levels in the flesh of squalene, a liver oil present in
deepsea sharks (Nichols et al, 2001). While these oils may play a similar role in buoyancy control, their exact role
is unknown.
The chisel-edged jaws of Tubbia species are lined with fine teeth adapted to firstly grasping and then slicing
through the soft and flexible tissues of pelagic invertebrates, such as coelenterates and ctenophores. The lower jaw,
which internally overlaps the upper jaw, probably uses a scissor-like action to cleanly remove lumps of tissue. This
feeding action is likely to be possessed by other centrolophid taxa. Tubbia stewarti has larger maximum size, and
larger mouth and gill rakers than T. tasmanica, and probably feeds on larger prey species.
Tubbia tasmanica Whitley
Tasmanian Rudderfish
Figs 1B, 2B, 4, 5B, 6; Tables 1–3
Tubbia tasmanica Whitley, 1943: 179, Fig. 9. Holotype: AMS IB.1148, off Tasmania, Australia.
Schedophilus huttoni (not Waite): Haedrich, 1967, 62 (misidentification).
Holotype. AMS IB 1148, 73 mm SL, off Maria Island, Tasmania, 42º42′ S, 148º34′ E, 50 m, 22 Jan 1941.
Other material. 52 specimens. AMS I 45431–001, 276 mm SL, Pedra Branca seamount, Tasmania, 44º56′ S,
147º16′ E, 525–650 m, 28 Nov 1992; AMS I 45441–002, 350 mm SL, south Tasman Sea, Tasman Sea, 40º58′ S,
164º24′ E, 590–763 m, 19 Jun 2009; CSIRO H 495, 390 mm SL, South Tasman Rise, Tasmania, 46º58′ S, 148º09′
E, 1020–1070 m, 2 Apr 1986; CSIRO H 1600–03, 354 mm SL, South Tasman Rise, Tasmania, 47º40′ S, 148º02′ E,
1200 m, 22 Mar 1986; CSIRO H 1603–01, 276 mm SL, South Tasman Rise, Tasmania, 47º05′ S, 148º27′ E,
1060–1090 m, 2 Apr 1986; CSIRO H 1615–02, 364 mm SL, off Cape Sorell, Tasmania, 42º23′ S, 144º38′ E,
1380–1400 m, 18 May 1986; CSIRO H 2667–01, 311 mm SL, off Studland Bay, Tasmania, 40º49′ S, 143º37′ E,
845–890 m, 8 Mar 1989; CSIRO H 2667–02, 323 mm SL, off Studland Bay, Tasmania, 40º49′ S, 143º37′ E,
845–890 m, 8 Mar 1989; CSIRO H 2812–03, 325 mm SL, South Tasman Rise, Tasman Sea, 47º34′ S, 148º38′ E,
1115–1150 m, 16 Jan 1992; CSIRO H 2815–01, 277 mm SL, South Tasman Rise, Tasman Sea, 47º23′ S, 148º52′ E,
1110–1180 m, 1 Feb 1992 CSIRO H 2830–01, 288 mm SL, South Tasman Rise, Tasman Sea, 47º23′ S, 148º50′ E,
1120–1145 m, 3 Feb 1992; CSIRO H 2831–01, 269 mm SL, South Tasman Rise, Tasman Sea, 47º37′ S, 148º13′ E,
1155–1164 m, 11 Feb 1992; CSIRO H 3272–01, 268 mm SL, off Maria Island, Tasmania, 42º45′ S, 148º16′ E,
85–96 m, 5 Jun 1992; CSIRO H 3273–01, 265 mm SL, off Tasmania, 43º19′ S, 139º40′ E, 16 Jul 1992; CSIRO H
3274–01, 269 mm SL, off Tasmania, 44º42′ S, 142º27′ E, 14 Dec 1992; CSIRO H 3563–01, 321 mm SL, Pedra
Branca seamount, Tasmania, 44º46′ S, 147º08′ E, 660–785 m, 28 Nov 1992; CSIRO H 3563–02, 302 mm SL,
Pedra Branca seamount, Tasmania, 44º46′ S, 147º08′ E, 660–785 m, 28 Nov 1992; CSIRO H 3584–01, 231 mm
SL, Tasman Plateau, off Tasmania, 45º00′ S, 142º30′ E, 20 Nov 1993; CSIRO H 3585–01, 209 mm SL, Tasman
Plateau, off Tasmania, 45º04′ S, 142º44′ E, 20 Nov 1993; CSIRO H 3967–01, 392 mm SL, Pedra Branca seamount,
Tasmania, 44º12′ S, 147º18 E, 650–775 m, 12 Nov 1992; CSIRO H 3967–03, 392 mm SL, Pedra Branca seamount,
Tasmania, 44º12′ S, 147º18′ E, 650–775 m, 12 Nov 1992; CSIRO H 3967–04, 362 mm SL, Pedra Branca
seamount, Tasmania, 44º12′ S, 147º18′ E, 650–775 m, 12 Nov 1992; CSIRO H 3968–02, 402 mm SL, Pedra
Branca seamount, Tasmania, 44º20′ S, 147º22 E, 705–725 m, 24 Nov 1992; CSIRO H 4157–03, 332 mm SL, Pedra
Branca seamount, Tasmania, 44º11′ S, 147º13′ E, 715–900 m, 13 Nov 1992; CSIRO H 6853–12, 306 mm SL,
south Tasman Sea, Tasman Sea, 40º31′ S, 158º16′ E, 591–768m, 18 Jun 2008; CSIRO H 6969–04, 292 mm SL,
south Tasman Sea, Tasman Sea, 40º57′ S, 164º25′ E, 763–912 m, 19 Jun 2009; CSIRO H 6978–05, 363 mm SL,

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south Tasman Sea, Tasman Sea, 40º58′ S, 164º24′ E, 590–763 m, 19 Jun 2009; CSIRO H 6978–06, 351 mm SL,
south Tasman Sea, Tasman Sea, 40º58′ S, 164º24′ E, 590–763 m, 19 Jun 2009; CSIRO H 6979–02, 343 mm SL,
south Tasman Sea, Tasman Sea, 40º55′ S, 166º53′ E, 610–800 m, 18 Jun 2009; CSIRO H 6979–03, 314 mm SL,
south Tasman Sea, Tasman Sea, 40º55′ S, 166º53′ E, 610–800 m, 18 Jun 2009; CSIRO H 6979–04, 324 mm SL,
south Tasman Sea, Tasman Sea, 40º55′ S, 166º53′ E, 610–800 m, 18 Jun 2009; CSIRO H 6979–05, 323 mm SL,
south Tasman Sea, Tasman Sea, 40º55′ S, 166º53 E, 610–800 m, 18 Jun 2009; CSIRO H 7130–01, 282 mm SL,
Pedra Branca seamount, Tasmania, 44º11′ S, 147º21′ E, 525–650 m, 13 Nov 1992; CSIRO T 33, 378 mm SL, off
King Island, Tasmania, Oct 1983; CSIRO T 45, 355 mm SL, off King Island, Tasmania, 40º26′ S, 143º19′ E,
920–940 m, 12 Oct 1983; CSIRO T 1865–01, 378 mm SL, off Trial Harbour, Tasmania, 41º53′ S, 144º29′ E,
923–977 m, 5 Jul 1983; CSIRO T 1892–01, 222 mm SL, South Tasman Rise, Tasmania, 47º10′ S, 147º52 E,
15–200 m, 2 Jan 1980; MNHN 1998–0465, 407 mm SL, St Paul and Amsterdam Islands, Indian Ocean, 38º 06′ S,
77º 45′ E, 754–809 m, 7 Jul 1996; NMNZ P. 007613, 110 mm SL, Puysegur Bank, New Zealand, 46º 21′ S, 165º
31′ E, 167 m, 16 Jan 1976; NMNZ P. 014554, 365 mm SL, Challenger Plateau, New Zealand, 40º 11′ S, 168º 50′ E,
855–857 m, 28 Aug 1983; NMNZ P. 014734, 403 mm SL, Challenger Plateau, New Zealand, 40º 18′ S, 169º 08′ E,
862–864 m, 18 Oct 1983; NMNZ P. 015814, 386 mm SL, off Canterbury Bight, New Zealand, 44º 37′ S, 173º 26′
E, 772–790 m, 11 Jun 1984; NMNZ P. 015932, 310 mm SL, Challenger Plateau, New Zealand, 40º 00′ S, 167º 42′
E, 1064–1082 m, 8 Jul 1984; NMNZ P. 018859, 84 mm SL, off Dunedin, New Zealand, 45º 59′ S, 170º 33′ E, 48 m,
15 Jan 1982; NMNZ P. 025990, 172 mm SL, Campbell Plateau, New Zealand, 51º 43′ S, 174º 06′ E, 779–789 m,
17 Aug 1990; NMNZ P. 027112, 194 mm SL, Chatham Rise, New Zealand, 44º 23′ S, 179º 40′ W, 910–1000 m, 10
Nov 1990; NMNZ P. 027973, 334 mm SL, Campbell Plateau, New Zealand, 48º 32′ S, 169º 11′ E, 699–712 m, 18
Dec 1991; NMNZ P. 029664, 302–357 mm SL, Snares Shelf, New Zealand, 48º 51′ S, 166º 26′ E, 540–560 m, 23
Oct 1992; NMNZ P. 036059, 143 mm SL, Puysegur Trench, New Zealand, 45º 44′ S, 165º 33′ E, 100 m, 12 May
1998; NMNZ P. 041938, 69 mm SL, off Timaru, New Zealand, 44º 16′ S, 171º 30′ E, 20–23 m, 5 Jun 1999; NMV
A 26359–001, 269 mm SL, Pedra Branca seamount, Tasmania, 44º56′ S, 147º16′ E, 650–525 m, 28 Nov 1992;
NMV A 26360–001, 323 mm SL, south Tasman Sea, Tasman Sea, 40º58′ S, 164º24′ E, 763–590 m, 19 Jun 2009.
Diagnosis. A species of Tubbia with the following combination of characters: head of adult moderately
compressed with a strongly arched interorbit (its width 6.4–7.9% SL in specimens exceeding 300 mm SL); nostrils
small, width of posterior nostril 14–26 times in postorbital head length in adults (exceeding 300 mm SL); eye small
in adults (diameter 5.2–6.3% SL) and well removed from dorsal margin of head (closest horizontal distance
1.5–1.9 times in diameter of orbit); jaws relatively small, head length 2.5–2.8 times length of lower jaw; head pores
dense; caudal peduncle relatively broad at insertion of anal fin, width 3.5–4.7% SL in juveniles (less than 110 mm
SL), 5.3–6.1% SL in adults (exceeding 300 mm SL); vertebral centra 40–43.
Size. Reaches at least 407 mm SL based on material form this study; recorded to 371 mm SL by McDowall
(1979).
Distribution. Widely distributed in the temperate mesopelagic zone of the Southern Hemisphere, off
Australia, New Zealand, and St Paul Island (southwestern Indian Ocean), but probably more widespread; occurs
over continental slopes as well as offshore seamounts and plateaus (Fig. 7) . A record of T. tasmanica from Natal
(southwestern Indian Ocean) may be of T. stewarti and warrants further investigation. Most juveniles (68–268 mm
SL) were taken in the epipelagic zone at 15–200 m; larger individuals (172–402 mm SL) for which depth data exist
were caught by midwater and demersal trawl gear at 52–1400 m.
Remarks. The two species of Tubbia are very similar to each other and it is not surprising that past authors
failed to identify a second species. Features used in the literature to define T. tasmanica equally define T. stewarti,
and no apomorphic character (with the possible exception of vertebral counts) has been identified to diagnose
either species. As a consequence, T. tasmanica was not redescribed herein. Rather, a composite of characters
distinguishing the two species are given in the diagnoses and these were discussed in the Comparisons section of T.
stewarti. In summary, T. tasmanica has a more compressed head, less slender body, more convex interorbit, shorter
jaws, fewer dense head pores, relatively smaller eyes and nostrils, a broader caudal peduncle and fewer vertebral
centra, than T. stewarti.
There are also important molecular differences between the two species (see Fig. 8). A neighbour joining tree,
including Tubbia species and their close relatives, identified individuals into their six respective taxa with good
bootstrap support (>70%): i.e. Tubbia tasmanica, T. stewarti, Schedophilus huttoni, S. maculatus, S. velaini and S.
labyrinthicus. Intragroup variation within five of the groups was low and Bootstrap values exceeded 85%

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supporting morphological evidence that these groups represent single species. Some variability was found in the S.
labyrinthicus group which comprises two genetically distinct sympatric forms in New Zealand waters.
FIGURE 4. Lateral views of juveniles of Tubbia tasmanica: A, holotype AMS IB 1148, 72.5 mm TL (preserved); B, NMNZ P.
018859, 83.7 mm TL (preserved); C, NMNZ P. 007613, 110 mm TL (preserved).

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FIGURE 5. Ventral view of the lower jaw of adult: A, Tubbia stewarti sp. nov., paratype, NMNZ P. 025954, 367 mm SL
(preserved); B, T. tasmanica, CSIRO H 3967–03, 392 mm TL (preserved).
Most importantly, the molecular analysis distinguished the two species of Tubbia supporting findings from an
earlier muscle protein analysis (Daley et al., 1997). Our analysis is not fully reliable from a phylogenetic aspect as
it uses only a single molecular marker, but the clustering of S. huttoni with Tubbia away from the 3–4 other ‘firm-
bodied’ Schedophilus species, supports earlier suggestions that the genus Schedophilus is polyphyletic (McDowall,
1979; Bolch et al., 1994). Also, conspecificity of Schedophilus velaini and S. labyrinthicus has been proposed
(Andrew et al., 1995; Eschmeyer & Fricke, 2012) but these taxa appear to be distinct at the species level.

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FIGURE 6. Skin peeled from the trunk of Tubbia tasmanica (NMNZ P. 044416) to reveal its inner surface (left section)
showing distribution and density of pores and the corresponding underlying body surface (right section) with its fibrous
attachments.
FIGURE 7. Map of the Australasian region showing the distributions of Tubbia stewarti (encircled diamonds) and T.
tasmanica (triangles); holotypes of Tubbia stewarti (encircled cross) and T. tasmanica (encircled star).
Key to species of the genus Tubbia
1. Head of adult robust with a relatively broad and moderately arched interorbit; nostrils large, width of posterior nostril 10–12
times in postorbital head length in adults (exceeding 300 mm SL); eye large in adults and close to dorsal margin of head (clos-
est horizontal distance 2.0–3.2 times in diameter of orbit); jaws large, head length 2.2–2.5 times length of lower jaw; caudal
peduncle narrow at insertion of anal fin, width 2.4–3.0% SL in juveniles (less than 110 mm SL), 3.3–5.2% SL in adults
(exceeding 300 mm SL); vertebral centra 44–45 . . . . . . . . . . . . . . . . . . .Tubbia stewarti sp. nov. (Australia and New Zealand)
- Head of adult moderately compressed with a strongly arched; nostrils small, width of posterior nostril 14–26 times in postor-
bital head length in adults (exceeding 300 mm SL); eye small in adults and well removed from dorsal margin of head (closest
horizontal distance 1.5–1.9 times in diameter of orbit); jaws small, head length 2.5–2.8 times length of lower jaw; caudal
peduncle broad at insertion of anal fin, width 3.5–4.7% SL in juveniles (less than 110 mm SL), 5.3–6.1% SL in adults (exceed-
ing 300 mm SL); vertebral centra 40–43 . . . . . Tubbia tasmanica Whitley, 1943 (Australia, New Zealand, and St Paul Island)

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FIGURE 8. Neighbour-joining tree of nucleotide sequence divergence at the barcoding region of the COI gene for species of
Tubbia and Schedophilus (Centrolophidae) from the Southern Hemisphere. Scale bar represents 1% K2P following 10000
bootstraps. Tissue sample numbers relate to those present in the Barcode of Life Database (www.barcodinglife.org).
Acknowledgments
This research was facilitated by a grant from the New Zealand Foundation for Research, Science and Technology
(Biosystematics of NZ EEZ Fishes programme, FRST contract MNZX0203; project leader Dr C. Roberts), as well
as the CSIRO Wealth from Oceans Flagship. Collection staff at various museums, in particular Patrice Provost
(NMHN), Mark McGrouther (AMS), and Andrew Stewart and Clive Roberts (NMNZ), kindly made valuable loan
material available for this study; Carl Struthers (NMNZ) produced X-rays of NMNZ material and figures 1A, 3C,
4B & C and 6, and Jeremy Barker (NMNZ) created the map. Alastair Graham (ANFC) assisted with materials
sections and John Pogonoski (ANFC) compiled meristic details from CSIRO radiographs. Sharon Appleyard
(CSIRO) provided invaluable assistance with the molecular analysis. Carlie Devine (ANFC) prepared digital
images of material used in the figures. The authors wish to thank the journal editor and reviewers of the manuscript
for their useful comments.

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