Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The
nean Society of London, 2004? 2004
141?
179205
Original Article
Lin-
R. D. MOOI and A. C. GILLPHYLOGENETIC POSITION OF NOTOGRAPTIDAE
Zoological Journal of the Linnean Society, 2004, 141, 179–205. With 14 figures
Notograptidae, sister to Acanthoplesiops Regan
(Teleostei: Plesiopidae: Acanthoclininae), with comments
on biogeography, diet and morphological convergence
with Congrogadinae (Teleostei: Pseudochromidae)
RANDALL D. MOOI1* and ANTHONY C. GILL2†
1
Milwaukee Public Museum, 800 West Wells Street, Milwaukee, Wisconsin 53233–1478, USA
2
Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK
Received February 2003; accepted for publication January 2004
The Notograptidae contains one genus, Notograptus Günther, and five nominal species from northern Australia and
southern New Guinea. Morphological evidence places Notograptus among acanthoclinine plesiopids (continuous free
margin of lower lip; head naked; dorsal and anal fins with many spines and few segmented rays; no extensor proprius; reduced number of caudal-fin rays) and supports a sister relationship with Acanthoplesiops (symphyseal flap
on lower lip; reduced hypural 5; reduced hypurapophysis). This hypothesis resolves the relationships within Acanthoplesiops, clarifying the polarity of autogenous middle radials of dorsal- and anal-fin pterygiophores. The proposed
relationships among acanthoclinines are: Acanthoclinus (Belonepterygion (Beliops (Notograptus (Acanthoplesiops
hiatti (A. indicus (A. psilogaster (A. echinatus))))))). The distribution of Notograptus compliments that of its proposed
sister clade in that Acanthoplesiops is unknown from northern Australia or southern New Guinea. There are
repeated geographical patterns among several groups suggesting that Australia is a basal area to a broader IndoPacific region. Similarity between the Congrogadinae (Pseudochromidae) and Notograptus has long been noted, both
having a loosely connected suspensorium and elongate body which were mistakenly considered indicators of relationship; we add reduced branchial arches, straight, tube-like gut and highly expandable anus. We examine these
similarities as an indication of a shared specialized feeding habit. Notograptus is an alpheid shrimp predator, able
to swallow its large prey whole. Most species of congrogadines eat whole, large crustaceans. This is probably an
example of convergent adaptation to a particular selective regime. © 2004 The Linnean Society of London,
Zoological Journal of the Linnean Society, 2004, 141, 179–205.
ADDITIONAL KEYWORDS: adaptation – biology – classification – feeding behaviour – myology – osteology –
phylogeny.
‘. . . in acanthomorphs as a whole there are about 280 families.
Many of these families seem to have come down like the tablets, unblemished by analysis since their names were inscribed
on stone in the days of Günther or Gill or Regan’ (Patterson,
1993: 29).
INTRODUCTION
The Notograptidae, bearded eel-blennies or dirkfishes,
comprises small (<200 mm SL), elongate, shallow*Corresponding author. E-mail: mooi@mpm.edu
†Current address: School of Life Sciences, Arizona State
University, Tempe, AZ 85287–4501, USA.
water fishes (Fig. 1). There is one recognized genus,
Notograptus Günther, and five nominal species
restricted to the northern coast of Australia and
southern coast of New Guinea (Gill & Mooi, 1993;
Mooi, 1999). The family is in need of revision.
The phylogenetic position of the family has long
been in question. Günther (1867) originally described
the genus in the Blenniidae, a placement followed by
McCulloch (1918). Regan (1912) erected a (then)
monotypic family Notograptidae for this unusual fish
and since that time, it has predominantly remained
with the Blennioidei (e.g. Jordan, 1923; de Beaufort,
1951; Greenwood et al., 1966; Norman, 1966; see
review by Gill & Mooi, 1993). However, in addition to
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
179
180
R. D. MOOI and A. C. GILL
Figure 1. Notograptus sp., BMNH 2002.1.2.1–2, 100.5 mm SL. Scale bar = 50 mm.
the true blennioids (sensu Springer, 1993), the Blennioidei of these authors variously included taxa now
assigned to the perciform suborders Percoidei, Gobioidei, Trachinoidei and Zoarcoidei, and to the paracanthopterygian order Ophidiiformes.
Greenwood et al. (1966: 401) included the Notograptidae in the Blennioidei, listing without comment the
Stichariidae (classified in the clinid subfamily Ophiclininae by George & Springer, 1980) as a synonym.
Gosline (1968: 60), following the lead of Regan
(1912), suggested a relationship of notograptids to
congrogadids and less certainly to peronedyids [sic],
placing the three taxa in his blennioid superfamily
Congrogadoidae. However, Godkin & Winterbottom
(1985) provided evidence for the inclusion of the Congrogadidae as a subfamily of the percoid Pseudochromidae, and George & Springer (1980) assigned
peronedysids to the blennioid clinid tribe Ophiclinini.
Nelson (1984), without evidence, placed notograptids
among the trachinoids; Mooi & Johnson (1997) provided arguments to dismantle the Trachinoidei, making inclusion of the notograptids in this unnatural
‘group’ uninformative.
Gill & Mooi (1993) listed apomorphic features of
Notograptus and considered its phylogenetic position. We noted that it shared numerous features
with other elongate perciforms (in particular, elongate blennioids, zoarcoids, pholidichthyids and congrogadine pseudochromids) but concluded that many
of these features are a consequence of elongation
(e.g. numerous vertebrae and dorsal- and anal-fin
rays; highly fused caudal skeleton; reduced pelvic
fins and girdle) and thus, not necessarily indicative
of close relationship. Considering characters that are
not obviously associated with elongation (e.g. egg
morphology; dorsal- and anal-fin spine-bearing pterygiophore construction; dorsal and anal fins comprising almost entirely spinous rays), we concluded that
available evidence best supported a relationship with
acanthoclinine plesiopids, a proposal first made by
Smith-Vaniz & Johnson (1990). However, we elected
not to place Notograptus in the Acanthoclininae,
pending the discovery of additional corroborating
evidence.
The purposes of this paper are to test the possibility
of an acanthoclinine relationship by including
Notograptus in a parsimony analysis that combines
characters and taxa employed by Smith-Vaniz &
Johnson (1990) in their examination of relationships
within the Acanthoclininae, and by Mooi (1993) in his
analysis of plesiopid monophyly and relationships
among nonacanthoclinine plesiopid genera. We also
examine the biogeographical implication of this formalization of our hypothesis that Notograptus has its
closest relatives among acanthoclinine plesiopids.
Lastly, we suggest that the convergence in morphology
of Notograptus and congrogadine pseudochromids is
due to a specific selective regime (similar specialized
feeding behaviour) and provides an example of adaptation in the historical sense of Coddington (1988) and
Larson & Losos (1996).
MATERIAL AND METHODS
External and myological characters were scored by
examination of alcohol-preserved specimens. Osteological characters were examined using cleared and
stained and X-radiographed material. Character
states of nonacanthoclinine plesiopids were obtained
from the literature (Smith-Vaniz & Johnson, 1990;
Mooi, 1993). Character states of acanthoclinine plesiopids were obtained from specimens where available
and taken from the literature when not (Smith-Vaniz
& Johnson, 1990). Acanthoplesiops naka Mooi & Gill
from Tonga (USNM 327794), known only from one
small ethanol specimen (9.9 mm SL), was excluded
from the phylogenetic analysis as it would not impact
the placement of Notograptus and, having only external characters available for analysis, would introduce
a series of unknowns to the data table.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
Character analysis was performed using PAUP*
Version 4.0 beta10 (Swofford, 2001) and results
explored using MacClade 4.05 (Maddison & Maddison,
2001). An initial branch-and-bound search was undertaken, with all characters unordered excepting characters 3, 10, 11, 19, 38 and 40, following previously
published interpretations. Equally parsimonious trees
were combined using strict consensus methods. The
data matrix (Table 1) differentiates those characters
for which taxa had no observations and were unknown
(?) from those that were inapplicable (n). Two analyses
were performed to deal with inapplicable characters.
One used the suggestion of Maddison (1993) that
inapplicable states be included as part of an unordered multistate character (composite coding as
described in Strong & Lipscomb, 1999). The other used
reductive coding where taxa for which a character is
inapplicable are coded the same as if the state were
unknown (using ‘?’) (recommended by Strong & Lipscomb, 1999). The possible effect of these codings on
the position of Notograptus was also examined by
deleting the characters inapplicable to Notograptus
and repeating the analysis. Additionally, the analysis
was repeated with the removal of all inapplicable and
unknown states in the matrix by deleting all inapplicable characters (1, 2, 15, 33, 41, 43, 49, 59), deletion
of the two taxa with most unexamined (unknown)
states (Beliops batanensis Smith-Vaniz & Johnson and
181
Acanthoclinus psilogaster Hardy), along with elimination of three remaining characters (10, 13 and 24) that
still exhibited unknown values in a few taxa. All analyses were repeated with all characters unordered to
examine any effects on topology. Finally, the tree was
constrained to have Notograptus as sister to the Plesiopidae and effects on tree length noted. Tree statistics reported are length (number of steps), consistency
index (CI), rescaled consistency index (RC) and retention index (RI) (Farris, 1989).
Distributional records were taken from the literature, examined specimens and museum catalogue
records, the latter focusing on the collections of
BMNH, MPM, USNM and AMS.
Gut contents were examined chiefly using Xradiographs (X-ray) and cleared and stained (CS)
specimens as noted above. Occasionally, ethanolpreserved (EtOH) specimens were dissected.
Institutional abbreviations follow Leviton et al.
(1985).
NOTOGRAPTUS
(16 lots, 99 specimens: 44–178 mm SL): AMNH
216034, 100.5 mm, North West Cape, Western Australia, 3 April 1969; ANSP 109653, 11: 63–105 mm
(EtOH, X-ray), 4: 70–90 mm (CS), Little Hope Is.,
Queensland, 17 January 1969; ANSP 109654, 38 mm
Table 1. Data matrix for examining the phylogenetic position of Notograptus among the Plesiopidae. ? – not examined;
n – not applicable due to modification
Character number
1 11111 11112 22222 22223 33333 33334 44444 44445 55555 5555
12345 67890 12345 67890 12345 67890 12345 67890 12345 67890 12345 6789
Outgroup
Trachinops
Assessor
Paraplesiops
Calloplesiops
Steeneichthys
Fraudella
Plesiops
Acanthoclinus fuscus
A. littoreus
A. rua
A. marilynae
A. matti
Belonepterygion
Beliops xanthokrossos
Beliops batanensis
Notograptus
Acanthoplesiops indicus
A. hiatti
A. psilogaster
A. echinatus
00000
11111
11111
11111
11111
11111
11111
11111
11221
11221
112?1
112?1
11221
11221
112?1
??2?1
nn221
11221
11221
??2??
??221
00000
11100
11111
11111
11111
11112
11111
11112
11112
11112
11112
1111?
11112
11112
1101?
?10??
00011
1101?
1101?
?10??
01012
00000
00000
10000
11111
11110
21111
11111
11111
1111n
1111n
1111n
1111n
1111n
1111n
1111n
1???n
2101n
1111n
1111n
11??n
11?1n
00000
00000
00000
11000
01111
11111
00000
10000
10110
10110
10110
10110
10110
10110
10120
00120
00130
10120
10120
1?12?
10120
00000
00000
00000
00000
00000
00001
11100
11111
11111
11111
111?1
111?1
11111
11111
001?1
?????
00001
001?0
001?0
?????
00110
00000
10000
00000
00000
00000
01000
00000
11000
10111
10111
10111
10111
10111
10111
00111
10111
10111
10111
10111
??111
10111
00000
00001
00001
00000
00000
33n01
00000
00000
11100
11100
11100
11100
11100
10011
00010
00011
22011
23n11
23n11
23n11
23n11
00000
00000
00000
00000
00000
01200
00000
00000
01000
00000
00100
00000
00000
11100
11111
11111
11102
11111
11111
01211
01211
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
11111
11111
n1n00
11000
11000
11000
11000
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
00000
00000
00000
00000
00000
00100
00000
00000
10000
10000
00000
00000
00000
00000
11000
11000
111n1
00111
01111
00111
00111
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
00000
10100
11111
11111
11111
11111
0000
0000
0000
0000
0000
0000
0000
0000
0010
0010
0000
0000
0000
0001
0000
0000
001n
1101
1101
1101
1101
182
R. D. MOOI and A. C. GILL
(EtOH, X-ray), Little Hope Is., Queensland, 26 January 1969; ANSP 165418, 98 mm (EtOH, X-ray), Cape
Arnhem, Northern Australia, 14 July 1948; BMNH
1867.5.13.16 + 1867.6.6.42, 2: 114–153.5 mm, Cape
York, Queensland (syntypes of N. guttatus Günther,
1867); BMNH 2002.1.19.16, 128.5 mm (CS), Clarence Strait, East Vernon Is., Northern Territory;
BMNH 2002.1.2.1–2, 2: 65–99 mm (EtOH, X-ray),
Burnside Is., Western Australia, 19 May 1996; MPM
32586, 29: 44–112 mm (EtOH, X-ray), Locker Is.,
Western Australia, 16 May 1996; ROM 38369, 8: 58–
101 mm, Wonga Beach just south of Daintree R.,
Queensland, 25 September 1981; ROM 717CS,
82.2 mm (CS), Wonga Beach just south of Daintree
R., Queensland, 25 September 1981; USNM 173796,
3: 77–159 mm (EtOH, X-ray), Groote Eylandt, Northern Territory, 19–25 April 1948; USNM 173797, 6:
45–178 mm (EtOH, X-ray), Groote Eylandt, Northern
Territory, 7 June 1948; USNM 173798, 11: 81–
170 mm (EtOH, X-ray), Cape Arnhem, Northern Territory, 14 July 1948; USNM 222134, 4: 87–120 mm
(EtOH, X-ray), 31 May 1979, Clarence Strait, East
Vernon Is., Northern Territory; USNM 325199, 14:
68–101 mm (EtOH, X-ray), Darwin, Northern Territory, 18–19 February 1988.
OTHER PLESIOPIDAE (ALL REMAINING MATERIALS
X-RADIOGRAPHS UNLESS OTHERWISE NOTED)
Acanthoclinus fuscus Jenyns (5 lots, 68 specimens: 28–
220 mm SL): ANSP 165085, 2: 46–47 mm (CS); MPM
32616, 3: 160–220 (EtOH); USNM 200547, 4: 55–
84 mm; USNM 200548, 6: 53–95 mm; USNM 339246,
53: 28–99 mm.
A. littoreus (Forster) (2 lots, 7 specimens: 56–
119 mm): ANSP 165089, 2: 56–78 mm (CS); USNM
339230, 5: 98–119 mm.
A. marilynae (Hardy): ANSP 134947, 2: 90–95 mm
(CS).
A. matti (Hardy): ANSP 165088, 52 mm (EtOH, suspensorium CS, X-ray).
A. rua (Hardy): ANSP 165087, 46 mm (CS, gutted).
Acanthoplesiops echinatus Smith-Vaniz & Johnson:
ANSP 166316, 21 mm (EtOH, gill arches and suspensorium CS, X-ray).
A. hiatti Schultz (3 lots, 10 specimens: 15–20 mm
SL): ANSP 165421, 20 mm (CS, gutted); USNM
135783, 2: 15–16 mm; USNM 257874, 7: 15–20 mm.
A. indicus (Day) (5 lots, 5 specimens: 19–27 mm SL):
ANSP 122483, 27 mm (CS); ANSP 165570, 22 mm
(CS, gutted); BMNH 1889.8.17.5, 19 mm (holotype);
RUSI 17291, 22 mm (CS); RUSI 17293, 22 mm
(EtOH).
A. psilogaster Hardy (3 lots, 4 specimens: 12–22 mm
SL): USNM 288813, 12 mm; USNM 318027, 22 mm;
USNM 326763, 2 : 15–16 mm.
A. naka Mooi & Gill: USNM 327794, 9.9 mm (holotype) (EtOH).
Assessor flavissimus Allen & Kuiter: MPM 40273,
36.0 mm (CS).
A. macneilli Whitley: MPM 40268, 39.0 mm (EtOH).
Beliops xanthokrossos Hardy: ANSP 165557,
26 mm (EtOH, X-ray), 26 mm (CS, gutted).
2:
Belonepterygion fasciolatum (Ogilby) (4 lots, 43 specimens: 12–42 mm SL): ANSP 142690, 42 mm (EtOH,
X-ray); BMNH 1914.12.28.1–2, 2: 30–39 mm; MPM
40265, 2: 47–48 mm (EtOH); USNM 257875, 31: 12–
36 mm; USNM 257876, 9: 15–38 mm.
Paraplesiops poweri Ogilby: MPM 40288, 50.0 mm
(EtOH).
Plesiops coeruleolineatus Rüppell: MPM SOL 98–20,
46.5 mm (EtOH).
Trachinops taeniatus Günther: MPM 40291, 48.0 mm
(EtOH).
PSEUDOCHROMIDAE
Anisochromis kenyae Smith: (10 lots, 46 specimens:
13.8–25.6 mm SL): AMS I.28113–064, 2: 13.9–
22.1 mm; ANSP 134469, 25.6 mm; ROM 56501, 2:
22.1–23.0 mm; ROM 56502, 9: 13.8–22.7 mm; ROM
56711, 21.6 mm; RUSI 149, 21.3 mm (holotype); RUSI
854, 14: 17.5–25.0 mm (paratypes); RUSI 4905, 3:
22.3–23.9 mm; RUSI 4906, 7: 21.6–25.0 mm
(23.3 mm, CS); USNM 216415, 6: 21.7–24.5 mm
(paratypes).
A. mascarenensis Gill & Fricke (7 lots, 11 specimens:
13.3–25.5 mm SL): BMNH 2001.3.8.2, 23.3 mm (paratype, CS); BPBM 16277, 13.3 mm (paratype);
MNHN 2001–494, 24.1 mm (paratype); SMNS 20933,
2: 19.7–25.5 mm (paratypes); SMNS 21025, 4: 19.7–
25.2 mm (partypes); SMNS 23037, 23.9 mm (holotype); USNM 364534, 19.6 mm (paratypes).
A. straussi Springer, Smith & Fraser (9 lots, 82 specimens: 16.1–28.3 mm SL): AMNH 35892, 6: 22.0–
28.0 mm SL (paratypes); BMNH 1976.8.24.1–10, 10:
16.8–25.4 mm SL (paratypes; 21.5 mm CS); CAS
37640, 14: 16.1–24.9 mm (paratypes); USNM 215859,
26: 18.7–26.1 mm (paratypes); USNM 216462,
23.9 mm (holotype); USNM 216463, 19: 16.2–27.0 mm
(paratypes); USNM 216464, 26.8 mm (paratype);
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
USNM 216465, 3: 22.6–23.9 mm (paratypes); USNM
216466, 2: 26.5–28.3 mm (paratypes).
Blennodesmus scapularis Günther (15 lots, 120 specimens: 28.5–87 mm SL): AMS IA. 606–8, 7: 41–
77 mm; AMS I.7072–4, 7076, 8: 45–71.6 mm; AMS
I.17445–151, 13: 47–87 mm. AMS I.20200–029, 19:
34–87 mm; BPBM 17412, 6: 57–87; MPM field no.
RDM96-20, 6: 28.5–60 mm; MPM field no. RDM96-28,
10: 30–73 mm; ROM 43211, 9: 45–85 mm; ROM
43212?, 2: 46.4–48.6 mm; ROM RW93-12, 5: 46.4–
55.6 mm; ROM RW93-16, 6: 42–55 mm; ROM RW9321, 53 mm; USNM 205026, 26: 38–53 mm; WAM
P.25112–014, 74 mm; WAM P.4645, 57.6 mm.
Congrogadus hierichthys Jordan & Richardson (9 lots,
30 specimens: 50–159 mm SL): BMNH 1933.3.11 :
727–728, 2: 101–102 mm; CAS 40138, 125 mm;
CAS SU20208, 108 mm (holotype); CAS SU34113,
159 mm; CAS SU33664, 8 : 74–102 mm; CAS
SU33665, 3 : 100–123 mm; ROM 46818, 12: 50–
68 mm; SOSC SP 1978–17, 70.5 mm; USNM 61684,
92 mm (paratype).
C. malayanus (Weber) (3 lots, 12 specimens: 30.8–
70.0 mm SL): AMS I.20828–018, 2: 38.2–62.5 mm;
QM I.17705, 2: 61–63 mm; ZMA 112.577, 8: 30.8–
70.0 mm (syntypes).
C. spinifer Borodin (11 lots, 34 specimens: 35.5–
121 mm SL): AMS IA.839, 6: 55–70 mm; AMS
IA.4239, 5 : 37–100 mm; AMS IA.4266, 3 : 35.5–
84.3 mm; AMS IA.4675, 81 mm; AMS I.15557–215, 3:
72–130 mm; AMS I.21842–011, 6: 49.2–121 mm; AMS
I.21943, 121 mm; BMNH 1911.1.4 : 3–4, 116 mm;
BMNH 1933.8.14 : 18–21, 5: 57–89 mm; SAMA
F.1494, 2: 44–63.5 mm; WAM P.5520, 83 mm.
C. subducens (Richardson) (25 lots, 75 specimens: 31–
340 mm SL): BPBM 14415, 2: 38–62 mm; CAS SU
7120, 100 mm; CAS SU 33860, 2: 97.5–116; MNHN
6716, 171 mm; MPM 32613, 340 mm (EtOH); MPM
32617, 205 mm (EtOH); MPM field no. RDM96-10,
65 mm; ORI 333144, 1; RMNH 6720, 2: 146–332 mm;
RMNH 6721, 342 mm; ROM 3911, 281 mm; SOSC ref.
no. BBC 1676 A, 2 : 170–191 mm; SOSC ref. no. BBC
1679, 10: 158–210 mm; SOSC SP-78, 71.3; USNM
122460, 102.2 mm; USNM 173804, 10 : 150–302 mm;
USNM 212291, 7: 31–156 mm; USNM 287587, 12: 92–
320 mm; USNM 287588, 10: 227–314 mm; USNM
287589, 3: 107–279 mm; WAM P.10070, 108.5 mm;
WAM P.22454, 31.2 mm; WAM P.25532–005, 75 mm;
WAM P.22670, 101.3 mm; WAM P.31013–025,
183 mm.
C. winterbottomi Gill, Mooi & Hutchins (15 lots, 31
specimens: 61.3–119 mm SL): AMS I.39770–001, 2:
66.9–74.3 mm; BMNH 1999.9.21.1–3, 3: 77.6–
115.4 mm; BMNH 1999.9.21.4–5, 2: 71.5–84.5 mm;
183
CSIRO H5237-01, 92.5 mm; MPM 32574, 5: 61.3–
101.8 mm; NTM S.14970–001, 80.9 mm; QM I.31415,
107.3 mm; ROM 71992, 2: 85.3–107.0 mm; SAMA
F9302, 81.5 mm; USNM 358035, 97.5 mm; WAM
P.31013–046, 2: 66.1–111.5 mm; WAM P.31017–022, 3:
71.8–115.7 mm; WAM P.31018–012, 4: 66.8–
119.0 mm; (all preceding are paratypes); WAM
P.31582–001, 85.1 mm (holotype); WAM P.31582–002,
2: 68.5–80.0 mm (paratypes).
Halidesmus polytretus Winterbottom (2 lots, 2 specimens: 57.0–57.3 mm SL): LACM 30695–13, 57.3 mm
(holotype); LACM 30695–29, 57.0 mm (paratype).
H. scapularis Günther: RUSI
97.5 mm.
11057,
20:
45.5–
H. socotraensis Gill & Zajonz (4 lots, 6 specimens:
39.6–69.5 mm SL): BMNH 2002.1.19.3, 63.3 mm SL
(paratype); ROM 72697, 60.0 mm (paratype); SMF
29223, 64.3 mm (holotype); SMF 29293, 3: 39.6–
69.5 mm (paratype).
H. thomaseni (Nielsen) (4 lots, 78 specimens: 29–
134 mm SL): USNM SOSC ref. 4, 40: 30–82 mm;
USNM SOSC acc. no. 23, 30: 29–90 mm; USNM
Cruise 4–8, 5: 66–75 mm; ZMUC P.75396–98, 3: 60–
134 mm (paratypes).
Halimuraena hexagonata Smith (5 lots, 28 specimens:
22.5–60 mm SL): RUSI 863, 10: 45–60 mm; RUSI
4019, 4: 32.5–45 mm; RUSI 5326, 4: 22.4–33.5 mm;
RUSI 5402, 5: 28.2–31.9 mm; USNM SOSC ref. no.
145, 5: 22.5–51.0 mm.
H. shakai Winterbottom (16 lots, 57 specimens: 22–
55 mm): BPBM 21709, 48 mm; ROM 56788, 36 mm;
ROM 56789, 2: 25–44 mm; ROM 56790, 11: 22–
47 mm; ROM 76–10, 7: 42–50 mm; ROM 76–11,
54.8 mm; ROM 76–12, 3: 42–50; ROM 76–15, 9: 35–
52 mm; ROM 76–24, 3: 42–46 mm; RUSI 8955, 2:
43–48 mm; RUSI 8994, 2: 37–52 mm; RUSI 9199, 3:
46–52 mm; RUSI 9333, 3: 38–48 mm; RUSI 9415, 2:
44–54 mm; RUSI 9451, 3: 45–47 mm; RUSI 9800, 4:
35–55 mm.
Halimuraenoides isostigma Maugé & Bardach (3
lots, 18 specimens: 65–242 mm): MNHN 1985–240,
242 mm (holotype); MNHN 1985–241, 15: 65–278 mm
(paratypes); ROM 46134, 2: 78–117 mm (paratypes).
Haliophis aethiopus Winterbottom (2 lots, 2 specimens: 49–50 mm SL): BPBM 20920, 49 mm (holotype); ROM 38419, 50 mm (paratype).
H. guttatus (Forsskål) (38 lots, 308 specimens: 20.7–
132 mm SL): BMNH 1951.1.16 : 606–608, 2: 48.6–
66 mm; BMNH 1960.3.15 : 1591–1603, 13: 44–
100 mm; BPBM 18361, 4: 65–132 mm; BPBM 19883,
4: 36–88 mm; BPBM 22600, 3: 34–50; LACM 30859–
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
184
R. D. MOOI and A. C. GILL
16, 14: 30–50 mm; ROM 38245, 102 mm; ROM 38346,
9: 20.7–57 mm; ROM X-ray C4, 13: 43.7–93.5 mm;
ROM X-ray C15, 56 mm; ROM X-ray plate no. 3, 3 :
34–80 mm; RUSI 5402, 38.1 mm; RUSI 5414, 4: 70–
78 mm; RUSI 5427, 10: 50–104 mm; RUSI 5858,
28 mm; RUSI uncat., 17: 56–82 mm; SMF 29198, 14:
38–79.5 mm; SMF 29224, 64.8 mm; SOSC acc. no. 54,
12: 63–84 mm; SOSC ref. no. 142, 86 mm; SOSC ref.
no. 145, 10: 50–104 mm; SOSC ref. no. 145, 16: 23–
100 mm; SOSC ref. no. 170, 10: 61–78 mm; SOSC ref.
no. 540, 125.4 mm; SOSC ref. no. 540, 3: 45–58 mm;
SOSC ref. no. 540, 3: 80–91 mm; SOSC ref. no. 540, 4:
80–127 mm; SOSC ref. no. 540, 13: 30–93 mm; TAU
P3744, 52.7; TAU P4911, 4: 39.2–77.5 mm; USNM
212290, 35: 24–84 mm; USNM 273294, 24: 29–
110 mm; USNM 279889, 16: 23–100 mm; USNM
285199, 10: 60–75 mm; USNM 308028, 10: 62–98 mm;
USNM uncat., Tulear, 87 mm; USNM uncat., Israel,
15: 35–98 mm; WAM P.4912, 4: 39–77 mm.
Natalichthys leptus Winterbottom (2 lots, 2 specimens:
52–56.4 mm SL): SAM 28938, 56.4 mm (holotype);
SAM 28939, 52.0 mm (paratype).
N. ori Winterbottom (2 lots, 2 specimens: 54–60.5 mm
SL): SAM 17340, 60.5 mm (holotype); SAM 28993,
54.0 mm (paratype).
N. sam Winterbottom (2 lots, 2 specimens: 40.4–
43.2 mm SL): SAM 21915, 43.2 mm (holotype); SAM
28940, 40.4 mm (paratype).
Rusichthys explicitus Winterbottom (3 lots, 3 specimens: 39.8–52 mm): BPBM 35886, 52 mm (holotype);
ROM 68794, 39.8 mm (paratype); ROM 1555CS,
44.4 mm (paratype).
R. plesiomorphus Winterbottom:
39.7 mm (paratype).
CHARACTER
USNM
218164,
in Fig. 2) or taxa are noted. If only a single node is
listed, the character exhibits no homoplasy. Reference
to nodes or taxa that show reversals or independent
acquisitions for particular characters could vary from
those listed below, depending on optimization.
Character 1 (RDM 1, fig. 2; node A). Posterior subpelvic cavity on pelvic girdle: absent (state 0); present
(state 1). The pelvic girdle of Notograptus lacks a posterior subpelvic cavity, but the girdle is very reduced
(Gill & Mooi, 1993: figs 11, 12) and the absence may be
secondary. We therefore coded the character as inapplicable (n) for Notograptus.
Character 2 (RDM 2, fig. 2; node A). Subpelvic shelf
on pelvic girdle: absent (state 0); present (state 1). The
pelvic girdle of Notograptus lacks a subpelvic shelf
but, as noted in the previous character, the girdle is
very reduced and the absence may be secondary. We
therefore coded the character as inapplicable (n) for
Notograptus.
Character 3 (RDM 3, SVJ 4; node A state 1, node H
state 2). Number of pelvic-fin rays: I,5 (state 0); I,4
(state 1); I,2 (state 2). Notograptus has I,2 pelvic-fin
rays (Gill & Mooi, 1993: 339, figs 11, 12). This character was originally interpreted as an ordered transformation series and we see no reason to alter this
interpretation.
Character 4 (RDM 4; node A state 1, node H state 2).
Extensor proprius insertion: on one or two of innermost rays (state 0); on second, third and fourth rays
(state 1); muscle absent (state 2). Notograptus and
acanthoclinines lack an extensor proprius. The original interpretation did not include state 2, treating the
absence of the muscle as an autapomorphy of the
Acanthoclininae. With the addition of state 2 as potentially informative, we have opted to run this character
as unordered.
ANALYSIS
Characters are numbered 1–59, but are referenced to
those as numbered in earlier papers using the designation RDM for Mooi (1993) and SVJ for Smith-Vaniz
& Johnson (1990) (e.g. character 3 here is the same as
character 3 in Mooi, 1993 and character 4 in SmithVaniz & Johnson, 1990). Figures of the states of many
of these characters can be found in Mooi (1993) and
Smith-Vaniz & Johnson (1990), and are referenced
after the character number of the appropriate abbreviation. Interpretation for conditions of these characters in plesiopids and acanthoclinines are also found
in these publications. Some character conditions have
been reinterpreted with respect to those of RDM and
SVJ, and have been recoded to reflect our current
interpretation. Any such changes are noted under the
character description. After reference to literature
character numbering, the relevant nodes (as lettered
Character 5 (RDM 5, figs 3, 5; node A). Distal radials of spine-bearing dorsal pterygiophores: autogenous
(state 0); associated with the following proximalmiddle pterygiophore to form a complete bony ring
that interlocks with the articulating spine (state 1).
Notograptus has the derived state for this character
(Fig. 3; Gill & Mooi, 1993: 340, fig. 13). A similar morphology appears in some other taxa (e.g. blennioids,
labrids) (Gill & Mooi, 1993; Mooi, 1993), although we
have interpreted it as nonhomologous.
Character 6 (RDM 6, fig. 10; node A, reversal in
Notograptus and Acanthoplesiops echinatus). Parasphenoid keel: absent (state 0); present (state 1).
Notograptus lacks a parasphenoid keel (Gill & Mooi,
1993: 346, fig. 4) and we could not find one in Acanthoplesiops echinatus (although the specimen was
damaged); these are interpreted as reversals.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
185
Tra
ch
ino
ps
As
se
sso
Pa
r
rap
les
Ca
i
llop ops
Ste lesio
ps
en
e
Fra ichth
y
ud
ella s
Ple
si o
ps
Ac
an
tho
c
A.
litto linus
r
fus
e
u
A.
cu
rua s
s
A.
ma
rily
A.
na
ma
e
t
Be ti
lon
ep
ter
Be
yg
liop
ion
sb
Be
a
t
an
liop
en
sx
si s
No
an
tog
tho
rap
k ro
sso
tus
Ac
an
s
tho
p
A.
ind lesio
i cu
ps
s
hia
A.
ps
tti
ilog
A.
as
ter
ec
hin
atu
s
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
E
D2
1
2
J
3
Q
2
P1
O
N3 7
M
1I
L
K
3
7
H
G 7
3
F
2
A
B
C4
2
8
Figure 2. Strict consensus cladogram from two equally parsimonious trees from a branch-and-bound search using composite coding of the data set in Table 1 (no. of steps = 123; CI = 0.642; RC = 0.547; RI = 0.852). Nodes are lettered as in the
text. Characters supporting each node, those without homoplasy in bold, are: A – 1, 2, 3, 4(1), 5, 6, 7, 8; B – 9, 10(1), 11(1);
C – 12, 13, 14, 15, 16; D – 17; E – 18, 19(1), 20; F – 21, 22, 23; G – 10(2), 24, 25, 26; H – 4(2), 18, 19(1), 28, 29, 30, 31(1); I
– 32(1), 33; J – 46, 58; K – 34, 35, 36, 37, 38(1); L – 19(2), 39, 40(1), 41, 42, 46, 47; M – 43, 44, 45; N – 31(2), 48, 50, 51, 53;
O – 32(3), 49, 52, 54, 55, 56, 57, 59; P – 47; Q – 36, 38(2). Numbers below nodes are decay indices.
li
ds1
pm1
d4+pm5
v1
e1
r4
e3
v7
r1
Figure 3. Notograptus sp., BMNH 2002.1.19.16, 128.5 mm SL, first seven vertebrae and associated dorsal-fin structures in
left lateral view. Abbreviations: d4 + pm5, fourth distal radial fused with fifth proximal and middle radials; ds1, first dorsal
spine; e1, first epineural; e3, third epineural; li, ligament; r1, first rib; r4, fourth rib; v1, first vertebra; v7, seventh vertebra.
Scale bar = 5 mm.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
186
R. D. MOOI and A. C. GILL
Character 7 [RDM 7, fig. 11 (SVJ, fig. 1); node A, reversal in Notograptus]. Notch or rounded projection on
posterolateral margin of branchiostegal membrane:
absent (state 0); present (state 1). Notograptus lacks a
notch or rounded projection on the posterolateral margin of the branchiostegal membrane (Fig. 4; Gill &
Mooi, 1993: 346); this is interpreted as a reversal.
Character 8 [RDM 8, fig. 12 (SVJ 17, fig. 16); nodes A,
reversal at L]. Posterior border of preopercular sensory canal: closed (state 0); open (state 1). The posterior border of the preopercular canal is closed in
Notograptus (Gill & Mooi, 1993: fig. 7). Derived acanthoclinines exhibit a similar apparent reversal from
the derived condition.
Character 9 (RDM 9, fig. 14; node B). Basioccipital/
parasphenoid foramen: large (state 0); small (state 1).
The foramen is very small in Notograptus, providing
evidence of higher plesiopid affinity.
Character 10 (RDM 10, fig. 15; node B state 1, node G
state 2, independent acquisition of state 2 in Steeneichthys, reversal to state 1 in Notograptus). Adductor superficialis pelvicus inserts: on spine and first
three or more segmented rays (state 0); on spine and
first one or two segmented rays (state 1); on spine and
first ray only (state 2). Notograptus has this muscle
inserting on the spine and first two segmented rays
(state 1), but the reduced pelvic girdle supports only
two rays (Gill & Mooi, 1993: fig. 12). Hence, Notograptus may exhibit state 1 only as a result of reducing the
number of pelvic-fin rays and the associated muscle
insertions. If Notograptus is an acanthoclinine, its condition must be interpreted as independently acquired
or reversed to a condition of inserting on both segmented pelvic-fin rays. Treated as ordered as originally interpreted by Mooi (1993).
Character 11 (RDM 11; node B state 1, independent
acquisition of state 2 in Steeneichthys and
Notograptus). Branches on first segmented pelvic-fin
ray: three or more (state 0); two (state 1); one (state 2).
This was originally interpreted as an ordered transformation series, which seems a reasonable interpretation so is maintained here. Notograptus has an
unbranched first pelvic-fin ray (Gill & Mooi, 1993:
339), which is shared with Steeneichthys, but is considered an independent derivation.
Character 12 (node C). Swim bladder: present (state
0); absent (state 1). Notograptus and all plesiopids
other than Trachinops and Assessor do not have swim
bladders.
Character 13 [RDM 12, fig. 16 (SVJ fig. 18); node C,
reversal in Notograptus]. Zygapophysis on second
vertebra: small and dorsally placed (state 0);
expanded and displaced ventrally (state 1). Dorsally
placed in Notograptus, requiring interpretation as an
apparent reversal (Fig. 3).
Character 14 (RDM 13, fig. 14; node C). Posterior tip
of parasphenoid: not bifurcate (state 0); deeply bifurcate (state 1). Deeply bifurcate in Notograptus (Fig. 5)
placing the taxon among higher plesiopids. Note this
is a correction to the observations of Gill & Mooi (1993:
fig. 4c).
Character 15 (RDM 14, fig. 17; node C, reversal in
Calloplesiops). Base of fourth segmented pelvic-fin
ray: square-shaped with sharp angles forming the
articulation with the pelvic girdle and providing sites
for muscle attachment (state 0); not angular (state 1).
Notograptus has only two segmented pelvic-fin rays,
so this character was considered inapplicable (n) for
the genus, as it is for acanthoclinines.
Character 16 (RDM 15; node C, reversals in Calloplesiops, Fraudella, Notograptus). Spinous dorsal-fin
membranes: not incised (state 0); incised (state 1). The
spinous dorsal fin of Notograptus is not incised.
Character 17 (RDM 16, fig. 18; node D). Posterodistal portion of first proximal-middle radial of first anal
pterygiophore: does not contact second pterygiophore
(state 0); contacts second anal pterygiophore (state 1).
Notograptids exhibit state 0 (Fig. 6).
Character 18 (RDM 17, SVJ 1, fig. 2; node E, independently derived at node H). Lower lip configuration:
interrupted by isthmus (state 0); continuous (state 1).
The lower lip is continuous in Notograptus (Gill &
Mooi, 1993: 341), which supports a position among the
Acanthoclininae. An independent occurrence of this
condition is shared by Calloplesiops and Steeneichthys. Mooi (1993) was hesitant to include this character due to its variability and polymorphism in
pseudochromid taxa reported by Gill (1990). Such
polymorphism has not been observed among
plesiopids.
Character 19 (RDM 18, SVJ 6; node E state 1, independently derived at node H, node L state 2, state 3
autapomorphic for Notograptus). Number of total
caudal-fin rays: 27–29 (state 0); 24 (state 1); 18–22
(state 2); 13 (state 3). Notograptus has 13 caudal-fin
rays (Gill & Mooi, 1993: 341, fig. 13). This character
was run ordered in the analysis to be consistent with
the interpretations of RDM and SVJ.
Character 20 (RDM 19, fig. 13; node E). Base of pu2
haemal spine: broad (state 0); constricted (state 1).
Notograptus has a relatively broad base on the pu2
haemal spine (Gill & Mooi, 1993: 13).
Character 21 [RDM 20, fig. 19 (SVJ fig. 18); node F,
reversal at node L]. Ventral surface of anterior second and third vertebrae: ridged (state 0); smooth
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
A
187
pp
aip
pip
potp
np
itp
sotp
B
dp
atp
sobp
scp
ptp
llp
pop
bf
C
dp
Figure 4. Notograptus sp., BMNH 2002.1.2.1–2, 100.5 mm SL, dorsal, left lateral and ventral views of head. Mechanical
stippling is branchiostegal membranes. Abbreviations: aip, anterior interorbital pores; atp, anterior temporal pore; bf, barbel-like flap; dp, dentary pores; itp, intertemporal pore; llp, lateral-line pores; np, nasal pores; pip, posterior interorbital
pore; pop, preopercular pores; popt, posterior otic pore; pp, parietal pores; ptp, posttemporal pore; scp, supracleithral pore;
sobp, suborbital pore; sotp, supraotic pore. Scale bar = 5 mm.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
188
R. D. MOOI and A. C. GILL
Figure 5. Notograptus sp., ANSP 109653, 63 mm SL, ventral view of parasphenoid. Anterior to left. Parasphenoid stippled. Scale bar = 1 mm.
hs3
hs1
d2+pm3
pm1
as3
li
as1
Figure 6. Notograptus sp., BMNH 2002.1.19.16, 128.5 mm SL, first three anal-fin spines and their pterygiophores in left
lateral view. Abbreviations: as1, first anal spine; as3, third anal spine; d2 + pm3, second distal radial fused to third proximal and middle radials; hs1, first haemal spine; hs3, third haemal spine; li, ligament. Scale bar = 2 mm.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
(state 1). Ridged in Notograptus (Fig. 7). This apparent reversal also occurs in Beliops and Acanthoplesiops and serves to unite these taxa.
Character 22 (RDM 21, fig. 14; node F, reversal at
node L). Attachment sight on basioccipital for Baudelot’s ligament: on lateral fossa (state 0); on medial, triangular, slightly raised process (state 1). Raised
processes occur in Notograptus, but from lateral fossa
(Gill & Mooi, 1993: fig. 4). A similar apparent reversal
occurs in the derived acanthoclinines Beliops and
Acanthoplesiops.
Character 23 (RDM 22, fig. 20; node F, reversal in
Notograptus). Posterior sphenotic spur: small and
closely applied to anterior spur (state 0); large and
widely separated from anterior spur (state 1). Sphenotic spur absent or perhaps closely applied in
Notograptus making this a reversal.
Character 24 (RDM 23, fig. 21; node G, reversal in
Notograptus). Abductor superficialis pelvicus: overlies arrector ventralis pelvicus posterior to the infracarinalis anterior (state 0); does not overlie arrector
ventralis pelvicus posterior to the infracarinalis
189
anterior (state 1). In Notograptus the abductor
superficialis pelvicus overlies the arrector ventralis
pelvicus posterior to the infracarinalis anterior (Gill
& Mooi, 1993: 347, fig. 12), making this an apparent
reversal.
Character 25 (RDM 24, fig. 22; node G, reversal at
node O, independent acquisition in Steeneichthys).
Dorsal process for muscle attachment on segmented
pelvic-fin rays: large on first two or more rays (state 0);
large on first ray only (state 1). In Notograptus only
the first segmented ray bears a large dorsal attachment for muscle attachment (Gill & Mooi, 1993: 346,
fig. 11a). This places the genus among higher plesiopids. The character reverses in Acanthoplesiops,
the hypothesized sister taxon of Notograptus.
Character 26 (RDM 25, figs 23, 24; node G, independent loss in Trachinops, reversal in Beliops
xanthokrossos). Lateral process on posterior (middle
radial) portion of proximal-middle radial of spinebearing pterygiophores: present (state 0); absent
(state 1). The lateral processes are absent in Notograptus (Fig. 3; Gill & Mooi, 1993: fig. 13). Beliops xantho-
v5
r3
v1
e1
e3
r1
Figure 7. Notograptus sp., BMNH 2002.1.19.16, 128.5 mm SL, first five vertebrae in ventral view. Abbreviations: e1, first
epineural; e3, third epineural; r1, first rib; r3, third rib; v1, first vertebra; v5, fifth vertebra. Scale bar = 1 mm.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
190
R. D. MOOI and A. C. GILL
krossos has very small lateral processes on the first
two pterygiophores, but they are absent on remaining
posterior spine-bearing pterygiophores; we have coded
the species conservatively as showing a reversal.
Character 27 (RDM 26, figs 23, 24; independent acquisitions in Plesiops and Steeneichthys). Anterior process on distal-radial portion of spine-bearing dorsal
pterygiophores: present (state 0); absent (state 1).
Notograptus has an anterior process on all dorsal-fin
pterygiophores (Fig. 3).
Character 28 (SVJ 2; node H). Squamation:
head
scaled (state 0); head naked (state 1). The head is
naked in Notograptus.
Character 29 (SVJ 3; node H). Number of dorsal- and
anal-fin rays: low number of spines (7–16 dorsal and 3
anal) and high number of segmented rays (6–21 dorsal
and 7–23 anal) (state 0); high number of spines (17–26
dorsal and 7–16 anal) and low number of segmented
rays (2–6 dorsal and 2–6 anal) (state 1). Notograptus
has a high number of dorsal and anal-fin spines
(62–69 and 37–43, respectively) and low number of
segmented rays (1–2 in each fin) (Gill & Mooi, 1993:
340). This provides evidence for Notograptus as an
acanthoclinine.
Character 30 (SVJ 5; node H). Number of branched
caudal-fin rays: 15–17 (state 0); 14 or fewer (state 1).
Notograptus has 11 branched caudal-fin rays and is
coded as state 1. As acanthoclinines have 12 or 14, the
condition in Notograptus could be interpreted as autapomorphic. Here we have chosen to interpret the
reduced number of branched rays as homologous, i.e.
assuming homology in the absence of contrary evidence (Hennig’s Auxiliary Principle).
Character 31 (SVJ 7; nodes H state 1, reversal at node
L, node N state 2, Steeneichthys state 3). Number of
lateral lines: two (state 0); three (state 1); one (state 2);
none (state 3). Notograptus has a single lateral line
consisting of enlarged ossicles (Gill & Mooi, 1993: 342,
fig. 10). This character was run as unordered as there
is no evidence that the defined states form an ordered
transformation series.
Character 32 (SVJ 8, fig. 13; node I state 1, Notograptus state 2, node O state 3, independent acquisition of
state 3 in Steeneichthys). Infraorbital bones: five
(state 0); six (state 1); four (state 2); one (state 3).
Notograptus has four infraorbital bones (Gill & Mooi,
1993: fig. 6). The character was run unordered, there
being no clear polarity demonstrated and homology
among retained infraorbital elements unknown. It
could be interpreted as providing weak evidence of a
Notograptus + Acanthoplesiops clade, if polarized as a
reduction in number of elements among acanthoclinines being derived.
Character 33 (SVJ 9, fig. 13; node I). Suborbital
shelf: present (state 0); absent (1). Notograptus has a
suborbital shelf on infraorbital 3. The character is not
applicable to Acanthoplesiops so provides no data
regarding possible affinities with Notograptus.
Character 34 (SVJ 10, fig. 2; node K). Gill membranes: separate (state 0); united (state 1). Notograptus has the gill membranes united to each other and,
additionally, has them fused to the isthmus (Fig. 4).
The character supports Notograptus as a derived
acanthoclinine.
Character 35 (SVJ 11, fig. 14; node K, independent
acquisitions in Trachinops, Assessor and Steeneichthys, reversal in Beliops xanthokrossos). Supramaxilla: present (state 0); absent (state 1). Notograptus
lacks a supramaxilla (Gill & Mooi, 1993: fig. 6), supporting its placement among higher acanthoclinines.
Character 36 (SVJ 12, fig. 15; node K, reversal at node
Q). Teeth on infrapharyngobranchial 2: present
(state 0); absent (state 1). Notograptus lacks teeth
on infrapharyngobranchial 2 (Gill & Mooi, 1993:
338, fig. 9) like its hypothesized relatives among
acanthoclinines.
Character 37 [SVJ 13, fig. 17 (RDM, fig. 13); node K,
independent acquisitions in Steeneichthys and Acanthoclinus fuscus]. Haemal spine of pu2: autogenous
(state 0); united with vertebral centrum (state 1).
Notograptus has the haemal spine of pu2 united with
the vertebral centrum (Gill & Mooi, 1993: 341, fig. 13).
Although exhibiting some homoplasy, the character
supports a position of Notograptus among higher
acanthoclinines.
Character 38 (SVJ 14, fig. 2; node K state 1, node Q
state 2, independent acquisitions of state 1 in
Acanthoclinus rua and state 2 in Steeneichthys).
Number of dentary pore positions: five (state 0); four
(state 1); three (state 2). Notograptus has four dentary
pore positions (Fig. 4). This character was treated as
ordered.
Character 39 (SVJ 16, fig. 16; node L, reversal in
Notograptus). Primary opercular spine: plate-like or
fimbriate (state 0); pungent (state 1). Notograptus has
a fimbriate opercular margin, an apparent reversal
(Gill & Mooi, 1993: fig. 7).
Character 40 (SVJ 18, fig. 15; node L state 1, state 2
autapomorphic for Notograptus). Interarcual cartilage size: relatively long, almost as long as or longer
than pharyngobranchial 1 (state 0); relatively short,
less than half as long as pharyngobranchial 1 (1);
absent (2). Notograptus lacks an interarcual cartilage
(Gill & Mooi, 1993: 338, fig. 9). This character was
treated as ordered.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
Character 41 (SVJ 19, fig. 17; node L). Second and
third epurals: separate (state 0); fused (state 1).
Notograptus lacks epurals (Gill & Mooi, 1993: 341,
fig. 13), a state considered inapplicable (n) for this
character.
Character 42 (SVJ 20, fig. 18; node L). First neural
spine: autogenous (state 0); joined to centrum (state
1). The first neural spine is joined to its centrum in
Notograptus (Fig. 3).
Character 43 (SVJ 21, fig. 15; node M). Interarcual
cartilage shape: rod-shaped (state 0); cone-shaped (1).
As noted under Character 40, Notograptus lacks an
interarcual cartilage (Gill & Mooi, 1993: 338, fig. 9),
and interarcual morphology characters are inapplicable (n).
Character 44 (SVJ 22, fig. 16; node M). Metapterygoid-quadrate joint: smooth (state 0); interdigitated
(state 1). Notograptus has a smooth joint between the
metapterygoid and the quadrate (Gill & Mooi, 1993:
fig. 5).
Character 45 (SVJ 23, fig. 19; node M). Scapulocoracoid joint: smooth (state 0); interdigitated (state
1). Notograptus has a smooth joint between the scapula and the coracoid (Gill & Mooi, 1993: fig. 10).
Character 46 (SVJ 24; nodes J, independently acquired at node L, reversal at node O). Supernumerary spines on first anal-fin pterygiophore: two (state
0); one (state 1). Notograptus has one supernumerary
spine, which, in combination with the other characters, places it among higher acanthoclinines (Fig. 6).
This character exhibits considerable homoplasy
among acanthoclinines, however.
Character 47 (SVJ 25; nodes L, reversal at node P).
Middle radials of segmented-ray-bearing dorsal- and
anal-fin pterygiophores: autogenous (state 0); united
with proximal radials (state 1). The middle radials of
segmented-ray-bearing dorsal- and anal-fin pterygiophores of Notograptus form a single element with the
proximal radials (Figs 3 and 6; Gill & Mooi, 1993:
fig. 13), as they are in Beliops species and Acanthoplesiops hiatti. Remaining species of Acanthoplesiops
form a clade on the basis of a reversal to autogenous
middle radials in median fins. This resolves the polytomy among Acanthoplesiops presented in the original
analysis of Smith-Vaniz & Johnson, 1990).
Character 48 (SVJ 27, figs 1, 11; node N, independent
acquisition in Steeneichthys). Symphyseal flap on
lower lip: absent (state 0); present (state 1). Notograptus has an elongate, barbel-like flap on the lower lip
(Figs 1 and 4; Gill & Mooi, 1993: 341), which is interpreted as a modified symphyseal flap as exhibited by
191
Acanthoplesiops, which in some individuals can be
quite long.
Character 49 (SVJ 28, fig. 15; node O). Uncinate process on epibranchial 1: not parallel to main arm, so
that junction between two arms is ‘V’- shaped (state 0);
parallel to main arm, so that junction between two
arms is ‘U’-shaped (state 1). Notograptus lacks an uncinate process on epibranchial 1 (Gill & Mooi, 1993:
338, fig. 9), so the character states are inapplicable (n)
for this feature.
Character 50 (SVJ 29, fig. 17; node N). Size of hypural 5: large to moderate (state 0); very small or
absent (state 1). Notograptus lacks hypural 5 (Gill &
Mooi, 1993: 341, fig. 13). It is suggested here that the
loss of hypural 5 in this taxon is a direct modification
of the condition in Acanthoplesiops of a reduced
element.
Character 51 (SVJ 30, fig. 17; node N). Hypurapophysis: present (state 0); absent (state 1). Notograptus
lacks a hypurapophysis (Gill & Mooi, 1993: 341,
fig. 13), sharing this condition with Acanthoplesiops.
Character 52 (SVJ 31, fig. 16; node O). Secondary
opercular spine: absent (state 0); present (state 1).
Notograptus lacks a secondary spine on the opercle,
although it does bear a slight expansion in this region
of the bone (Gill & Mooi, 1993: fig. 7). Smith-Vaniz &
Johnson (1990: 249) incorrectly reported that
“. . . Fraudella has a series of prominent spines on the
posterior margin of the opercle”; Fraudella exhibits a
typical perciform condition with a single primary opercular spine.
Character 53 (SVJ 32, fig. 19; Node N). Ventral arm
of coracoid: moderately slender (state 0); robust (state
1). The ventral arm of the coracoid of Notograptus is
relatively robust (Gill & Mooi, 1993: fig. 10), suggesting a relationship to Acanthoplesiops, although the
general shape of these elements differs among these
taxa (cf. SVJ, fig. 19d, e).
Character 54 (SVJ 33, fig. 19; node O). Pectoral
radial formula: 2-1-1 (state 0); 3-0-1 (state 1).
Notograptus has a 2-1-1 radial formula (Gill & Mooi,
1993: fig. 10).
Character 55 (SVJ 34; node O). Supracleithral lateral-line canal: present (state 0); absent (state 1).
Notograptus has a lateral-line canal in the supracleithrum (Gill & Mooi, 1993: fig. 10).
Character 56 (SVJ 35; node O). Anterior-posterior
ceratohyal suturing: medial only (state 0); on both
medial and lateral surfaces (state 1). Notograptus has
the anterior and posterior ceratohyals sutured on the
medial surface only (Gill & Mooi, 1993: fig. 15).
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
192
R. D. MOOI and A. C. GILL
Character 57 (SVJ 36, fig. 20; node O). Scales in
mid-lateral series: not bilobed (state 0); bilobed (state
1). The mid-lateral series of scales of Notograptus are
not bilobed.
Character 58 (SVJ 37, fig. 21; node J, independent
acquisition in Notograptus). Body scales: some ctenoid (state 0); cycloid (state 1). Notograptus has cycloid scales (Gill & Mooi, 1993: 342), apparently an
autapomorphy among higher acanthoclinines.
Character 59 (SVJ 38, fig. 22; node O, independent
acquisition in Belonepterygion). Adductor mandibulae A2 section: laterally exposed (state 0); covered by A1
laterally (state 1). The adductor mandibulae of
Notograptus is modified, lacking distinct A1 and A2 sections (Gill & Mooi, 1993: 338, fig. 8); this character
was therefore considered inapplicable (n) for
Notograptus and uninformative regarding its phylogenetic position among acanthoclinines.
Excluded characters from previous studies
SVJ 15. Maximum standard length: c. 46–200 mm
(state 0); <27 mm (state 1). Notograptus exceeds
46 mm standard length (largest specimen examined
178 mm; 185 mm SL reported by Taylor, 1964), suggesting a reversal to larger size. This is a questionable
character; it is difficult to use SL as an index of size
when plesiopids vary so much in body shape. The
change in maximum size is not a particularly convincing characteristic for building phylogenies, with substantial changes in size notable among many
perciform groups as well as within the Plesiopidae
(e.g. Plesiopinae, Paraplesiopinae).
SVJ 26. Pale spot on pectoral-fin base: absent (state
0); present (state 1). Smith-Vaniz & Johnson (1990)
proposed that a pale spot on the pectoral-fin base is an
autapomorphy of the acanthoclinine genus Acanthoplesiops, but noted (p. 249) that the spot ‘is difficult to
discern in preserved specimens, but in fresh material
it is usually conspicuous’. Our survey indicates that
the derived state is more widely distributed, though
we acknowledge that we had difficulty in determining
its presence in some taxa. It is present in at least Beliops xanthokrossos (see Hardy, 1994: fig. 496), all four
species of Acanthoclinus (Paulin & Roberts, 1992:
pl. 8A–D), possibly Beliops batanensis (see SmithVaniz & Johnson, 1990: fig. 7), and several species
of Plesiops (e.g. P. cephalotaenia, P. corallicola,
P. coeruleolineatus, P. oxycephalus; see Masuda et al.,
1984: pl. 126I–M; Mooi, 1995: figs 11–13, 15 and 29).
Notograptus lacks a pale spot on the pectoral-fin
(Fig. 1). However, because of our difficulty in determining its distribution, this character was not
included in the analysis.
PHYLOGENETIC
ANALYSIS
Of the 61 characters surveyed for Notograptus and
plesiopids, two were excluded from the analysis and
seven could not be scored for Notograptus (Table 1).
Characters 1, 2 and 15 were not applicable due to modification and autapomorphic reduction of the pelvic
girdle in Notograptus. Character 41 could not be
scored because of the lack of epurals in Notograptus,
and 43 and 49 were inapplicable due to absence of particular dorsal gill elements. The adductor mandibulae
of Notograptus is autapomorphically modified and
obscures its interpretation for Character 59.
Several other characters have no direct bearing on
the position of Notograptus among plesiopids (17, 20,
27, 44, 45, 52, 54–57) but have been retained in this
analysis to maintain the structure of the original plesiopid tree of Mooi (1993) and acanthoclinine topology
of Smith-Vaniz & Johnson (1990). Hence, 42 characters are potentially informative regarding the relationships of Notograptus among plesiopids.
Analysis using all 59 characters (79 steps minimum) with composite coding for inapplicable characters and six characters ordered resulted in two equally
parsimonious trees (no. of steps = 123; CI = 0.642;
RC = 0.547; RI = 0.852). Both trees placed Notograptus as the sister taxon to Acanthoplesiops among the
Acanthoclininae as defined by Mooi (1993) and SmithVaniz & Johnson (1990). Topology changes involved
only the relationships among species of Acanthoclinus,
shown as a polytomy in the strict consensus tree
(Fig. 2). This is a relatively robust tree, particularly at
nodes A, H, L and O (note decay indices on Fig. 2).
With such decay values, strict consensus of trees even
six steps longer than the most parsimonious topology
retained these nodes and left Notograptus among a
polytomy of the derived acanthoclinines Beliops xanthokrossos, B. batanensis and Acanthoplesiops. Various analytical manipulations (e.g. unordering all
characters, deleting characters with unknown inapplicable states) resulted in identical strict consensus
topologies (excepting collapse of node I and loss of
Acanthoclinus monophyly due to character deletion)
and only slight decreases in CI, RC and RI. Constraining the placement of Notograptus as sister of the Plesiopidae lengthened the tree by 20 steps. Reductive
coding of inapplicable states (Strong & Lipscomb,
1999) resulted in the same two most parsimonious
trees and strict consensus result as the initial composite coding, with slightly different tree statistics as a
consequence of treating inapplicable states as
unknowns rather than as a new state (no. of
steps = 112; CI = 0.625; RC = 0.533; RI = 0.853). Our
interpretation and coding of these characters has no
affect on the conclusion that Notograptus is an
acanthoclinine.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
Of the eight characters that Mooi (1993) (Fig. 2,
node A) used to define plesiopids, Notograptus shares
three: a reduced number of pelvic-fin rays (3), modification or loss of the extensor proprius (4) and a bony
pterygiophore ring articulating with dorsal-fin spine
bases (5). Although several nonplesiopids also share
these conditions (see Mooi, 1993 and Gill & Mooi,
1993), two of the eight characters cannot be scored for
Notograptus (as noted above). Of the three others, a
secondary loss of a parasphenoid keel (6) and open
preopercular canal (8) are known to occur among
higher acanthoclinine plesiopids; the absence of a
branchiostegal notch (7) is a novel reversal in our
hypothesis. However, convincing evidence positions
Notograptus as a derived plesiopid through a series of
nested characters as reviewed below.
Placing Notograptus among higher plesiopids
(Fig. 2, nodes B–G) are six characters: small aortic
foramen (9), two or fewer branches on the first pelvicfin ray (11), loss of swim bladder (12), posterior end of
parasphenoid deeply bifurcate (14), large dorsal process on first pelvic-fin ray only (25), loss of posterior
lateral processes on dorsal-fin spine-bearing pterygiophores (26).
Notograptus also exhibits all six of the characters
listed by Smith-Vaniz & Johnson (1990) as defining
the Acanthoclininae (Fig. 2, node H): only two pelvicfin rays (3), complete lower lip (18), reduced number of
caudal-fin rays (19), head scaleless (28), dorsal and
anal fins with high numbers of spines (29) and reduced
number of branched caudal-fin rays (30). Smith-Vaniz
& Johnson (1990: 220) provided a list of characters
that differ between Notograptus and acanthoclinines,
but all (excepting perhaps the absence of a branchiostegal notch, character 7) could be explained as a
result of an autapomorphic feeding morphology and
behaviour in Notograptus. Notograptus and acanthoclinines also share an absence of the extensor proprius
pelvicus (4); because other plesiopids have modified
this muscle, we originally interpreted its absence as
‘inapplicable’ (n) but have reinterpreted it as an autapomorphy of acanthoclinines, including Notograptus.
Notograptus can be hypothesized to be among
‘higher’ acanthoclinines (Belonepterygion, Beliops,
Acanthoplesiops; Fig. 2, node K) with the following five
characters: fusion of gill membranes (34), loss of
supramaxilla (35), no teeth on second infrapharyngobranchial (36), haemal spine of pu2 united with centrum (37) and reduced number of dentary pores (38).
Although the conditions are unknown for the other
taxa, Notograptus and Belonepterygion share an
extraordinarily similar egg surface morphology modified by multiarmed projections raised above the
chorion by a central pedicel. This might be further
indication of a close relationship (Gill & Mooi, 1993:
Fig. 14). However, Acanthoclinus and some other
193
plesiopids (e.g. Assessor) have similar, although not
identical, egg surface morphology. Until homologies
are understood and character distribution among
other acanthoclinines is determined, the character
remains merely a tantalizing similarity.
Notograptus is related to Beliops and Acanthoplesiops (Fig. 2, node L) based on a further four characters:
reduced or absent interarcual cartilage (40), first neural spine fused to centrum (42), one supernumerary
spine on first anal-fin pterygiophore (46) and middle
radials of pterygiophores supporting dorsal- and analfin segmented rays forming a single element with
proximal radials (47). This relationship is also weakly
supported by three apparent reversals in these taxa:
preopercular canal no longer open (8), ventral surfaces
of three anterior vertebrae no longer smooth (21) and
Baudelot’s ligament again originating from a lateral
position on the basioccipital (22).
We have placed Notograptus as the sister taxon to
the genus Acanthoplesiops based on four characters:
presence of a symphyseal flap (48), reduced or absent
hypural 5 (50), loss of hypurapophysis (51) and robust
coracoid arm (53). The presence of only one lateral line
(31, state 2) can also be considered as evidence uniting
Notograptus and Acanthoplesiops, although this is a
relatively labile feature exhibiting several states for
which homology is difficult to determine. As we note
below, Notograptus and Acanthoplesiops species examined have straight guts with no bends or constrictions
demarcating a stomach, an apparent additional synapomorphy for these taxa, although character distribution of this feature has not been fully explored.
Speaking against the inclusion of Notograptus in
the Plesiopidae are nine characters: lack of a parasphenoid keel (6), no branchiostegal notch (7), a primitive condition of the insertion of the adductor
superficialis pelvicus to both segmented rays (10, state
1), dorsally positioned zygapophysis (13), dorsal fin
membranes not incised (16), posterior sphenotic spur
absent (23), abductor superficialis pelvicus overlies
the arrector ventralis pelvicus (24), opercular spine
flattened and fimbriate (39). Among these characters,
6 and 16 are known to reverse among other acanthoclinines, so could be considered somewhat more labile
and less informative. In our estimation, characters 7
and 10 provide the strongest evidence against the
inclusion of Notograptus among plesiopids. We have
no reasonable arguments to explain their apparent
reversal to the primitive condition necessitated by our
hypothesis. Character states for several others are not
presently known for all acanthoclinines (13, 23, 24)
and might exhibit a similar lability to 6 and 16,
although unlikely. The interpretation of characters 13
and 24 is somewhat subjective; additionally, the modifications in the morphology of the pelvic girdle, and
size and shape of the zygapophysis, might suggest
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
194
R. D. MOOI and A. C. GILL
that these are autapomorphic states in Notograptus. It
could be argued that branchiostegal and opercular
morphology (characters 7 and 39) has been modified as
a result of a unique jaw mechanism and feeding
behaviour. In any event, none of the nine homoplastic
characters associated with the position of Notograptus
as the sister to Acanthoplesiops provides substantial
evidence to overturn the hypothesis.
IMPACT
ON CLASSIFICATION AND RELATIONSHIPS
AMONG ACANTHOCLININES
The realignment of the Notograptidae as a sister
genus to the plesiopid acanthoclinine genus Acanthoplesiops resolves a longstanding phylogenetic enigma.
After being included among a fluid ‘Blennioidea’ for
over 100 years at the familial rank, and bounced to
other wastepaper basket higher taxa such as the
Trachinoidei, the peculiar genus Notograptus can settle among a growing Plesiopidae. Relationships of the
Plesiopidae to other perciforms remain problematic.
As sister to Acanthoplesiops, Notograptus exhibits a
condition that polarizes a previously equivocal character, middle radials of pterygiophores bearing segmented rays free or forming a single element with the
proximal radials (47). In Smith-Vaniz & Johnson
(1990: 248, fig. 12), the presence of trisegmented
40°
60°
80°
100°
pterygiophores (free proximal, middle and distal radials) is considered primitive and the presence of united
proximal-middle radials derived, occurring once in the
genus Beliops and interpreted as independently
derived in Acanthoplesiops hiatti. As Smith-Vaniz &
Johnson (1990: 255) noted, it is equally parsimonious
to interpret the character of having united proximalmiddle radials arising in a common ancestor of Beliops
+ Acanthoplesiops with a reversal to a trisegmented
condition in Acanthoplesiops indicus + (A. echinatus +
A. psilogaster). With the insertion of Notograptus as
sister to Acanthoplesiops, a choice can now be made
between these alternative interpretations in favour of
the latter. Notograptus has proximal-middle radials as
a single element, which suggests that Acanthoplesiops
hiatti is the sister to the remaining Acanthoplesiops
species (Fig. 2). We have been unable to determine the
condition of this or several other characters in the
new species of Acanthoplesiops naka; its position
remains equivocal (Mooi & Gill, 2004).
BIOGEOGRAPHY
With the inclusion of Notograptus, the distribution of
the Acanthoclininae expands to an area previously
unrecognized as being occupied by the subfamily
(Fig. 8; Smith-Vaniz & Johnson, 1990: fig. 3). In effect,
140°
120°
160°
180°
160°
40°
40°
4
4
4
4
20°
3
20°
2
2 2
12
Equator
3
12
3
3
2
20°
5
20°
3
3
40°
40°
40°
60°
80°
100°
120°
140°
160°
180°
160°
Figure 8. Distribution of Notograptus (hatching) and Acanthoplesiops (1: A. echinatus; 2: A. hiatti; 3: A. indicus; 4:
A. psilogaster; 5: A. naka).
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
an unexplained ‘hole’ in the distribution of acanthoclinines, northern Australia, has been plugged with
the addition of Notograptus. We also note that the distribution of Belonepterygion should be modified to
include at least Thevenard Island of north-western
Australia (MPM), Santa Cruz Islands of the Solomons
(USNM), Shepherd and Erromango Islands of Vanuatu (USNM), Loyalty Islands (USNM), and Middleton and Elizabeth Reefs (AMS; Gill & Reader, 1992).
As the sister group, Notograptus provides resolution
of the relationships among Acanthoplesiops through
the reinterpretation of the evolution of middle/
proximal radial association (character 46; described
above). Within Acanthoplesiops, A. hiatti, a West
Pacific species, is sister to a three species clade that is
Indo-West Pacific. Within this clade, the Indian Ocean
taxon, A. indicus, is sister to two allopatric West
Pacific species, A. psilogaster to the north (Japan, Taiwan, Batanes) and A. echinatus to the south (southern
Philippines and Moluccas) (Figs 8, 9A). This interpretation suggests that an endemic northern Australian
A
Japan, Phil (A. psilogaster)
Sulu, Banda (A. echinatus)
WCInd (A. indicus)
WCPac (A. hiatti)
NAust, Papu (Notograptus)
B
IndO, WPac (Congrogadinae)
WIndO (Anisochrominae)
IndO, WPac (Pseudoplesiopinae)
NAust (Assiculoides)
NAust (Assiculus)
C
Mala
EAsi
Phil
Sula
Papu
IndO
WPac
Aust
Figure 9. Area/taxon cladograms of A, Notograptus +
Acanthoplesiops; B, derived Pseudochromidae; C, marine
water striders (after Andersen, 1998: fig. 7e).
195
taxon (Notograptus) is sister to a more broadly distributed Indo-West Pacific taxon (Acanthoplesiops) with a
more complicated biogeographical history, perhaps
influenced by some of the factors outlined by Springer
& Williams (1990). Where Acanthoplesiops naka from
Tonga fits into this history cannot be determined at
this time (Mooi & Gill, 2004).
The basic Notograptus/Acanthoplesiops area relationships are broadly similar to those of derived
pseudochromids (Gill & Hutchins, 1997; Gill &
Edwards, 1999; Fig. 9B) if the Acanthoplesiops distribution pattern is viewed as a potentially repeating
Western Indian Ocean versus Pacific + Eastern Indian
Ocean pattern. This is the same pattern found in the
Congrogadinae. Their sister group, Anisochrominae, is
Western Indian; the sister to those is another repeating
pattern (Lubbockichthys – Pacific/Eastern Indian
Oceans ((Amsichthys + Pseudoplesiops – both Pacific/
East Indian Oceans) + (Chlidichthys + Pectinochromis
– both Western Indian Ocean))). The sister to all these
is Assiculoides (Kimberley District of Western Australia) and the sister to all of these is Assiculus (northwestern Australia) (Fig. 9B). Hence, the Notograptus/
Acanthoplesiops and the derived pseudochromid area
relationships can be reduced to the repeating Pacific/
Indian Ocean pattern with a sister relationship to
northern (or perhaps north-western) Australia. This
pattern is roughly equivalent to that seen in some
invertebrate taxa such as marine water striders
(Andersen, 1998) (Fig. 9C). Other fish taxa have an
Australian (Pacific/Indian Ocean) pattern, but the Australian distribution is temperate or southern rather
than northern (e.g. Pempheridae, R. Mooi, unpubl.
data; other Plesiopidae, Paraplesiops + Calloplesiops +
Steeneichthys). The Australian region has numerous
marine endemic families (Brachionichthyidae, Pataecidae, Gnathanacanthidae, Dinolestidae, Leptobramidae, Enoplosidae, Arripidae, Odacidae, Leptoscopidae)
and genera (e.g. certain aplodactylids, gobiesocids,
clinids, monacanthids, antenariids, gobiids, syngnathids etc.) that might provide similar repeated patterns (Mooi & Gill, 2002). To make strides in
understanding the biogeographical history of Australia, we need to determine the relatives to these groups
and search for repeated patterns of distribution.
This is in contrast to the hypothesis proposed by
Santini & Winterbottom (2002) using Brook’s Parsimony Analysis (BPA). Although a ‘South Australia
Basin’ is shown basal to all areas excluding New
Zealand, Indian Ocean areas form a series of basal
areas to several Indonesian and west and central
Pacific areas that are then more closely related to
independent ‘West’ and ‘North Australia’ basins (the
latter encompassing the distribution of Notograptus)
(Fig. 10A). Their analysis used several groups, including acanthoclinines; Notograptus was not included as
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
196
sin
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R. D. MOOI and A. C. GILL
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2
2
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© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
Figure 10. Area cladogram of the Indo-Pacific determined from a combined data set of 13 cladograms using Brooks Parsimony Analysis (A) after Santini & Winterbottom (2002); (B) with Plesiops and acanthoclinine data corrected, single most
parsimonious tree from a branch-and-bound search (no. of steps = 283; CI = 0.601; RC = 0.382; RI = 0.635). Numbers under
nodes are decay indices.
3
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PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
its position was unclear. With our hypothesis that
Notograptus is an acanthocline, we wanted to examine
what impact adding it to this data base might have on
the results. However, upon closer inspection of the
Santini and Winterbottom data set, we discovered
that the acanthoclinine and Plesiops portions have
several errors in coding of internal nodes and occurrence of taxa in identified areas. In addition, two taxa
are missing from the Plesiops cladogram (cf. Mooi,
1995: fig. 34 and Santini & Winterbottom, 2002: fig. 1b
VIII), and their inclusion alters coding of internal
nodes. With corrections, using BPA we found a single
most parsimonious tree with considerable difference
from that reported by Santini & Winterbottom (2002)
(Fig. 10B). This corrected tree perhaps reflects more
closely the general description of biogeography of the
region discussed earlier, where west and north Australian areas are found farther ‘down’ the tree and are
sister to a broader western Pacific and Indonesian
region, although it differs in that Indian Ocean
regions remain as a series of sister areas to Australian, Indonesian and West Pacific areas. Adding
Notograptus to this corrected data set has no impact
on the tree topology, not surprisingly given the size of
the data set. However, even this corrected biogeographical tree should be treated with some scepticism
because it exhibits low decay indices (Fig. 10B) and
further errors or omissions might exist in the coding of
the several included groups that we did not reexamine. There are also methodological questions that
should be reconsidered, for example, how to delimit
areas (six of the regions in the data set of Santini &
Winterbottom are not defined by endemic taxa), and
several listed nudibranch taxa do not occur in any of
the areas (i.e. they appear in the cladograms of the
analysed groups and in the data set, but occur outside
of the Indo-West Pacific in the Caribbean, Atlantic and
Mediterranean). There continues to be considerable
discussion on how to perform BPA and whether or not
it is the most appropriate method of analysis (Ebach,
Humphries & Williams, 2003; van Veller, Brooks &
Zandee, 2003; references therein). We have not pursued other methods with these data because of the
questionable area designations and unverified coding
for most taxa.
BIOLOGY
OF
NOTOGRAPTUS
The largest specimen examined was 178 mm SL,
although a 185 mm SL specimen was reported by
Taylor (1964). Gill & Mooi (1993: 342, fig. 14a, b)
described the eggs of Notograptus: 29–35 roughly cruciform chorionic projections arranged in a narrow ring
closer to one pole of the egg than the other, 1–3 rows of
projections wide. The projections are raised above the
surface by a short pedicel, and the projection’s arms
197
are produced into filaments, two to three greatly elongate. In other acanthoclinines, similar-looking eggs
bind together via the filaments and the egg mass is
guarded by the male in a burrow; similar behaviour is
expected in Notograptus. In the specimen with largest
ovarian eggs (USNM 173798, 170 mm SL), the eggs
come in three basic size classes: very small (0.5–
0.6 mm in diameter), small (0.9–1.2 mm in diameter)
and large (2.5–3.4 mm in diameter). This size distribution is indicative of a cyclical breeding cycle, perhaps lunar. Gravid females ranged in size from 88 mm
SL to 170 mm SL and were found in collections made
in February, April, May, June and September. Because
our sample is small and collections were restricted to
January through September, reproduction taking
place in other months cannot be precluded. The largest specimens carried the most eggs (170 mm SL, 63
right ovary + 53 left ovary = 116 mature eggs; 170 mm
SL, 47 + 42 = 89; 152 mm SL, 41 + 38 = 79; 103 mm
SL, 24 total; 88 mm SL, 18 + 14 = 32). Note that the
right ovary always contained more eggs than the left.
Males do not have a modified intromittent organ, and
eggs are likely fertilized after laying.
We have examined 99 specimens of Notograptus and
found 32 with identifiable gut contents (Table 2).
Eighteen of these contained whole alpheid shrimp,
always swallowed tail first (Fig. 11A, B). Thirteen
(usually smaller) specimens contained only one or two
claws, suggesting that smaller individuals are only
able to obtain these parts. However, a 51 mm fish
engulfed a whole 23 mm shrimp (claw tips to telson
tip) that filled the entire gut from the anus to well into
the buccal chamber (Fig. 11C). The largest individual
examined (USNM 173797, 178 mm SL) had eaten a
24 mm SL gobiid. This apparent exception to a strict
alpheid diet is likely an artefact of collection methods;
rotenone collecting kills smaller fishes first that are
often eaten by as yet unaffected bigger individuals
that may not be piscivores under normal circumstances. Considering that the gobiid was in excellent
condition in the gut (scales still intact, no digestion),
and that the specimen was collected with ‘barbasco
root’ (J. T. Williams, pers. comm.; a source of rotenone),
opportunistic feeding is a likely explanation for this
anomalous food item. Our observations strongly indicate that Notograptus are alpheid shrimp specialists.
Many morphological features of Notograptus appear
to be adapted to accommodate their feeding speciality.
The elongate body would permit entry into shrimp
burrows. The extremely large gape, knobbly teeth and
reduced gill arches would all facilitate eating large
prey whole. The gut is straight, lacking the complicated intestinal bends that would hamper ingestion
of large prey. Additionally, pleated skin around the
anus (reminiscent of a baleen whale throat) allows
evacuation of large indigestible items (Fig. 12A). In
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
198
Species
N : size range
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
Notograptus
99 : 44–178
Acanthoclinus fuscus
68 : 28–220
A. littoreus
7 : 56–119
A. marilynae
2 : 90–95
A. matti
1 : 52
Acanthoplesiops echinatus
1 : 21
A. hiatti
9 : 15–20
A. indicus
4 : 19–27
Acanthoplesiops naka
1 : 9.9
A. psilogaster
4 : 12–22
Beliops xanthokrossos
1 : 26
Belonepterygion fasciolatum43 : 12–42
Anisochromis kenyae
46 : 13.8–25.6
A. mascarenensis
11 : 13.3–25.5
A. straussi
82 : 16.1–28.3
Blennodesmus scapularis
120 : 28.5–87
Congrogadus hierichthys
30 : 50–159
Alpheids
whole (w),
parts (p)
Stomatopods (s)
or shrimps (h)
Amphi/Isopods
18w, 13p
–
–
2
–
–
–
–
11
–
–
–
–
4
–
–
–
–
–
1p
UnID
Crust.
Digested fish
Undigested
fish
UnID
Empty
Other
(*incidental)
–
65
–
5
48
–
1 (gobiid
with scales)
2 (1 with 3
tripterygiids)
–
–
2
1 snail,
1 limpet
1 mollusc
1
–
–
–
–
1 snail
–
–
–
–
–
1
–
–
–
–
–
–
–
–
–
–
–
–
6
–
–
–
3
–
1p
–
–
1
–
–
–
2
–
–
–
–
–
–
–
–
1
–
–
–
–
–
–
–
–
4
–
–
–
–
–
–
–
–
1
–
–
–
–
–
–
–
–
43
–
2?
1s
–
9
–
–
2
28
–
1h
–
6
–
–
–
3
3 ostracods
1 crab
1 ostracods
–
3 s,2 h
1i
15
–
–
11
46
2p
2 s,2 h
2a
10
–
–
24
78
3 gastropods
1 ostracod
1* mollusc
1p
1h
1
5
1
–
4
17
–
R. D. MOOI and A. C. GILL
Table 2. Frequency of specimens with specific gut contents of Notograptus and selected plesiopids, anisochromines, and congrogadines. Size ranges in mm SL
Stomatopods (s)
or shrimps (h)
Congrogadus malayanus
12 : 31–70
C. spinifer
33 : 36–121
C. subducens
75 : 31–340
–
–
–
3
–
–
8
1
1w
3 s,2 h
–
10
–
–
2
15
4w
1 s,1 h
–
10
2
29
–
3* small
snails
–
C. winterbottomi
31 : 66.1–119
Halidesmus polytretus
2 : 57–57.3
H. scapularis
20 : 46–98
–
5h
12a
2
–
3
8
1 mollusc
–
–
–
2
–
–
–
–
–
–
–
–
–
–
–
H. socotraensis
6 : 39.6–69.5
H. thomaseni
78 : 29–134
Halimuraena hexagonata
28 : 22–60
H. shakai
57 : 22–55
Halimuraenoides isostigma
18 : 65–278
Haliophis aethiopus
2 : 49–50
H. guttatus
308 : 21–132
Natalichthys leptus
2 : 52–56
N. ori
2 : 54–60.5
N. sam
2 : 40–43
Rusichthys explicitus
3 : 40–52
R. plesiomorphus
1 : 40
Amphi/Isopods
UnID
Crust.
Digested fish
28 (a few of
these with
undigested fish,
but often mixed
with digested;
3 with Crustacea;
8 with 2–3 fish)
–
Undigested
fish
UnID
Empty
Other
(*incidental)
–
–
–
–
–
–
–
20 (some
might be
shrimps)
4
–
–
–
2
–
–
–
12
–
–
40
26
–
–
–
8
9
–
–
–
11
–
1p
1 s,11 h
2
11
1
–
17
13
–
1w?
–
–
4
–
–
8
5
–
–
1h
–
1
–
–
–
–
–
1w
2 s,6 h
24
105
2
6
30
132
–
–
–
–
–
–
–
2
2* sponge
spicules
–
–
–
–
–
–
–
–
2
–
–
–
–
–
–
–
–
1
–
–
–
–
–
–
–
3
1
pycnogonid?
–
–
–
–
–
–
–
–
1
–
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
Alpheids
whole (w),
parts (p)
Species
N : size range
199
200
R. D. MOOI and A. C. GILL
A
B
C
Figure 11. X-radiographs of Notograptus sp., MPM 32586, showing whole alpheid shrimps in their guts. A, 73 mm SL,
scale bar = 10 mm. B, Close-up of A, scale bar = 5 mm. C, 51 mm SL, scale bar = 5 mm, note claws of shrimp extend into
buccal cavity.
comparison, Plesiops and Belonepterygion have recognizable stomachs with sharp bends, and Acanthoclinus has a bend apparently confined to a shorter
portion of the posterior intestine in some species but
substantially convoluted with two loops in A. fuscus.
Beliops xanthokrossos has a straight gut with a substantial constriction that demarcates a short posterior
intestine. Acanthoplesiops examined have a straight
gut, which would serve as an additional synapomorphy with Notograptus. Pleated skin around the anus is
found to a lesser degree in Acanthoclinus (Fig. 12B),
and Acanthoplesiops has only slight anterior pleating.
Such pleating does not occur in other acanthoclinines,
Plesiops [P. nigricans (Rüppell), MPM 31314], Steeneichthys (S. nativitatus Allen, MPM SOL 98–32;
S. plesiopsus Allen & Randall, WAM P30629.016) or
other percoids [e.g. Cephalopholis cyanostigma (Valenciennes), MPM 31524; Perca flavescens (Mitchill),
MPM 20093]. A reviewer pointed out what appears to
be anal pleating in at least some Ecsenius (Springer,
1988: figs 22, 23), but this pleating is of very limited
extent being only about 6% of head length compared
with 30 + % of head length in Notograptus.
Diet of acanthoclinines other than Notograptus has
been difficult to determine, as the guts of most specimens examined were empty (105 specimens of 141)
(Table 2). Only Acanthoclinus and Acanthoplesiops
had specimens with identifiable gut contents. Most
Acanthoclinus contained unidentified crustaceans
(16); remaining specimens contained various molluscs
(four) and fishes (two). All of the few Acanthoplesiops
with gut contents contained crustaceans (nine), with
two of these having parts of alpheid shrimps. A further
outgroup, the genus Plesiops, feeds mostly on small
crustaceans, or parts of larger ones, and gastropods
(64% crustaceans, 32% gastropods, 2% fishes, 1% pelecypods, 1% ophiuroid arms of 322 specimens with
identifiable gut contents; 451 specimens had empty
guts). In Plesiops, at least 20% of the gastropod shells
contained hermit crabs, although most did not; some
had opercula intact, and one gut contained an abalone,
indicating that gastropods are a true portion of the
diet. Overall, data are limited for plesiopids, but it
appears that an alpheid diet is a specialization among
derived acanthoclinines and is likely an autapomorphy of Notograptus.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
201
A
p
a
gp
sr
a
gp
B
p
sr
p
a
gp
C
sr
Figure 12. Pleated anal region of A, Notograptus sp., USNM 173797, 148 mm SL; B, Acanthoclinus fuscus, MPM 32616,
190 mm SL; C, Congrogadus subducens, MPM 32613, 340 mm SL. Anterior to left. Note that pleated region in the
Notograptus specimen is as large as the others’ despite being the smallest individual illustrated. Abbreviations: gp, genital
papilla; a, anus; p, pleating; sr, first anal-fin element, spine in (A) and (B), ray in (C). Scale bar = 4 mm.
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
202
R. D. MOOI and A. C. GILL
COMPARISON
WITH CONGROGADINES: ADAPTIVE
CONVERGENCE
The Congrogadinae, or eel blennies, and Notograptus
share a long taxonomic history where they were
regarded as potential relatives in the Blennioidei of
Regan (1912) and Gosline (1968), or even the Trachinoidei (Nelson, 1984). Today, congrogadines are firmly
ensconced as a derived subfamily among the Pseudochromidae by Godkin & Winterbottom (1985) with corroboration by Gill (1990), and any similarity between
congrogadine and Notograptus specializations is
undoubtedly a result of convergence, i.e. independent
derivation from different pre-existing characters (Gill
& Mooi, 1993).
Knowing this, their morphological convergence is
quite extraordinary and the original hypotheses of
close relationship of these taxa by Regan (1912) and
Gosline (1968) are understandable. Along with the
elongate body (cf. Figs 1 and 13), Gosline (1968: 45, 60)
noted that both taxa have the anterior portion of the
suspensorium only weakly connected to the posterior
portion. However, the condition in each is clearly not
homologous; in congrogadines the endopterygoid has a
weak connection to the metapterygoid and no attachment to the ectopterygoid, whereas in Notograptus the
loose connection is between the metapterygoid and
hyomandibular (cf. Gill & Mooi, 1993: fig. 5 and Godkin & Winterbottom, 1985: fig. 6). In both, this convergence presumably permits the mouth to open widely to
engulf large, whole prey. Both groups have reduced
branchial elements, although to a higher degree in
Notograptus. The gut of congrogadines is a straight
tube except for a very small S-shaped bend just before
the anus, very similar to the completely straight gut
found in Notograptus. In addition, the anus and surrounding skin is pleated, presumably to permit wide
expansion to ease excretion of large indigestible items
(Fig. 12C). Anisochromines, the sister taxon of congrogadines, also show slight pleating around the anus
that is otherwise absent in pseudochromids [e.g.
Cypho purpurascens (De Vis), MPM 32315]. The
remarkable convergence of derived and specialized
morphology between congrogadines and Notograptus
suggests that these are the result of adaptation
through similar selective regimes, perhaps as evidenced through diet and behavioural data.
Data on congrogadine diets are scarce. HarmelinVivien (1979) reported that Haliophis guttatus of
Madagascar is essentially a diurnal predator feeding
primarily on shrimp (45% of diet) and brachyuran
crabs (22% of diet), and secondarily on galatheid
crabs, amphipods, fish eggs and hermit crabs. Only a
few fishes were found in the 132 specimens examined.
Maugé & Bardach (1985: 376; our translation) stated
for Halimuraenoides isostigma that ‘the stomach
contents, visible in radiographs, chiefly are shrimps of
the Alpheidae and very rarely fishes’. Our own
observations of 830 congrogadine specimens using Xradiography (Table 2) indicate that most eat crustaceans (306 specimens), with stomatopods, penaeids
and amphipods or isopods being the most common
identifiable types (87 specimens). Evidence of alpheid
shrimps was found in only 11 specimens (five whole
shrimps in larger Congrogadus specimens, one whole
shrimp in Haliophis guttatus, Fig. 14), although a further 208 specimens contained unidentified crustaceans. Some specimens, mostly larger Congrogadus
subducens, contained fishes (40), many of which were
well digested, ruling out rotenone collecting artefact.
Molluscs were found only as incidental food items in
five specimens, so are not generally a part of a normal
congrogadine diet. One Natalichthys specimen contained what appeared to be a pycnogonid, and two
Haliophis guttatus specimens had incidental sponge
spicules ingested, suggesting foraging among sponges
(Table 2). The diet of Rusichthys, sister to all other
congrogadines (Winterbottom, 1986), is unknown. The
sister group of congrogadines, the Anisochrominae,
eat mostly crustaceans (shrimps, crabs, ostracods in
46 of 49 with identifiable stomach contents) and only
rarely molluscs (three of 49) (Table 2).
Congrogadine morphology is well suited to a behaviour of engulfing large crustaceans from confined
spaces such as burrows or narrow coral interstices.
Such behaviour would likely be similar to that of
Notograptus, which is inferred to involve entering alpheid shrimp burrows and eating the shrimp whole.
The convergence of morphology and feeding specialization between Notograptus and basal congrogadines
Figure 13. Exemplar of the Congrogadinae, Congrogadus winterbottomi, WAM P.31582–001, 85.1 mm SL (holotype).
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
PHYLOGENETIC POSITION OF NOTOGRAPTIDAE
203
Figure 14. X-radiograph of Haliophis guttatus, SMF 29198, 62.5 mm SL. Note the whole alpheid shrimp in the gut.
provides a pattern attributable to adaptation in the
historical sense (Coddington, 1988; Larson & Losos,
1996). The feeding modes might have provided a common selective regime for convergence of morphology
among these taxa.
Indeed, feeding behaviour of Notograptus and congrogadines might yet prove more similar than we have
demonstrated here. Halimuraenoides isostigma, a
basal member of the congrogadines (Winterbottom,
1986), is reported as eating mostly alpheids (Maugé &
Bardach, 1985). If the diet of Rusichthys, the sister to
remaining congrogadines, is found to be predominantly burrow-inhabiting alpheid shrimps, the case
for convergent adaptation would be even more palatable. The congrogadine body form and other unique
features would be an adaptation to this specialized
diet, with a secondary broadening of food preference to
other crustaceans and fishes as body size increases.
As noted by de Quieroz (1998), repeating phylogenetic patterns of morphology and behaviour might be
consistent with an adaptive explanation, but alternative explanations are not falsified. For example, a
straight gut might be strictly a function of being narrow-bodied, as perhaps a folded gut cannot be accommodated in the confines of an eel-like body. However,
true eels (Anguilliformes) seem not to be so restricted,
having a separate stomach overlying the intestine [e.g.
Moringua edwardsi (Jordan & Bollman), MPM 24972;
Gymnothorax moringa (Cuvier), MPM 30833]. Func-
tional studies could be undertaken to examine the
enlarged gape and whether or not reduced gill arches
provide an advantage for eating large prey. We think
that the loose connection in the suspensorium of congrogadines and Notograptus might function somewhat
like the distensible jaws of snakes to permit the
engulfing of large prey (and breaking the general rule
of never eat anything larger than your head). However, if other examples of fishes with elongate body,
large gape, reduced branchial arches, straight gut and
stretchy anus can be correlated with eating alpheid
shrimps or similarly hard-bodied, relatively large,
burrow-inhabiting or otherwise confined prey, an even
stronger case for convergent evolution and adaptation
to a particular selective regime could be put forward.
ACKNOWLEDGEMENTS
Specimens and/or X-radiographs were kindly provided
by: C. D. Roberts and A. Stewart (NMNZ); M. Burridge, M. Rouse, R. Winterbottom and M. Zur (ROM); S.
Jewett, S. Raredon, V. G. Springer and J. Williams
(USNM); W. Saul (ANSP); J. B. Hutchins and S. Morrison (WAM); M. McGrouther (AMS); P. Pruvost
(MNHN); U. Zajonz (SMF); R. Fricke (SMNS); E.
Anderson and A. Bentley (RUSI). X-ray material listed
from other institutions was provided through R. Winterbottom (ROM) who also kindly provided access to
reference material otherwise difficult to obtain and
© 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141, 179–205
204
R. D. MOOI and A. C. GILL
commented on aspects of an earlier version of the
manuscript. Special thanks are due to S. Raredon for
X-raying hundreds of specimens. MPM and BMNH
material of Notograptus was collected with the cooperation of B. Hutchins (WAM). We are grateful to D. F.
Hoese for initially bringing to our attention that
Notograptus is an alpheid shrimp predator. P. Hurst
photographed the whole specimens in Figures 1 and
13. This work was supported, in part, by the United
States National Science Foundation under grant no.
DEB-9317695 to RDM.
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