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The New Zealand triplefin Grahaminasignata (Teleostei; Tripterygiidae): Ajunior synonym of G. gymnota fromTasmaniaKendall D. Clements a , Laith A. Jawad a & Andrew L. Stewart ba School of Biological Sciences, University of Auckland, Private Bag92019, Auckland, New Zealandb Museum of New Zealand Te Papa Tongarewa, P.O. Box 467,Wellington, New ZealandPublished online: 30 Mar 2010.

To cite this article: Kendall D. Clements , Laith A. Jawad & Andrew L. Stewart (2000): The NewZealand triplefin Grahamina signata (Teleostei; Tripterygiidae): A junior synonym of G. gymnota fromTasmania, Journal of the Royal Society of New Zealand, 30:4, 373-383

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f Journal of The Roved Society of New Zealand,Volume 30, Number 4. December 2000. pp 373 384

The New Zealand triplefin Grahamina signata (Teleostei;Tripterygiidae): a junior synonym of G. gymnota fromTasmania

Kendall D. Clements*, Laith A. Jawad* and Andrew L. Stewart*

The triptcrygiid fish Grahamina signata Fricke & Roberts, 1993 was described from174 specimens collected from several exposed locations around the coast of NewZealand. It was distinguished from the Tasmanian G. gymnota (Scott 1977) by thenumber of pored scales in the lower lateral line, the number of branches of thesupraorbital tentacle, and the colouration of the pectoral-fin base. Examination of newmaterial of G. signata from the Auckland region, and re-examination of type materialof G. gymnota, show that the supraorbital tentacle and lower lateral line are notdifferences of taxonomic significance. Pectoral fin colouration in a published underwaterphoto of G. gymnota suggests that the colour difference was due to the effects ofpreservation. We therefore refer G. signata to the synonymy of G. gymnota. Thissynonymy, in combination with the restricted Tasmanian distribution and dispersalcapabilities of the species, suggests that G. gymnota was introduced to Tasmania fromNew Zealand.

Keywords triplefin; Tripterygiidac; Grahamina gymnota; Grahamina signata; synonymy

INTRODUCTION

The triplefin Grahamina gymnota (Scott 1977) was described from six specimens collectedfrom Montague Bay, south-eastern Tasmania, in August 1976. More recently, the species hasbeen observed around rocks, wharves and jetties in bays and estuaries in the Hobart region ofsouth-eastern Tasmania (Edgar et al. 1983; Last et al. 1983; Edgar 2000). G. gymnota hasbeen considered a Tasmanian endemic, although its nearest relatives are all found in NewZealand (Fricke 1994).

Originally placed by Scott in the genus Forsterygion, G. gymnota was subsequentlyassigned to the new genus Grahamina by Fricke & Roberts (1993). They included three otherspecies of triplefins in the genus, all from New Zealand: G. capito (Jenyns 1841) fromsheltered marine areas, G. nigripenne (Valenciennes 1836) from estuarine areas, and a newspecies, G. signata, from exposed marine areas. These species share no unique synapomorphies,although Fricke & Roberts (1993) believed that they were "easy to separate visually from thesimilar genus Forsterygion" in that they had "a bully head and robust body". Thus at presentit is not possible to diagnose Grahamina and Forsterygion as discrete monophyletic taxa. Weare currently reviewing the phylogenetic status and validity of these and related genera usingmolecular and morphological characters.

R00002. Received 14 February 2000; accepted 11 July 2000*School of Biological Sciences, University of Auckland, Private Bag 92019, Auckland, New Zealand†Museum of New Zealand Te Papa Tongarewa, P.O. Box 467, Wellington, New Zealand

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Fricke & Roberts (1993) diagnose the four Grahamina species by various combinations ofthe number of tubular pored scales in the upper lateral line, the height of the first dorsal finrelative to the second dorsal fin, the number of branches on the supraorbital tentacle, thenumber of basally pored scales in the lower lateral line, and the colouration of the pectoral fin(Fricke & Roberts 1993). Fricke (1994) considered G. signata most similar to G. gymnota,and distinguished the two species on the basis of three characters: (i) pored scales in thelower lateral line (10-15 in G. gymnota, 20-24 in G. signata), (ii) number of branches of thesupraorbital tentacle (3-7 in G. gymnota, 6-10 in G. signata), and (iii) the colouration of thepectoral fin base (uniform brown in G. gymnota, with a black and a white band in G. signata).

On 14 August 1999 the senior author collected two specimens (56 and 65 mm SL) of aGrahamina species in bait catchers off rocks at Whatipu, Auckland, which brought intoquestion the diagnosis of Grahamina species. The height of the first dorsal fin and the bluntshape of the snout excluded G. nigripenne and G. capita, respectively. The number of poredscales in the lower lateral line fell within the range reported lor G signata by Fricke (1994),although these were difficult to count. The pored scales were discontinuous anteriorly, so itwas difficult to determine where the lower canal originated. Both the left and right supraorbitaltentacles of the larger specimen fell within the range reported by Fricke (1994) for theTasmanian G. gymnota. In the smaller specimen the left and right supraorbital tentacles fellwithin the ranges of G. signata and G. gymnota, respectively, suggesting that this character isunreliable. In colour and shape, both specimens resembled the G. gymnota figured in Edgaret al. (1983). This underwater photograph of a Tasmanian fish clearly shows a black and awhite band at the base of the pectoral fin, a character diagnostic for G. signata in Fricke &Roberts (1993).

This confusion over the identification of the two Whatipu specimens led us to examine thestatus of nominal species in the genus Grahamina. The aims of the present study are: (a) todetermine the diagnosis of nominal Grahamina species, (b) to determine the number of validGrahamina species, and (c) to determine the correct nomenclature for these species. Thethird aim requires a re-examination of the status of Forsterygion robustum (Clarke 1879), aspecies referred to the synonymy of/7, varium by Fricke (1994).

METHODS

We attempted to identify material examined to genus and species following Fricke & Roberts(1993) and Fricke (1994). Counts and measurements follow Hubbs & Lagler (1958).Measurements were performed with digital calipers. Lateral line scale counts included allpored scales. Supraorbital tentacle counts included tentacles greater in length than width.Where possible, lateral line scales and supraorbital tentacles were counted on both sides ofthe body (thus each specimen has two counts). Material examined from the Museum of NewZealand Te Papa Tongarewa (NMNZ), the Queen Victoria Museum, Launceston, Tasmania(QVMT) and our own collections held at the University of Auckland is presented in Appendix1.

The total sample sizes of specimens examined for various characters are presented inTables 1-6. It was not possible to count pored scales in the lower lateral line of many of theWhatipu G. signata specimens. These pored scales are visible against the darker backgroundpigmentation in darker coloured males and in specimens preserved for a length of time, butnot in the lighter coloured females from the Whatipu site.

RESULTS AND DISCUSSION

Meristic counts for the four recognised Grahamina species are presented in Tables 1—4.Although the meristics of G. gymnota and G. signata overlap with G. capito, modal differences

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Table 1 Frequency distribution of dorsal spines and rays of Grahamina species. Asteriskindicates holotype.

Species (n)

Dl spines D2 spines D3 rays

7 8 18 19 20 21 22 12 13 14 15 16

G. gymnota (39) - 14 24* 1 - 1 28* 9 1 - - 4 27 8*G. signata (82) - 24 55* 3 - 1 35 42* 4 - - 41 38* 3G. capilo (75) 4 64 7 - 2 36 30 7 - - 22 50 3G. nigripenne (29)- 29 - - 9 1 9 - 1 - 1 0 1 8 1

2103021

20*54*344

17172_

Table 2 Frequency distribution of anal fin rays of Grahamina species. Asterisk indicatesholotype.

Species (n) 24 25 26 27

G. gymnota (39) -G. signata (82) 1G. capita (75) 9G. nigripenne (29) 5

Table 3 Frequency distribution of scales in the upper and lower lateral line of Grahaminaspecies. Asterisks indicate left and right sides of holotype.

Species (2n) 14 15 16 17 18 19 20 21 22 23 24 25 26

Upper lateral lineG. gymnota (76) - - - - - - 1 4 23 27 17 3G. signata (164) - - 6 12 53 67** 24 2G. capita (150) - 15 40 28 45 17 5G. nigripenne (54) 1 7 1 5 1 5 5 2Lower lateral lineG. gymnota (75) - - _ _ - - 3 12 1 2 3 1 * * 1 2 5G. signata (132) - - - - - - 8 47 26* 29* 17 5G. capita (172) - - 18 1 1 13 29 34 38 5 11 2 7 4G. nigripenne (22) - - _ _ _ 1 1 _ 4 1 2 4

Table 4 Frequency distribution of branches on the supraorbital tentacle of Grahaminaspecies. Asterisks indicate left and right sides of holotype.

Species (2n) 1 2 3 4 5 6 7 8 9

G. gymnota (1%) 28 39 6** 3 1 1G. signata 0^) - ~ - 35 74 39* 15* 1G. capita (25\) 16 88 55 33 37 14 8 - -G. nigripenne (24) 24 - - - - - - - -

(especially in scales in the upper lateral line) strongly suggest that the latter is a distinctspecies. These modal differences were also apparent in specimens of G. signata and G.capita collected from the same location at Orapiu Wharf, Waiheke Island, so are unlikely tobe the result of environmental influences. The presence of G. signata at Orapiu Wharfindicates that this species is not confined to exposed locations, as was thought previously

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Fig. 1 A Grahamina gymnota (signata), 63 mm SL, Whatipu, Manukau Heads, Auckland. B Grahaminagymnota, NMNZ P. 30579, 82 mm SL, Wrest Point, Derwent River Estuary, Tasmania. C Grahaminacapito, 78 mm SL, Island Bay, Wellington. D Grahamina nigripenne, 58 mm SL, upper creek,Mangawhai estuary.

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(Fricke & Roberts 1993; Fricke 1994). Instead, our collections indicate that the habitat of G.signata may be more broadly characterized as areas with a cover of encrusting invertebrates,especially the green-lipped mussel Perna canaliculus (C. Duffy pers. comm. 21 December1999 and pers. obs.).

Ranges for all of the body proportions we examined (Tables 5 and 6) overlapped broadlyfor all four species. This was surprising, given that G. signata and G. gymnota were readilydistinguishable (in the vast majority of cases) from G. capito and G. nigripenne on the basisof head shape (Fig. 1). The former two species are characterised by a relatively steep snoutprofile, although this is not reflected in snout length. Grahamina nigripenne is easilydistinguishable from congeners by its high first dorsal fin relative to the second dorsal fin, butwe required new diagnostic characters to separate the other three species.

We identified three external characters additional to snout profile that served to separateG. signata and G. gymnota from G. capito. First, we were able to define the difference inhead shape between G. signata/G. gymnota and G. capito by drawing a line through the tip ofthe premaxilla (i.e., the anteroventral tip of the upper lip) across the ventral margin of theorbit (Fig. 2). In G. signata and G. gymnota this line touches or passes anterodorsally to thedorsal extremity of the preoperculum (as visible externally), while in G. capito this line cutsthe preoperculum or touches its dorsal extremity. Second, in G. signata and G. gymnota the

Table 5 Morphometric measurements for Grahamina species as percentage standard length.Number of specimens examined: G. gymnota 11 (39-82 mm SL), G. signata 17 (50-74 mmSL), G. capito 10 (55-78 mm SL), G. nigripenne 10 (48-67 mm SL). Data for holotypes ofG. signata and G. gymnota given in parentheses.

Character

Length to 1sl dorsal originLength to 2nd dorsal originLength to 3rd dorsal originLength to anal originLength to pelvic originLength to ventLength to pectoral originHead lengthBody width at ventMaximum body widthCaudal peduncle depth

G. gymnota

12.2-22.5(17.1)20.7-31.3(31.3)67.9-74.5 (70.3)42.7-48.7 (45.3)18.5-27.0 (20.3)39.0-48.2 (40.6)24.6-30.8(31.3)22.7-28.0 (23.6)17.2-21.4(18.0)18.4-24.3(20.8)6.4-8.6 (7.9)

G. signata

12.9-18.5(15.1)27.9-33.3 (32.9)68.3-74.5 (73.9)40.4-50.0 (45.2)16.8-22.3(16.8)37.8-47.2 (43.8)23.3-29.2 (23.6)21.1-27.9(23.3)17.3-22.0(19.1)18.3-23.3(20.4)6.5-8.6 (7.8)

G. capito

16.2-22.223.6-35.269.5-74.744.1-50.920.6-23.039.0-49.128.6-33.825.7-28.417.2-21.217.8-20.86.1-8.1

G. nigripenne

16.4-25.032.7-40.069.1-75.945.0-52.719.8-32.041.4-49.129.3-35.424.0-28.715.9-20.514.9-19.98.4-11.4

Table 6 Morphometric measurements for Grahamina species as percentage head length.Specimens examined as for Table 5. Data for holotypes of G. signata and G. gymnota givenin parentheses.

Character

Snout lengthOrbit diameterInterorbital widthHead width at rear of orbitHead width at operculumMaximum head widthJaw length

G. gymnota

28.4-34.7(33.1)26.9-36.2 (32.8)9.8-16.3(13.2)50.9-62.9 (59.8)51.0-79.9(73.1)63.8-73.7 (72.2)33.7-44.2 (44.2)

G. signata

27.2-36.5 (27.9)28.1^2.4(28.5)9.0-18.9(17.7)57.5-65.9 (65.9)62.6-86.5 (64.9)65.4-79.2 (67.7)38.6-44.9 (42.0)

G. capito

25.1-30.521.4-35.08.0-14.857.0-62.660.8-69.960.1-70.732.8-45.6

G. nigripenne

24.6-34.127.6-37.810.4-17.956.1-66.554.2-82.356.1-71.625.5-39.9

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nape (i.e., the triangle formed by the base of thefirst dorsal fin and the tops of the opercularopenings) always has either superficial orembedded ctenoid scales. In G. capita this area iseither naked or has superficial or embeddedcycloid scales. Third, G. signata and G. gymnotahave a distinct groove passing transversely overthe top of the head in front of the first dorsal fin.This groove contains cephalic sensory pores ofthe supratemporal canal. In G. capito this lateralgroove is absent, but the central pore of thesupratemporal canal usually sits in a depression.

While the four characters listed above areuseful in separating G. signata and G. gymnotafrom G. capito, we observed several specimensof the latter species that were intermediate in oneor more (but never all) characters. Specimensfrom the Chatham Islands intermediate betweenG. signata and G. capito prompted Fricke (1994)to suggest that these species hybridized at thislocality. We re-examined these Chatham Islandspecimens, several of which had the steep snoutprofile characteristic of G. signata and G.gymnota. Fricke also noted the high number ofbranches on the supraorbital tentacles of theseChatham Island specimens. Since we found ahigh number of supraorbital tentacle branches onGrahamina specimens from elsewhere that areattributable to G. capito in all other respects, wedo not consider this character reliable. On thebasis of the other three diagnostic characters listedabove, the Chatham Island specimens discussed

by Fricke appear to be G. capito. The similarity between G. signata and G. capito suggeststhat these species are closely related, but at present we have no morphological or molecularevidence of hybridization.

Unlike Fricke & Roberts (1993) and Fricke (1994), we observed no separation between G.signata and G. gymnota in either pored scales in the lower lateral line or branches of thesupraorbital tentacle (Tables 3 and 4). Furthermore, the number of pored scales in the lowerlateral line did not allow the separation of any of the four Grahamina species. In any case thischaracter is problematical since these structures are (a) discontinuous and therefore difficultto count, and (b) almost impossible to observe in light-coloured fresh specimens. To ourknowledge this character has not been used by any other workers studying the group.

We observed 10 specimens of G. capito (collected from the Whangateau Estuary, WaihekeIsland and Wellington) with simple, unbranched supraorbital tentacles, a character previouslythought to be unique in the genus to G. nigripenne (Fricke 1994). The number of branches ofthe supraorbital tentacle thus served only to separate G. nigripenne from G. gymnota and G.signata. The unsatisfactory nature of pored scales in the lower lateral line and branches of thesupraorbital tentacle as key characters has led to the recent misidentification of Grahamina

Fig. 2 a, Grahamina gymnota showingrelationship of line drawn through tip ofpremaxilla (i.e. anteroventral tip of upperlip) across ventral margin of orbit to dorsalextremity of preoperculum. b, Grahaminacapito showing relationship of line drawnthrough tip of premaxilla (i.e. anteroventraltip of upper lip) across ventral margin oforbit to dorsal extremity of preoperculum.

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species in the literature. For example, a specimen of G. signata used by Brix et al. (1999) andEyton (1999) was found to be G. capito upon re-examination.

Fricke & Roberts (1993) and Fricke (1994) give pectoral fin colouration as the thirdcharacter that distinguishes G. gymnota from G. signata. Fricke (1994) describes the pectoralfin base as "brown in G. gymnota, without black and white streaks," while G. signata has a"pectoral fin base with a black and a white band." The underwater photograph of a TasmanianG. gymnota in Edgar et al. (1983) clearly shows a black and a white band at the base of thepectoral fin, and this banding was apparent on G. gymnota specimens we observed andcollected underwater in Hobart. We have also observed this banding in both G. signata andG. capito, thus this character is unreliable. We believe Fricke may have been misled in thischaracter because he compared freshly caught specimens of G. signata (specimens hecollected in 1992) with preserved museum specimens of G. gymnota collected between 1966and 1982. It is likely that the pectoral fin colouration has faded in the latter.

While we are able consistently to diagnose G. nigripenne and G. capito, we have beenunable to find any colour pattern, morphometric or meristic characters which separate G.signata from G. gymnota. This has led us to conclude that G. signata is a junior synonym ofG. gymnota. This conclusion is supported by mitochondrial DNA sequence data (A. Hickeyand K. D. Clements, pers. obs.).

The status of Forsterygion robustum

The systematics of the genus are further clouded by the confused status of Forsterygionrobustum (Clarke, 1879). The type specimen of F. robustum is lost. Waite (1913) and Parrott(1936) considered F. robustum a junior synonym of/7, varium, while Hardy (1989) andPaulin & Roberts (1992) treated these species as distinct. Paulin & Roberts (1992) used thename F. robustum informally to refer to taxa that were subsequently placed in Grahamina byFricke and Roberts (1993). F. robustum was referred to the synonymy of/7, varium by Fricke(1994), who designated a specimen of the latter as a neotype for F. robustum, despite the factthat the meristics of Clarke's specimen do not match F. varium (Paulin 1995). Fricke spenttwo days in 1992 at Clarke's type locality of Jackson's Bay on the west coast of the SouthIsland, where he caught two specimens of F. varium but no Grahamina specimens (Fricke1994). Fricke concluded that F. robustum was a specimen of/7, varium, and that Clarke hadmiscounted the dorsal rays in his description.

We agree with Paulin (1995) that a more likely explanation is that F. robustum was aGrahamina species and not F. varium, and that Clarke's description was accurate. Thisconclusion is almost inescapable, given that specimens of G gymnota have since beencollected at Jackson's Bay by the Museum of New Zealand (NMNZ P. 36246 and NMNZ P.36570). We therefore conclude that Fricke's neotype of/7, robustum does not represent thespecies originally described by Clarke (1879), and recommend that this neotype be consideredinvalid.

This conclusion raises the question of whether F. robustum is the junior synonym of G.capito (Jenyns, 1841) or the senior synonym of G. gymnota (Scott 1977). Scott (1977)referred to F. robustum as follows: "its (i.e. G. gymnota) nearest congener is F. robustumClarke 1879 - if that species is distinct; synonymized by Waite (1913) with F. varium(Forster 1801) - but it appears clearly to be distinct from all New Zealand forms." Thespecies encountered at Te Arai Point by Syms (1995) and referred to as 'Tripterygion'robustum is almost certainly G. gymnota. We found G. gymnota to be common at thislocation, but observed no G. capito.We are unable to establish the identity of/7, robustum with certainty in the absence of a typespecimen. Clarke's (1879) description, which includes fin ray counts but omits lateral line

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scale counts, agrees with both G. gymnota and G. capita. The "trifid supra-orbital tentacle"(Clarke 1879) tends to suggest G. capito, although this character is not diagnostic. The snoutprofile in his illustration is suggestive of G. gymnota. However, neither of our other headshape characters (line through tip of premaxilla along ventral edge of orbit in relation todorsal tip of preoperculum, groove in front of first dorsal fin) provide a definitive result fromthe illustration. Our other diagnostic character for the two species, nape scalation, is notdiscernable from the illustration. We are therefore unable to determine whether Clarke's F.robustum is G. capito or G. gymnota. Clarke may have been aware of Jenyns (1841)description of G. capito, but if so he does not seem to have taken it into account. Given thatwe cannot determine the identity of/7, robustum with certainty, we propose that it be declareda nomen dubium.

Key to the species of Grahamina

la. First dorsal fin high, often as high as second dorsal, with posterior spines longer thananterior (Fig. 3a); upper lateral line series with fewer than 19 scales

Grahamina nigripenne (Valenciennes in Cuvier and Valenciennes, 1836)lb. First dorsal fin low, shorter in height than the second dorsal fin, with posterior spines

equal to or shorter in length than anterior (Fig. 3b) 22a. Snout profile steep (Fig. la, b); line drawn through tip of premaxilla across ventral

margin of orbit touches or passes anterodorsally to dorsal extremity of preoperculum(Fig. 2a); nape with superficial or embedded ctenoid scales; transverse groove in front offirst dorsal fin containing cephalic sensory pores; New Zealand and Tasmania

Grahamina gymnota (Scott, 1977)2b. Snout profile sloping, concave or (rarely) steep (Fig. lc); line drawn through tip of

premaxilla across ventral margin of orbit cuts preoperculum or touches its dorsal extremity(Fig. 2b); nape naked or with superficial or embedded cycloid scales; no groove in frontof first dorsal fin, although medial sensory pore may sit at base of depression; NewZealand Grahamina capito (Jenyns, 1841)

Biogeography

The synonymising of G. gvmnota and G. signata raises the question of the biogeographicstatus of the Tasmanian population. This population of G. gymnota meets six of the 10criteria for inferring introduced species listed by Chapman & Carlton (1991), stronglysuggesting that they were established as a result of human activity.Criterion 1: Previously unknown in local region.There is no record of G. gymnota observed or collected in Tasmania prior to 1966 (i.e,.NMNZ P. 25423). Grahamina gymnota was described much more recently than most otherTasmanian triplefins, despite (a) the distinctiveness and large size of this species comparedwith other Tasmanian triplefins, and (b) the high likelihood of its detection given itsoccurrence around wharves and jetties in the Hobart region.Criterion 4: Association with or dependency on other introduced species.Four species of New Zealand marine invertebrates occur in the same Tasmanian locations asG. gymnota. It is thought that the chiton Chiton glaucus (F. Chitonidae), the crab Cancernovaezealandiae (F. Cancridae), the screw shell Maoricolpus roseus (F. Turritellidae) andthe starfish Patiriella regularis (F. Asterinidae) were accidentally transported from NewZealand to Tasmania around 1920 with live shipments of oysters (Edgar 2000).Criterion 5: Association with new or artificial environments.

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Fig. 3 a, Grahamina nigripenneshowing relative heights of first andsecond dorsal fins, b, Grahaminagymnota showing relative heightsof first and second dorsal fins.

G. gymnota is common around wharves and piers in theDerwent region (Last et al. 1983), and is thus prevalent inartificial environments in Tasmania. The senior authorfound the species to be abundant at Rosny Point,immediately opposite the main Hobart wharf area.Criterion 6: Relatively restricted distribution comparedto distributions of native species.The Tasmanian distribution of G. gymnota is limited tothe southeast corner of the island in the vicinity of Hobart(Edgar et al. 1983; Last et al. 1983). The only othertriplefin with an Australian distribution confined toTasmania is Forsterygion varium (Hardy 1989; Fricke1994), and this Tasmanian population is thought also tohave been introduced from New Zealand (Furlani 1996;Lockett & Gomon 1999). A recently established populationof F. lapillum in Port Phillip Bay, Victoria is thought tohave originated in the same way (Lockett & Gomon 1999).Criterion 9: Insufficient passive dispersal mechanisms.Prevailing ocean currents flow west to east across theTasman Sea (Brodie, 1960), thus it is unlikely that eggsand larvae of the benthic species G. gymnota could havereached Tasmania from New Zealand via passive dispersal.Criterion 10: Exotic evolutionary origin.All close relatives of G. gymnota are endemic to NewZealand.

CONCLUSION

The conclusion of this paper is that, ironically, the New Zealand endemic G. gymnota wasfirst described from specimens taken from a population introduced to Tasmania, Australia.

ACKNOWLEDGMENTSWe thank B. Smith at the Queen Victoria Museum, Launceston, for access to specimens; L. C.Clements, A. HickeyandB. Doak for help with collecting specimens; I. MacDonald for the photography;V. Ward for producing the line drawings; J. Montgomery for help with lateral line morphology; and R.McDowall, P. Last, C. Duffy, M. Lockett, J. Randall and J. Williams for helpful advice and comments.A. L. S. participated with support from the Biosystematics of N. Z. EEZ Fishes program, contractMNZX0003.

REFERENCESBrix, O.; Clements, K. D.; Wells, R. M. G. 1999: Haemoglobin components and oxygen transport in

relation to habitat distribution in triplefin fishes (Tripterygiidae). Journal of Comparative PhysiologyB 169: 329-334.

Brodie, J. W. 1960: Coastal surface currents around New Zealand. New Zealand Journal of Geologyand Geophysics 3: 235-252.

Chapman, J. W.; Carlton, J. T. 1991: A test of criteria for introduced species: the global invasion by theisopod Synidotea laevidorsalis (Miers, 1881). Journal of Crustacean Biology II: 386-400.

Clarke, F. E. 1879: On some new fishes. Transactions and Proceedings of the New Zealand Institute1878 11: 291-295, pi. XV.

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Edgar, G. J.; Last, P. R.; Wells, M. W. 1983: Coastal fishes of Tasmania and Bass Strait. Hobart, Catand Fiddle Press.

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Fricke, R.; Roberts, C. D. 1993: Grahamina, a new genus for robust-bodied triplefins (Teleostei:Tripterygiidae) from New Zealand and Australia, with description of a new species. StuttgarterBeiträge zur Naturkunde, (A) 504: 1-21.

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Hardy, G. S. 1989: The genus Forsterygion Whitley and Phillipps, 1939 (Pisces: Tripterygiidae) inNew Zealand and Australia, with descriptions of two new species. Journal of Natural History 23:491-512.

Hubbs, C. F.; Lagler, K. F. 1958: Fishes of the Great Lakes region. Ann Arbor, University of MichiganPress.

Jenyns, L. 1841: Fish. Part 16. In: Darwin, C. ed. The zoology of the voyage of H. M. S. Beagle duringthe years 1832-1836. Vol. 3. London, Smith, Elder and Co. Pp. 65-96, pls 16-20.

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Lockett, M. M.; Gomon, M. F. 1999: Occurrence and distribution of exotic fishes in Port Phillip Bay.In: Hewitt, C. L.; Campbell, M. L.; Thresher, R. E.; Martin, R. B. ed. Marine biological invasion ofPort Phillip Bay, Victoria. Centre for Research on Introduced Marine Pests, Technical Report No.20. Hobart, CSIRO Marine Research. Pp. 178-190.

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APPENDIX 1: MATERIAL EXAMINED

Museum materialG. gymnota New Zealand (G. signata of Fricke & Roberts, 1993) - NMNZ P. 30569, holotype of G.signata, 73 mm SL, Tauranga Bay, S side, opposite seal colony, 3 km SSW Cape Foulwind, 41° 47' S171° 27' E, rock pools and 0-1.5 m depth, rocks, sand and algae, exposed area, R. Fricke, 21 November1992; NMNZ P. 30570, paratypes, 4 spm.s, 51-62 mm SL, same data as holotype; NMNZ P. 13557, 4spm.s, 36-58 mm SL, Muriwai Beach, 36° 50' S 174° 26' E, intertidal pools, R. K. Dell and M. P. Dell,17 January 1962; NMNZ P. 17082, 3 spm.s, 43-68 mm SL, Castle Rock, Seal Rocks, Sugar LoafIslands, New Plymouth, 39° 03.5' S 174° 00.2' E, G. S. Hardy et al., 25 March 1985; NMNZ P. 24347,10 spm.s, 36-66 mm SL, Higgins Wharf, Napier Harbour; NMNZ P. 25282, 1 spm., 47 mm SL, TiPoint, 35° 16' S 174° 05' E, 4-6 m depth, 4 June 1971; NMNZ P. 30176, 3 spm.s, 68-76 mm SL, KapitiIsland, C.A. Fleming, December 1950; NMNZ P. 30691, 12 spm.s, 32-72 mm SL, Gisborne Wharf,Poverty Bay, 38° 40.8' S 178° 01.1' E, puerulus collector, T. Willis, 22 June 1993; NMNZ P. 32774, 5spm.s, 43-61 mm SL, 13-mile Point, Pyramid Rock, Greymouth, rotenone, rockpool, 3 August 1995,C. D. Paulin and P. Ryan; NMNZ P. 36246, 1 spm., 41 mm SL, -100 m North of Jackson's Bay Wharf,Jackson's Bay, 43° 58.9' S 168° 37.3' E, rotenone, 1.5-2.5 m, MoNZ Field Stn. H 02, 7 February 1999;NMNZ P. 36570, 2 spm.s, 39 and 44 mm SL, "Mock-maker Reef," Jackson's Bay, 43° 58.9' S, 168°37.3' E, rotenone, 0-3 m, MoNZ Field Stn. H 17, 14 February 1999.

G. gymnota Tasmania - QVMT 1976/5/204, QVMT Type No. 232, holotype, 64 mm SL, MontagueBay, south-eastern Tasmania, T. Walker, August 1976; QVMT 1976/5/205, 206, QVMT Type Nos.233 and 234, paratypes, 2 spm.s, 55 and 56 mm SL, same data as holotype; NMNZ P. 25422, 2 spm.s,

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39-40 mm SL, Triabunna, 42° 30' S 147° 55' E, M. Hortle, February 1982; NMNZ P. 25423, 2 spm.s,cleared and stained, Kettering Jetty, S. Bell, 11 June 1966; NMNZ P. 30579, 3 spm.s, 80-93 mm SL,Wrest Point, Hobart, Derwent River estuary, 2 m depth, K. Evans, 1982; NMNZ P. 30580, 2 spm.s, 47and 82 mm SL, Wrest Point, Hobart, Derwent River estuary, 2 m depth, K. Evans, 1982.

G. capita - NMNZ P. 13491, 5 spm.s, 55-71 mm SL, Portobello, Otago Harbour, 45° 50' S 170°39'E,R. J. Hoist, 13 August 1962; NMNZ P. 18137, 5 spm.s, 61-72 mm SL, Kawaroa, New Plymouth, 39°03.4'S 174° 03.7'E, 0-1 m depth, G. S. Hardy et al., 24 January 1986; NMNZ P. 21074, 3 spm.s, 59-73 mm SL, Waitangi, Chatham Island; NMNZ P. 25519, 5 spm.s, 60-76 mm SL, Owenga, ChathamIsland, 44° 02' S 176° 18' W, 26 January 1954; NMNZ P. 26640, 1 spm., 71 mm SL, Manukau Bay,Owenga, Chatham Islands; NMNZ P. 26658, 12 spm.s, 33-80 mm SL, Owenga Wharf rockpools,Chatham Island; NMNZ P. 26764, 14 spm.s, 54-89 mm SL, Napper Point, Port Hutt, Chatham Island;NMNZ P. 28332, 2 spm.s, 56-86 mm SL, Boat Cove, Lottin Point, 37° 33.0' S 178° 08.0' E, rotenone,rockpools in 0-0.5 m, East Cape Field Trip Stn. E6, I May 1992; NMNZ P. 28850, 2 spm.s, 46 and70 mm SL, Bare Island, Hawke's Bay, 39° 49.9' S 177° 1.5' E, rotenone, intertidal, C. Duffy, 9December 1991; NMNZ P. 30839, 5 spm.s, 59-79 mm SL, Lighthouse Reef, Cape Campbell, 41 ° 43.5'S 174° 16.5' E, rotenone, rockpool 0-0.5 m, Cape Campbell Field Trip Stn. CC1, 14 November 1993;NMNZ P. 30857, 8 spm.s, 35-80 mm SL, Limestone Reef, Ward Beach, 41° 50.5' S 174° 10.5' E,rotenone, 0.5 m, Cape Campbell Field Trip Stn. CC3, 15 November 1993; NMNZ P. 32725, 5 spm.s,60-75 mm SL, Lower Ahuriri Estuary, Napier, rotenone, mid-tide pool with limestone boulders andgravel, C. Duffy, 18 May 1995.

G. nigripenne - NMNZ P. 1327, 5 spm.s, 55-84 mm SL, Wanganui River Estuary; NMNZ P. 27830,10 spm.s, 40-69 mm SL, Mill Creek, Halfmoon Bay, Stewart Island, 44° 53.7' S 168° 07.5' E, 0-0.5 mdepth, 3 March 1992.

Material collected during this studyG. gymnota New Zealand (G. signala of Fricke & Roberts, 1993) - 2 spm.s, 56 and 65 mm SL,Whatipu, North Head Manukau Harbour, 37° 03' S, 174° 31' E, bait catchers, 2-3 m amongst rocks andkelp, 14 August 1999; 10 spm.s, 50-75 mm SL, Whatipu, North Head Manukau Harbour, 37° 03' S,174° 31' E, bait catchers, 2-3 m amongst rocks and kelp, 5 September 1999; 1 5 spm.s, 27-70 mm SL,Te Arai Point, 36° 10' S, 1 74° 39' E, slurp gun, 4-7 m on kelp-covered rocky reef on sand, mainly incaves,!5 December 1999; 12 spm.s, 43-56 mm SL, Orapiu Wharf, Waiheke Island, 36° 50.7' S, 175°08.9' E, bait catchers, under wharf, 29 and 30 December 1999, 1 January 2000.

G. gymnota Tasmania - 9 spm.s, 38-57 mm SL, Sandy Bay, Hobart, Derwent River Estuary, 42° 54' S,147° 20' E, slurp gun, 1-1.5 m under rock ledges on sand, 5 and 6 February 2000; 19 spm.s, 40-79 mmSL, Rosny Point, Hobart, Derwent River Estuary, 42° 52' S, 147° 21' E, slurp gun, 2-4 m amongstrocks on sand, 6 and 8 February 2000.

G. capita - 2 3 spm.s, 46-90 mm SL, Musick Point, Auckland, 36° 51' S, 174° 54'E, hand net, intertidalpools at low tide, 16 October 1997; 3 spm.s, 31-35 mm SL, Ulva Island, Patterson Inlet, Stewart Island,slurp gun, amongst red algae on sand, 1 February 1998; 6 spm.s, 38-79 mm SL, Island Bay, Wellington,41° 21' S, 174° 46' E, slurp gun, on cobbles in backreef inlet, 7 February 1998; 5 spm.s, 56-66 mm SL,Tinopai Wharf, Kaipara Harbour, 36° 15' S, 174° 15' E, hand net, intertidal pools at low tide, 11 April1998; 5 spm.s, 4 1 ^ 6 mm SL, Rocky Bay, Waiheke Island, 36° 49'S, 175° 03'E, hand net, intertidalpool at low tide, 4 June 1999; 6 spm.s, 36^46 mm SL, Whangateau Wharf, Whangateau Estuary, 36°20'S, 1 74° 45'E, rock on sand around wharf and boat ramp, 13 December 1999; 18 spm.s, 37-68 mmSL, Orapiu Wharf, Waiheke Island, 36° 50.7' S, 175° 08.9' E, bait catchers, under wharf, 29 and 30December 1999; 7 spm.s, 34.1-82.8 mm SL, Island Bay, Wellington, 41° 21' S, 174° 46' E, slurp gunand hand net, 1-2 m in subtidal inlet and rockpools, 7 and 8 January 2000; 5 spm.s, 57.3-73.3 mm SL,Seatoun Wharf, Wellington, 41° 19' S, 174° 50' E, bait catchers, 4-5 m on sand adjacent to wharf piles,8 January 2000.

G. nigripenne - 2 spm.s, both 55 mm SL, Pataua Estuary, Ngunguru Bay, 35° 43' S, 174° 30' E, baitcatchers, T. Hickey, 3 August 1999; 12 spm.s, 33-86 mm SL, Whangateau Wharf, WhangateauEstuary, 36° 20' S, 174° 45' E, hand net, rocks on sand around wharf and boat ramp, 13 December 1999.

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