Relationships and taxonomy of the New Zealand torrent fish, Cheimarrichthys fosteri Haast (Pisces: Mugiloididae)

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1 Journal of the Royal Society of New Zealand ISSN: (Print) (Online) Journal homepage: Relationships and taxonomy of the New Zealand torrent fish, Cheimarrichthys fosteri Haast (Pisces: Mugiloididae) R. M. McDowall To cite this article: R. M. McDowall (1973) Relationships and taxonomy of the New Zealand torrent fish, Cheimarrichthys fosteri Haast (Pisces: Mugiloididae), Journal of the Royal Society of New Zealand, 3:2, , DOI: / To link to this article: Published online: 14 Feb Submit your article to this journal Article views: 409 View related articles Citing articles: 17 View citing articles Full Terms & Conditions of access and use can be found at

2 Journal 0/ the Royal Society of New Zealand, 1973, Vol. 3, No.2, pp ,14 figs Relationships and Taxonomy of the New Zealand Torrent Fish, Cheimarrichthys /osteri Haast (Pisces: Mugiloididae) R. M. McDoWALL Fisheries Research Division Fisheries Research Publication 190 [Received by the Editor, 16 June 1972J Abstract Osteological study shows that Cheimarrichthys fosteri Haast, is very closely related to the mugiloidid Parapercis colias (Bloch and Schneider). As a result, Cheimarrichthys should be placed in the family Mugiloididae. C. /osteri is described and knowledge of its biology discussed and reviewed. INTRODUCTION Cheimarrichthys fosteri Haast, the torrent fish, is one of the least known species in the New Zealand freshwater fish fauna. During recent years, although knowledge of this fauna has grown rapidly, little work has been done on Cheimarrichthys. This paper examines the relationships of C. fosteri and attempts to clarify, expand, and synthesise current knowledge of the species. Previous papers about C. fosteri have included descriptive and distributional details, Maori utilisation, and a few brief notes on life history (Haast, 1874; McCulloch and Phillipps, 1923; Phillipps, 1926a, 1929; Whitley, 1927; McDowall, 1966). Two very brief papers (Lane, 1965; Topp, 1967) discussed the osteology of C. fosteri. MATERIALS AND METHODS A large series of samples of C. fosteri from throughout the known range of the species in New Zealand has been examined. Most of this material was collected by staff of the Fisheries Research Division; some is held in the collections of the Division but much is now in the collections of the National Museum, Wellington. Measurements of the fishes were taken with needle-point dividers, point to point, to the nearest half millimetre, except that a few small dimensions with clearly defined limits were measured to the nearest quarter millimetre. The following body dimensions were measured: total length (T.L.); standard length (S.L.); body depth at vent (B.D.V.); body width (v,reatest width behind head; B.W.); length and depth of caudal peduncle (L.C.P., D.C.P.); predorsal and preanal lengths (PreD., PreA.); basal length and greatest length of dorsal and anal fins (L.D.B., L.A.B., M.L.D., M.L.A.); pectoral and pelvic fin lengths (Pec., Pel.); prepelvic, pectoral-anal and pelvic-anal ler:gths (Prepe!., Pec..\:1., Pel.An.); head length, width and depth (length measured to edge of opercular flap, depth at occiput; H.L., H.W., H.D.); snout length (Sn.L.); postorbital head length (P.O.H.L.); interorbital width (I.O.W.); eye diarr:.eter (E.D.); length of upper and lower jaws (L.U.]., L.L.].); gape width (W.C.). Counts were taken of the following: pungent and non-pungent spines, and rays in dorsal fin, non-pungent spines, and rays in amd and pelvic fins, rays in pectoral and caudal fins, branchiostegals; gill rakers on first arch; vertebrae; pyloric caeca; pored scales in lateral line (pored scale number seems to vary greatly, probably because of the extension of a variable number of pored scales on to caudal fin base; these seem to vary in number and their small size makes them difficult to count).

3 200 JOURNAL OF THE ROYAL SOCIETY OF NEW ZEALAND Osteological study was made of material cleared in trypsin and alizarin stained (after Taylor, 1967). Abbreviations for bone names in Figs 1 to 6 are as follows:- actinost-act angular-ang articular-art basibranchial-bb basibranchial cartilage-bb c basihyal-bh basioccipital-boc branchiostegal-br ceratobranchial-cb cera tohyal-ch c1eithrum-cl coracoid-cor dentary-den ectopterygoid-ect epibranchial-eb epihyal-eh epiotic-epo epural-ep ethmoid-eth exoccipital-exoc frontal-fron hypobranchial-hpb haemal spine-h sp hyomandibula-hym hypohyal-hyh hypural-hyp hypurapophysis-hpp in terh yal-ih interopercular-iop lachrymal-lac lateral ethmoid-i eth lateral extrascapular-l ex maxilla-max median extrascapular-m ex mesopterygoid-mspt metapterygoid-mpt nasal-nas neural spine n!p opercular-op opisthotic--ops palatine-pal parasphenoid-ps parhypural-parh parietal-par pharyngobranchial-pb postcleithrum-pc1 posttemporal-ptm prefron tal-pref premaxilla-pnn preopercular-prop preural vertebra-pru v prootic-prt pterotic-pter quadrate-qu scapula-scap sphenotic-sph subopercular-sop suborbital-sorb supracleithrum-scl supra-intertemporal-s-it supraoccipital-soc symplectic-sym uroneural-uron urostylar vertebra-ur v vomer-vom RELATIONSHIPS " Inasmuch as Cheimarrichthys fosteri is the sole representative of the distinctive family Cheimarrichthyidae, its osteology and relationships are of more than casual interest "-Topp, Its relationships are little studied, although there has been some speculation about thi:;, C. fosteri generally is placed in the montypic family Cheimarrichthyidae, althoc:gh Schultz et at. (1960) considered that separate family status for this genus was undesirable. He placed C. fosteri, along with a broad assortment of genera, in the family Trichonotidae (including also H emerocoetes, another New Zealand genus customarily placed in a family of its own). Schultz acknowledged that placement of Cheimarrichthys in the Trichonotidae was to avoid large numbers of small or mono typic families. Phillipps (1929) stated that C. fostcri is related to the blue cod, Parapercis colias (Bloch and Schneider), and Allen ( 1957) expressed a similar view. Greenwood et al. ( 1966), while they combined many families in a wide-ranging study of higher fish classification, restored the separate family status of Cheimarrichthys. Although there is a marked similarity between C. fosteri and the New Zealand mugiloidids Parapercis colias and P. gilliesii (Hutton), none of these taxonomic changes has been supported by the necessary osteological and anatomical studies. Two brief papers on the osteology of C. fosteri (Lane, 1965; Topp, 1967) do not contribute materially to our understanding of the relationships of this species, except that Topp, in correcting errors in Lane's bone determinations showed that C. fosteri is a "typical" perciform. The relationships of C. fosteri remain somewhat unclear, and clarification awaits osteological comparisons with similar trachinoid fishes from the New Zealand region. New Zealand trachinoids comprise the genera Parapercis, Tewara,

4 McDowALL-Cheimarrichthys, the torrent fish 201 Hemerocoetes, Limnichthys, Leptoscopus, Geniagnus and Crapatulus, in addition to Cheimarrichthys, and it would seem logical to begin any comparisons with representatives of these genera. Tewara, Limnichthys, H emerocoetes, Leptoscopus, Geniagnus and Crapatulus are all rather specialised forms, especially the last three, and study of these seems unlikely to contribute much to our knowledge of the relationships of Cheimarrichthys. Certainly my brief examination of H emerocoetes, and Gosline's (1968) comments on this genus, suggest that H emerocoetes is a specialised and derived form. In contrast, Gosline (1968) considered that Parapercis is a generalised trachinoid, and therefore I have made comparisons between Parapercis and Cheimarrichthys. General form: Superficial similarity between Cheimarrichthys and Parapercis colias, the commonest New Zealand mugiloidid, is marked. The only obvious major difference between them is that whereas in P. colias the head is somewhat rounded, the eyes more or less lateral, the cheeks not expanded, and the mouth terminal, in Cheimarrichthys the head is strongly depressed, the eyes become dorso-iateral as the cheeks expand noticeably below the eyes, and the mouth is distinctly inferior in position. More minor differences include thickening of the fins and strengthening of the rays in Cheimarrichthys; the pelvic fins have moved apart so that they are widely separated. Scales are present on the head of P. colias, but not in C. fosteri. All these differences can be related to a change in habitat and adaptation to a very rapid water habitat by Cheimarrichthys. There are minor differences in fin spine and ray numbers, there being fewer spines and rays in the dors::ll fin of Cheimarrichthys, and vertebral COl.uts me higher by three to five vc:,teb::-ae. Only the inferior mouth, and these few minor meristic differer::ces exclude Cheimarrichthys from Parapcrcis as di9.gnosed bj Cantwell (1964). OSTEOLOGY Cheimarrichthys fosteri Previous work on the osteology of Cheimarrichthys comprises the two papers mentioned above (Lane, 1965; Topp, 1967) and discussion by Gosline (1968) on selected aspects of its osteology in relation to higher perciform classification. The following description is derived in part from these studies, but primarily from my own alizarin-stained material. Study is based on three specimens from the Waikanae River, New Zealand, 90, 95, 125 mm total length. The occurrence, disposition and general form of the bones is described as briefly as possible without prolonged discussion; reliance is placed on ilhstrations to indicate bone shapes. The bones of C. fosteri are heavily ossified; the head is strongly depressed and ventrally flattened, so that the orbits are dorsolateral in position. Neurocranium (Fig. 1, 2): There is a median dorsal ethmoid which projects forwards from the frontals, over the vomer as a narrow, vertical ridge with a cartilaginous tip. Paired prefrontals form the floors of the olfactory capsules, and lateral ethmoids project laterally to form a vertical wall between the olfactory capsule and the orbit on each side. Below the ethmoid region is a strong vomer, with a long, pointed shaft that covers the antcroventral surface of the parasphenoid. The head of the vomer projects downwards into the roof of the mouth behind the premaxilla as a crescent carrying an irregular band of small, but strong teeth. At the anterior of the cranium, the vomer folds upwards on each side of the rostral cartilage of the ethmoid region, behind the ascending and articular processes of the premaxillae, and forms the anterolateral margins of the cranium. A pair of small nasals (Fig. 3) occurs in the roof of the olfactory capsules, one on each side of the median rostral ethmoid ridge. In each orbit there is a pterosphenoid in the posteromedial orbit, a frontal forming the orbital roof, but also extending backwards to form the roof of the cranium, and a ring of bones around the eye. This ring comprises a lachrymal which has moved away from the orbital margin except for a narrow strut which does reach up to the orbit, and a series of six small sub orbital s, of which only the fifth and sixth border the fleshy orbit. A small dermal sphenotic forms a cap over a cartilage bone of the same name in the posterodorsal orbit, completing the chain of bones which carries the infraorbital laterosensory canal. The frontals, which are narrow above the orbits (the bony interorbital is narrow), broaden suddenly behind the orbits and form most of the cranial roof. Anteriorly

5 202 JOURNAL OF THE ROYAL SOCIETY OF N EW ZEALAND FIGS 1-2.-Cheimarrichthys fosteri neurocranium. Fig. 1. D orsal aspect. Fig. 2. Ventral aspect. they meet the ethmoid and posteriorly the supraoccipital. The supraoccipital is a large roofing bone which extends forwards between the parietals, which do not meet in the dorsal midline. The supraoccipital has a prominent median ridge, mostly on the posterior face of the bone. On each side, the parietals are small bones. D orsolaterajly, forming the lateral cranial margins are the sphenotics anteriorly, and larger, pterotics posteriorly; these form, with contributions from the parietals and epiotics, the floors of small temporal fossae. The epiotics form the posterodorsal corners of the cranial roof and a prominent spur projects posterodorsally to support one arm of the posttemporal. The exoccipitals are large, forming most of the posterior cranial wall, as well as making up part of the cranial roof between the epiotics and supraoccipital. They also contribute to the cranial floor between the prootic and foramen magnum. The exoccipitals meet posterodorsajly above the foramen magnum and thus exclude the supraoccipital from the margin of the foramen. The exoccipitals have small, stout condyles dorsolateral to the principal basioccipital condyle. The opisthotic is a small, flat, ovoid bone which overlies the suture ventrally between the pterotic and exoccipital. The basioccipital projects forwards as a broad, flat bone on the median floor of the cranium, between the exoccipitals, and projects medialiy above the parasphenoid and on each side meets the prootics as they project medially above the parasphenoid. The basioccipital forms a large, posterior condyle. The prootics are large

6 McDoWALL-Cheimarrichthys, the torrent fish 203 bones that form much of the floor of the auditory capsule. The parasphenoid is elongate and narrow, forming the anteromedial floor of the cranium, from the vomer, back to the basioccipital. Forming a rigid strut between the para sphenoid ventrally and the paired pterosphenoids, dorsally in the posterior orbit, is the basisphenoid, visible only from the lateral aspect. A series of dermal bones is present in the cranial roof; these bones carry laterosensory canals. They comprise supra-intertemporals along the dorsolateral cranial margins, fused to the sphenotics and pterotics, followed on each side by lateral extrascapulars; joining these, across the occipital region is a pair of medial extrascapulars which carry the supratemporal no s... ~. FIG. 3.-Lateral aspect of facial bones of left side of Cheimarrichthys fosteri. laterosensory commissure. One or other of these medial extrascapulars is sometimes divided. Behind each lateral extrascapular is a tri-radiate posttemporal. The dorsomedial arm overlies a spur projecting dorsolaterally from the epiotic, a very short dorsolateral arm carries a laterosensory canal confluent with that in the supra- and intertemporals, and a ventromedial arm has a strong ligamentous attachment to the opisthotic. Branchiocranium (Fig. 3, 4): The palatine is a flattened, ovoid bone, with an expansion anteriorly to form a facet which articulates with the maxilla, and posteriorly, a deep cleft into which projects a spur from the ectopterygoid. The palatine carries a few small teeth anteromedially. The triangular ectopterygoid forms a bridge between the palatine and quadrate, the narrow base of the triangle meeting and underlying a sector of the quadrate. The mesopterygoid is a flat, filamentous bone lying along the medial side of the ectopterygoid. There are no teeth on either the meso- or ectopterygoid. The quadrate is a flat, quadrant-shaped bone with a stout condyle. Ventrally, the quadrate is thickened into a ridge running back from the condyle and projecting beneath the symplectic. The metapterygoid forms a suture with the arc of the quadrate, rising to meet the forward projecting arm of the hyomandibula. The symplectic is a simple strut which fits into a trough above the thickened ventral border of the quadrate, and extends from behind the articular condyle on the quadrate backwards to meet the base of the hyomandibula. The hyomandibula, which fits up under the sphenotic and pterotic also fits inside the preopercular, and is much broadened dorsally at its articulation with the cranium, as well as projecting forwards and inwards, behind the eye, to meet the metapterygoid. The opercular series includes a stout, crescent-shaped preopercular which extends from the head of the hyomandibular downwards and forwards to the articular condyle on the quadrate, binding this section of the head together. There is a flat, elongate, interopercular which lies along the inner face of the ventral arm of the preopercular, and a boomerang-shaped subopercular, one arm of which projects vertically between the preopercular and opercular, and the other arm below and inside the opercular; the opercular is a roughly triangular, flat plate, the apex of the triangle, anterodorsally, has a well-developed facet that articulates loosely with the hyomandibula. The upper jaw comprises premaxilla and maxilla, there being no supramaxilla. The premaxilla forms most of the margin of the upper jaw. It carries a strong, though slender ascending process medially, and shorter, squat, round articular process lateral to the

7 204 JOURNAL OF THE ROYAL SOCIETY OF NEW ZEALAND ascending process, which articulates with the head of the maxilla. The alveolar process carries a strong crest distally ("post-maxillary process "-Topp, 1967: 920; "lateral process" Greenwood et al., 1966: 353), that fits below and behind the maxilia. The premaxilla carries an outer row of much enlarged teeth, with a band of smaller, villiform teeth inside. The maxiiia has a stout, hammer-like articular head that fits between the articular process of the premaxilla and an articular facet at the expanded anterior end of the palatine. The rest of the maxilla is an elongate, strap-shaped bone of simple form. The lower jaw comprises typical dentary, angular and articular; the dentary has a row of enlarged teeth marginally along about ft to! of its length with a band of villiform teeth inside. The tongue is very small, little more than a triangular protuberance at the anterior end of the hyoid arch. The basihyal is a short rod, scarcely projecting beyond the hypohyals; it is toothless. The upper and lower hypohyals are short, stout bones followed on each side by elongate ceratohyals, shorter, triangular epihyals, to which are attached tiny interhyals. The branchiostegals are in two groups on each side; two, larger and more widely spaced than the others, articulate on the middle of each ceratohyal and four smaller and more crowded ones on the distal end of the ceratohyal and on the epihyal. The urohyal comprises a flat horizontal plate with an elevated ridge posterodorsally between the urohyal and the basibranchials of the first arch. There are three basibranchials, the first two rather stout, the third amphora-shaped. The fourth basibranchial is represented by a cartilaginous block. The first two hypobranchials are elongated and carry a few tiny, toothed gill rakers distally. The third hypobranchial, however, is spatulate, the elongated arm lying forwards beneath the second. The five ceratobranchials form most of the pharyngeal floor. Each is an elongate bone, the first four carrying very short, toothed rabrs. The fourth articulates basally with the cartilage representing the fourth basibranchial. The fifth is rather broader than the others, curves forwards, and is heavily toothed on the upper, exposed, surface. The fifth ceratobranchials from each side almost meet medioventrally behind the fourth basibranchial cartilage. The four epibranchials arch back over the ceratobranchials forming much of the pharyngeal roof. The first is essentially a rod with a broad flat wing posterodorsally, the wing terminating abruptly on the medio-dorsal third. The second epibranchial is much shorter, and basically a stout block. The third and fourth are even shorter, and are essentially flat bones. There is no pharyngobranchial on the first arch (it may be fused with the epibranchial), but second, third and fourth pharyngobranchials are present as flat, toothed plates, lying above the toothed fifth ceratobranchial. The third pharyngobranchial is much larger than the others. Vertebral column and fin skeletons: The vertebral column comprises 12 thoracic vertebrae and 20 caudal vertebrae (plus the hypural) for a total of 32 (range 32-33). Vertebrae one to twenty carry epicentrals, and pleural ribs are present from vertebrae three to twelve. The supracleithrum of the pectoral girdle (Fig. 5) is a small, thin, triangular bone which originates underneath the posttemporal and lies over the outer face of the cleithrum. The cleithrum is the major bone of the girdle, being a curved, twisted bone. It has a vertical limb, the face of which is lateral, but as it extends ventrally, it twists and curves inwards to meet its opposite in the ventral midline. The scapula, which has a large foramen, and the coracoid, articulate onto a ridge deep in the medial surface of the cleithrum and form together a flat vertical plate onto the edge of which the four actinosts articulate, two and one-half on the scapula and one and one-half on the coracoid, the third bridging the scapulacoracoid suture. A slender, splint-like postcleithrum projects posteroventrally from the c1eithrum, behind the pectoral fin. The pelvic girdle lies beneath and between the pectoral girdles, just behind the symphysis of the two cleithra, but the two girdles have no bony attachment to one another. The pelvic girdle comprises two flat, triangular basipterygia with an extensive median suture. The fin rays are supported on the posterolateral edges of the basipterygia. The anterolateral edge of each basipterygium is folded downwards to fit inside the anteromedial ramus of the cleithrum. The fin carries a stout spine and five very stout and much branched rays. The dorsal fin comprises four (3-5) short and stout, pungent spines, one longer, more slender, non-pungent spine, and 20 (18-21) branched rays. The first spine has no pterygiophore. but fits into a notch on the anterior face of the pterygiophore which supports the second spine. This pterygiophore comprises fused proximal and medial elements, the bone being V-shaped. A small block-shaped distal pterygiophore fits between the diverging arms of the proximal-medial pterygiophore and is further enclosed by the diverging proximal end of the second spine. The second pterygiophore has similar structure and supports the third spine. Each of the branched rays is supported by a pterygiophore of similar form; the pterygiophores are a little crowded so that the first three overlie only two vertebrae, but thereafter, the pterygiophores interdigitate accurately between the neural spines of the vertebrae. The fin rays are stout and much branched. The anal fin has similar structure. The single spine is supported by the pterygiophore from the first branched ray; each of the 15 (14-16) rays is supported by its own pterygiophore; the pterygiophores interdigitate accurately with the haemal spines of the vertebral column.

8 McDowALL-Cheimarrichthys, the torrent fish 205 ci "'x. /., ~.. -.:.. eb 1-4 pb \ \ 1 6 FIGS 4-6.-Cheimarrichthys fosteri. Fig. 4. Dorsal (adpharyngeal) aspect of hyo-branchial skeleton: branchial skeleton of left side, and branchiostegals of right side not shown. Fig. 5. Lateral aspect of left pectoral girdle. Fig. 6. Left lateral aspect of caudal skeleton.

9 206 JOURNAL OF THE ROYAL SOCIETY OF NEW ZEALAND As with the dorsal fin, the pterygiophores of the anal fin consist of fused proximal and medial elements with a separate small block representing the distal one. The caudal skeleton (Fig. 6) comprises a dense fan of flat bony plates closely adjacent to the neural and haemal spines of the preural vertebra. In stained specimens I have studied, the preural centrum has a full neural spine, although Lane (1964) shows an abbreviated one and radiographs demonstrate that this is often the case. The single ural centrum carries a parhypural which basally imbricates with the base of the preceding (preural) haemal arch. The parhypural has a pair of very strong, laterally directed hypurapophyses. There are five hypurals, one and two more or less fused, a small gap to three which is similarly fused with four and five. There is a single pair of uroneurals which carries three epurals. The uroneurals enclose the fifth hypural anterodorsally and proximally, and are more or less fused with it. The uroneurals also enclose the base of hypural four, and again, are more or less fused with it. The limits of the various bones usually can be discerned but the bones cannot easily be prised apart from one another. The parhypural-hypural fan supports 14 (13-15) principal rays, the 12 (11-13) branched rays being very much branched. Eight or more procurrent rays are present dorsally and ventrally. There are many differences between my interpretation of bones (which agrees with that of Topp, 1967) and Lane's (1965). Topp has already shown that Lane has named the basisphenoid "orbitosphenoid", and also corrected Lane's interpretation of the frontoparietal region. My bone interpretations differ from Lane's in the following respects: (Lane's names in parentheses) :-prefrontal and lateral ethmoid, and part of vomer as seen from dorsal view (lateral ethmoid); part of frontal (parietal); parietal (epiotic); epiotic (part of exoccipital); basisphenoid (part of orbitosphenoid); pterosphenoid (part of orbitosphenoid); prootic (pterosphenoid); pterotic (part of exoccipital); lateral extrascapular (pterotic); symplectic (hyomandibula, which is shown reaching quadrate) ; cleithrum (coracoid); scapula (cleithrum); coracoid (scapula). Lane notes that "suborbitals and supraorbitals were cartilaginous and greatly reduced with the exception of the lachrymal in a specimen with a cranium about 18 mm long". In a fish somewhat larger, I have found the suborbital series well ossified, as did Topp (1967) in specimens with skulls 16 and 19 mm long. Lane also illustrated the eighth abdominal vertebra with two canals above the centrum, one enclosed by the neural arch, and a second by diverging arms of the neural spine. I can find nothing similar to this second canal and doubt its existence. eth ~ '\ \ '.. -- fron ~-=~. '1-./,,'eI" /./ _._ _---- ~ ) ~ '0' o,~ Ii ~::~~~ I ex FIG. 7.-Dorsal aspect of neurocranium of Parapercis colias.

10 McDowALL-Cheimarrichthys, the torrent fish 207 Parapercis colias Apart from comments on isolated aspects of the osteology of Parapercis by Gosline (1968), and some notes by Cantwell (1964), the osteology of Paraperns seems unstudied. The present study is based on alizarin-stained specimens 01 Parapercis colias, 280 and 200 mm long, from off Kapiti Island, New Zealand, and one 55 mm long from Foveaux Strait, New Zealand. The osteology of P. colias proved to be very similar to that of C. fosteri, and only points of difference are noted here. The same bones are present, they are usually the same shape, and differences generally are related to either (a) adoption of a rapid-water habitat by C. fosteri, or (b) movement of the lower jaw to take an inferior position in C. fosteri. C. fosteri lives in very swiftly flowing water. This seems to be related to much strengthening of the skeleton; ossification is generally heavy, fin supports are stouter, anu in particular, the fin rays have become very thick, strong, and much divided, especially in the pelvic and anal fins. The elements of the hypural fan are discrete in P. colias but fused into two plates in C. fosteri, and they are less firmly attached to the hypural centnun in the former. The hypurapophysis of P. colias is an elongate, slender spine, but a short, swu, spike in C. fosteri. The first two branchiostegals of C. fosteri are broadened but those ot P. colias are slender, like the others. Movement of the lower jaw to an inferior posltion nas had little impact on the osteology of C. fosteri; the basihyal is elongate in P. COiUlS nul very short in C. fosteri. P. colias has more numerous and slightly longer gill rakers; otherwise the hyobranchial skeletons of the two species are virtually indistinguishable. The interorbital of P. colias is narrower than that of C. fosteri (Fig. 7), and the suborbital series of P. colias rings the entire orbit, with the lachrymal also in tlte orbital margin (Fig. 8). The form of the maxilla is distinctive in P. colias; the head is cleft, with the articular process of the premaxilla entering tlte cleft. These few differences distinguish P. colias from C. fosteri. The two species are so similar that I have little hesitation in assigning them to the same family, thus not recognising the family Cheimarrichthyidae: this family was previously the only family of fishes endemic to New Zealand fresh waters. There seems little doubt that C. fosteri is derived from Parapercis, or a Parapercis-like ancestor, no longer in New Zealand seas. Although C. fosteri merits generic separation from Parapercis, there seems little justification for the retention of a separate family. Cheimarrichthys is therefore assigned to the family Mugiloididae, following the inclusion of the Parapercidae in the Mugiloididae by Greenwood et al. (1966)., " " --", //, /, / \, \ I \ I \ I J f ; fj \, /~\ ~~orb FIG. lac 8.-Suborbital-Iachrymal series of Parapercis colias. TAXONOMY Only one species of Cheimarrichthys has been described, C. fosteri Haast, 1874, although Stokell (1955) suggested that" there seems some doubt whether these large specimens [from the North Island] belong to the same species, as in addition to the difference in size, the habits appear to differ".

11 208 JOURNAL OF THE ROYAL SOCIETY OF NEW ZEALAND In the present study of a wide variety of taxonomic variates, I have been unable to find any characters which separate different stocks or taxa. Detailed comparisons of material from much of the range of the species in New Zealand demonstrated no differences of any taxonomic importance. Fishes from all areas were similar in both morphometric and meristic characters (see "Variation", also" Harper River Population "). It seems highly probable therefore, that there is but a single species of Cheimarrichthys. GENUS Cheimarrichthys Haast Cheimarrichthys Haast, 1874: 103 (type species Cheimarrichthys fosteri Haast by original monotypy). Chimarrhichthys Regan, 1913: (mis-spelling of Cheimarrichthys). Generic diagnosis: As a single species of C. fosteri is recognised, a generic diagnosis, distinct from the species description that follows would be redundant. FIG. 9.-Cheimarrichthys fosteri Haast, Waikanae River, 105 mm total length. Cheimarrichthys fosteri Haast (Fig. 9) Cheimarrichthys fosteri Haast, 1874: 103 (syntypes (3): Canterbury Museum, No. 79; type locality: Otira River "where that alpine torrent leaves its picturesque gorge"); Hutton, 1904: 43; Waite, 1907: 29; 1910: 389; Phillipps and Hodgkinson, 1922: 96; McCulloch and Phillipps, 1923: 19; Hiroa, 1926: 640; Phillipps, 1926a: 298; Whitley, 1927: 303; Phillipps, 1927a: 42; 1927b: 13; 1929: 166; Stokell, 1955: 62; Allen, 1956: 4; 1957: 5; McMillan, 1961: 143; Wisely, 1962: 213; Lane, 1965: 207; McDowall, 1966: 102; Skrzynski, 1967: 97; Topp, 1967: 189; McDowall, 1968: 13; Gosline, 1968: 41; Whitley, 1968: 68; Burnet, CTanfield and Benzie, 1969: 505; Hopkins and McDowall, 1970: Cheimarrichthys forsteri: Phillipps, 1926b: 485; Hefford, 1937: 73; Schultz et al. 1960: 275; McDowall, 1964a: 7; 1964b: 60. DESCRIPTION Trunk elongate but robust, cylindrical to slightly compressed behind head, becoming much compressed on caudal peduncle; dorsal profile arched, ventral profile rather flattened. Head of moderate length, broad and depressed, shaped like an inverted shovel, and ventrally flattened. Mouth of small to moderate size, inferior, lips thick and fleshy, cleft of mouth about horizontal. Eyes close to dorsal profile of head, quite close together, cheeks expanding broadly below eyes, so that the rather small eyes are directed dorsolaterally. Interorbital flat or slightly concave. The preopercular margin slightly crenulated, nonspinous, except in very small juveniles. Open pores on head as shown in Fig. 10. Lateral line simple, along mid-lateral trunk. Nostrils large, slightly tubular, anterior and posterior nostrils on each side close together. Isthmus narrow with gill openings extensive, opercular membranes from each side confluent, forming a free flap across isthmus. Gill rakers short, stout and toothed. Teeth on premaxillae and dentaries villiform with outer row conspicuously enlarged as canines, especially on premaxillae; a broad V-shaped band of teeth on vomer and a small patch on the head of each palatine, close to the vomerine teeth; tooth plates on pharyngobranchials and on fifth ceratobranchials. Three moderately long pyloric caeca. Scales moderately large, ciliated, extending forwards to nape, but absent from head and anterior two-thirds of belly; very small scales present on pectoral fin bases and, covering proximal half of caudal fin; scales on fins and belly cycloid. About 50 scale$ in lateral line.

12 McDowALL-Cheimarrichthys, the torrent fish 209 A B 0\ ',',':: :......;.' ~ : :. :i~.. ~.. FIG. 1O.-Disposition of cephalic laterosensory pores in Cheimarrichthys /osteri, left side of head; A. Dorsal aspect; B. Ventral aspect. Dorsal fin long, origin at about mid-abdomen, preceded by several very short, stout, pungent spines, Ijach with a discrete but rudimentary fin membrane. Fin proper with a longer, less stout spine, followed by branched rays, longest anteriorly becoming progressively shorter posteriorly, distat margin of fin smooth and about straight, high anteriorly, low posteriorly. Anal fin similar in form but lacking the short pungent spines anteriorly; fin membrane thick and fleshy, distal margin of fin strongly crenulate. Dorsal and anal fins terminate at about the same level on trunk. Caudal fin emarginate to truncated, depth of fin somewhat less than body depth, fin moderately long and fleshy, fin tips somewhat rounded. Upper pectoral fin insertion a little below mid-latedal trunk, fin broad based, upper rays short, lower rays up to twice length of upper ones, fin increasingly thickened and stiffened ventrally. Pelvic fins jugular and broadly separated from each other, fin broad based and semi-circular to ovoid in shape, fin projecting ventrolaterally to laterally below pectoral fins, thick and stiff, anterior ray much shorter than those following, the fourth of the five rays the longest. The sexes are similar in external morphology, although the abdominal wall of the male is thicker than that of the female. COLORATION In preservative, C. /osteri is grey to grey-brown, dark dorsally, paler laterally; ventrally it is a creamish-white. The head is dark dorsally with a darker stripe obliquely across the head, passing forwards and downwards ihrough the eyes; the lips are paler, the lateral margins and ventral surface of the head unpigmented. Along the trunk there are four dark bands that slope obliquely and irregularly forwards and downwards. The first of these bands begins at about the front of the dorsal fin and passes forwards behind the pectoral fin, the second extends from about the anterior third of the dorsal fin towards the vent, the third from near the hind end of the dorsal fin towards the posterior third of the anal fin and the last band crosses the caudal peduncle. The bands are caused by a gradual posteriad increase in the intensity of pigmentation, which reaches a peak density and abruptly fades, and is followed by a further increase in density. There is a dark vertical band across the base of the caudal fin. In the dorsal and anal fins there are series of dark pigment patches along the fin rays, and in the pectoral and caudal fins there are irregular bands of pigment

13 210 JOURNAL of THE ROYAL SOCIETY OF NEW ZEALAND across the fins. The pelvic fins have similar, if indistinct, bands across the anterior four fin rays, but the fin in the vicinity of the fifth ray is unpigmented. In life, C. fosteri is brown-black dorsally, paling to a dull orange-buff ventrally. Quite strong, orange to red coloration is commonly present in large adults, especially in the anterior and distal third of the dorsal fin, the anal fin, the lower lobe of the caudal fin, the lower third of the pectoral fin and also in the pelvic fins. This coloration extends onto the ventral surface of the belly in the most strongly pigmented individuals. SIZE Phillipps (1929) reports torrent fish growing to 6-8 inches long ( mm). The largest I have seen was 161 mm total length. C. fosteri commonly grows to about 100 mm, and fish more than about 125 mm long are not common. GROWTH Before differences between populations of C. fosteri can be examined, it is important to establish variations in body form with growth, because of differences between samples in their size distributions. To establish whether growth is iso- or allometric, a series of measurements was taken from 59 fishes from the Waikanae River, covering as complete as possible a range of sizes ( mm total length)..j: m» 0 '- U"l -I» Z 0» ;;0 0 :- ~ ~ : :. : ~--~--~~--~--~--~~~~--~--~~--~--~~~~~~--~ STANDARD LENGTH (mm) FIG. l1.-relationship between standard length and head length/standard length ratio in Cheimarrichthys fosteri from Waikanae River. With increase in size, the head of C. fosteri becomes relatively shorter (Fig. 11), and, perhaps partly as a result of this, the predorsal, preanal and prepelvic dimensions also decline relative to standard length. The pectoral-pelvic and pelvic-anal dimensions show a relative increase in length, and this induces a decrease in the pectoral fin/pectoral-anal length and pelvic fin/pelvic-anal length ratios. The head also becomes narrower and deeper, the interorbital narrower, the snout longer and the gape broader, relative to head length; but these differences are not striking. Although no dramatic changes in body form are evident, either from visual comparisons or from measurement, the fish nevertheless changes in form in minor ways as it grows so that comparisons of samples from different localities, comprising fishes of differing size distributions, could lead to faulty interpretations of geographic variation in this species. VARIATION AND POPULATION DIFFERENCES C. fosteri is widespread in New Zealand and could, perhaps, be expected to exhibit regional or local variations in some taxonomic characters. Measurements from three widely separated populations (Whanarua Bay, East Cape, 37 20'S latitude; Waikanae River, Wellington Province, 40 10'S; Ashley River, Canterbury Province, 45 15'S) failed to reveal any differences in taxonomic importance (Table 1). Meristic characters were also studied and the data revealed that caudal and pelvic fin ray, branchiostegal and vertebral numbers are almost constant. Variation in the number of dorsal, anal and pectoral fin rays was broader but no pattern of variation is evident (Table 2). There are no differences which indicate separate taxonomic status for any population or group of populations. HABITAT C. fosteri is found most commonly in unstable shingly rivers, mually larger rivers with broad open beds. It occurs in the tumbling, broken water of the swiftest rapids and seems to Iiv~ on the bottom amongst the rocks. Its habitat is very similar to that of Philypnodon

14 TABLE 1.-vanation 1Il body proportions of Cheimarrichthys fosteri (figures given as percentages of denominator in ratio). Whanarua Bay Str. Waikanae R. Ashley R. Harper R. Mean Min. Mean Max. Min. Mean Max. Min. Mean Max. #1 #2 TL/SL BW/SL IG BDV/SL :( ~ LCP/SL Cl DCP/LCP ~ PreD/SL ~ > PreD/PreA t"' t"' LDB/SL I MLD/LDB P"' LAB/SL ~ MLA/LAB i S!>l Pee/PecAn ::; Pel/Pel An CS P"' PrePel/SL P"' PecAn/SL '< v'" PelAn/SL HL/SL ;; HW/HL '" 0" HD/HL SnL/HL '" ~ POHL/HL " ~ IOW/HL '"' ;:s- ED/HL LUJ/HL LLJ/HL jl7 34, WG/HL No. measured 13 j r-:>

15 212 JOURNAL OF THE ROYAL SOCIETY OF NEW ZEALAND hubbsi Stokell and the two species are often taken together. A brief look at stomach contents showed that the torrent fish feeds on stream bottom invertebrates; Simuliidae, Chironomidae, Trichoptera and Plecoptera were observed. C. fosteri appears to be well adapted for its habitat; the depressed head, stout body with heavy ossification, and the thick, fleshy fins with very strong fin rays, seem well suited to life in very swift water. The dorsolateral eyes, and ventral mouth, the very well-developed nostrils and the numerous laterosensory pores, especially on the lateral and ventral surfaces of the head suggest that C. fosteri may locate its food in part, by olfaction and detection of movement through the latero-sensory system. LIFE HISTORY Little is known about the life history of C. fosteri. Stokell (1955) found that "Mature specimens on the point of spawning are taken in February and March", and Mair (1880) found them to be full of ova in February. Phillipps (1929), however, reported that spawning fish may be taken in June, July, December, January and February. Phillipps thought that there might be summer and winter spawnings. I have found fish with gonads much enlarged causing gross swelling of the abdomen from January through to April, although by March most adults have their gonads reduced to a slender thread. The data are limited but suggest a summer spawning. The eggs of C. fosteri are very small and numerous. They measure about 0.6 mm diameter; Phillipps (1929) reported that an adult 83 mm long contained 12,546 ova, and estimated 30,000 in one 150 mm long. Nothing is known about spawning habitat, spawning behaviour, whereabouts of the eggs during development, or larval and early juvenile life. Phillipps (1929) suggested that the torrent fish may make its way downstream,tb spawn, but the occurrence of mature adults many kilometres upstream, e.g., in the Wanganui River near Taumarunui, suggest that this is improbable. In April, 1970, I obtained four samples of C. fosteri in the Waikanae River system, near Wellington, at distances of 12.0, 10.6, 5.6, and 2.8 km upstream from the sea. The presence of fully ripe adults in the most upstream localities may also suggest that a downstream spawning migration is unlikely. From their very small size, I suspect that the eggs are probably washed downstream to the sea before hatching, and are planktonic. Occasionally, small C. fosteri, about 25 mm long, TABLE 2.-Meristic variation in Cheimarrichthys fosteri Dorsal fin IV V VI Whanarua Waikanae Ashley 6 5 Harper 2 Anal fin I 14 Whanarua 3 Waikanae Ashley 1 Harper 2 Pectoral fin 14 Whanarua Waikanae Ashley Harper Vertebrae 31 Whanarua 3 Waikanae 5 Ashley 4 Harper Gill rakers 2-7 Whanarua Waikanae 2 Ashley 3 Harper II Caudal fin Pelvic fin Branchiostegals

16 McDoWALL-Cheimarrichthys, the torrent fish 213 are captured from the sea, and small ones are sometimes caught in rivers near their mouths. At this stage they resemble the larger adults in both form and coloration. Study of the size distribution of the four samples from the Waikanae River system mentioned above shows a progressive upstream decline in small fish (Fig. 12). This is not A -n :::ti m 0 C m Zl () -< 5 c B 10 5 o STANDARD LENGTH (mm) FIG. 12.-Size distribution of Cheimarrichthys fosteri at four sites in Waikanae River, and tributary, April A. Ngatiawa Stream 12 km from sea, 24 fish, mean length 96 mm (indicated by arrow); B. Waikanae River 10.6 km from sea, 48 fish, mean length 100 mm; C. Waikanae River 5.6 km from sea, 65 fish, mean length 79 mm; D. Waikanae River 2.8 km from sea, 61 fish, mean length 86 mm. especially clear from the sample average, partiy because of the small size of the samples, but it is clear that the small fish are most abundant lower downstream. The same phenomenon was observed in Gobiomorphus huttoni (Ogilby) in which species the larval life is similarly marine (McDowall, 1965). In both G. huttoni and C. fosteri the exclusion of juveniles and small adults from upstream localities can be attributed to the young fish coming into the river from the sea, taking time to penetrate to upstream localities. HARPER RIVER POPULATION Occasionally specimens of C. /osteri are collected from the Harper River, above Lake Coleridge. Two large adults were collected there in December, Originally the Harper River was joined by the Wilberforce River and the combined flows joined the Rakaia. Lake Coleridge naturally drained into the Harper River via a small stream. However, beginning about 1911 (Dollimore, 1962) a hydro-electric station was constructed at Lake Coleridge, the waters of the Harper River were diverted into Lake Coleridge through a diversion race, and overflow from Lake Coleridge was fed through tunnels to a hydro-electric station on the Rakaia River (Fig. 13). Thus the bed of the Harper River, from the diversion race to the original confluence with the Wilberforce, and the stream which naturally drained Lake Coleridge, are now usually dry. Since C. fosteri is thought to be diadromous, it is somewhat surprising that C. fosteri is collected from the Harper River above the Lake Coleridge diversion, and also from the diversion race. It seems improbable that the fish can swim up the tunnels from the Hydro station into Lake Coleridge; if they could, they would still have to migrate up through Lake Coleridge and the diversion race to get to the Harper River. The presence of C. fosteri in tributaries of Lake Wairarapa does show that the young fish will migrate up through lakes to reach their rapid-water shingle stream habitats. So Lake Coleridge itself would probably offer no barrier to migration. Occasionally, only when the Harper River is in flood and when storage in Lake Coleridge is at its peak, the Harper River flows down its natural course. It is possible that in such circumstances, a few fish make their way up the Harper River from the Rakaia. An alternative possibility is that a lacustrine population of C. fosteri is established in Lake Coleridge, but such a population would be unique (unless the fish in Lake Wairarapa tributaries are similar in character). Study of the only two specimens available from the upper Harper River shows that they are similar in all examined characters to normal sea S"0in~ fishes a,pd th1,ls ~ives no support for a $"enetically i~olatecl lakt;-li)tii~ecl stoc;:k, Th\;

17 214 JOURNAL OF THE ROYAL SOCIETY OF NEW ZEALAND rareness of the species in the Harper River also gives no support for a self-sustaining lake stock. It seems most probable that the fish reach the Harper River from the Rakaia during floods when water is allowed to flow down the natural river course which is usually dry. : fi~ods when lak~.. ~HARPER....,. IS!.'-!.IL.::.~ I RIVER.. dry except during I.. t :::': /-;.<-;:.::':::-~ ::K~""'\ HARPER RIVER DIVERSION STREAM (natural lake outlet) (into lake).' WILBERFORCE RIVER LAKE COLERIDGE '" '.' OUTLET (tol.coleridge..' hydro-electric station, FIG. 13.-Map of Lake Coleridge, Harper and Wilberforce Rivers. DISTRIBUTION Stokell (1955) predicted that C. fosteri " will be found to be widely distributed", and this has proved to be true. It has the typical distribution of diadromous freshwater fishes in New Zealand. Although there are gaps in the known distribution, C. fosteri occurs throughout New Zealand in rivers it can reach from the sea, being limited only in inland penetration. Most localities are coastal, although C. fosteri has been collected from the Wanganui River at Taumarunui, about 250 km upstream from the sea. Gaps in the known range of C. fosteri are probably due, in part, to lack of sampling. Stokell (1955) reported it from Taranaki, but gave no precise locality, and this is the only Taranaki record known to me. It has not been recorded from Stewart Island, probably because of a lack of collecting. Skrzynski (1967) failed to find it on the Chatham Islands and suggested that suitable habitats are not present. C. fosteri is known from the following localities: Mangatete Stream, near Kaingaroa (Fig. 14: 1); Te Awa Puka Str., tributary of Oruru R. (2); tributary of Herekino R. (3); Mangamuka R. (4); Waima R. (5); Waiomio Str., trib. of Kawakawa R. (6); Kaihu R., trib. of Wairoa R. (7); Makarau R. (8); stream just north of Coromandel (9); trib. of Waiwawa R. (10); Kaueraunga R. (11); Matatoki Str., Waihou R. trib. (12); Pikowai Str. (13); Raroa Str. Waimana R. trib., (14); and Owhakatoro Str. (15), Whakatane R. system; Wairata and Omakora Str., Waioeka R. system (16); Whanarua Bay Str. (17); stream south-west of Whangaparaoa ( 18) ; trib. of Whangaparaoa at Waikura ( 19) ;

18 McDoWALL-Cheimarrichthys, the torrent fish " SO :: 49 S8 Sl FIG. H.-Distribution of Cheimarrichthys fosteri (numbers in figures) as indicated in text (pp. 214, 216).

19 216 JOURNAL OF THE ROYAL SOCIETY OF NEW ZEALAND Mangatote Str., trib. of Waiapu R. (20); trib. of Mangapoike Str., Wairoa System (21); trib. of Waipa R. near Te Pahu, Waikato system (22); Wanganui R. at Taumarunui (23), and trib. at Jerusalem (24); Waipawa R., trib. of Tukituki R. (25); Whakataki R. (26), Mikimiki Str. (27), Tauherenikau R. (28) and Tauanui R. (29), Ruamahanga system; Burlings Str., trib. L. Wairarapa (30); Otaki R. (31); Waikanae R. (32); Horokiwi Str. (33); Hutt R. (34); Wakamarina R., trib. of Pelorus R. (35); Waimea R. (36); Oparara Str. (37); Little Totara R. (38); Punekaiki R. (39); 13 Mile Str. (40); Kawhaka R., Arahura system (41); Hokitika R. (42); Waiau R. (43); Hurunui R. (44); Ashley R. (45); Purau Str. (46); Harper R., trib. of Rakaia R. (47); and Rakaia R. (48); Ashburton R. (49); Hinds R. (50); Rangitata R. (51); Wahapo R., Okarito system (52); Waikukupa R. (53); Manakaiau-ua R. (54); Haast R. (55); Turnbull R. (56); Waiatoto R. (57); Arawata R. (58); Awarua R. (59); Otaio R. (60); Waihau R. (61); Waianakarua R. (62); Waitati R. (63); Waiau R. at Blackmount (64). ACKNOWLEDGMENTS I am grateful to Mr J. M. Moreland, National Museum, Wellington, for the loan of specimens in the museum collections, and to Dr P. H. J. Castle, Victoria University of Wellington, for reading and commenting on the manuscript. REFERENCES ALLEN, K. R The geography of New Zealand's freshwater fish. New Zealand Science Review 14(3): Freshwater fish. In Science in New Zealand, pp Reed, Wellington. BURNET, A. M. R., CRANFIELD, H. J., and BENZIE, V. L The freshwater fishes. In Knox, G. A. (Ed.). The natural history of Canterbury, pp Reed, Wellington, 620 pp. CANTWELL, G. E A revision of the genus Parapercis, family Mugiloididae. Pacific Science 18(3): 239-PO. DOLLIMORE, E. S The ;{ew Zealand Guide. Wise, Dunedin. 981 pp. GoSLINE, W. A The suborders of perciform fishes. Proceedings of the United States National Musuem 124: GREENWOOD, P. H., ROSEN, D. E., WEITZMAN, S. H., and MYERS, G. S Phyletic studies of teleostean fishes with a provisional classification of living forms. Bulletin of the American Museum of Natural History 131(4): HAAST, J. VON On Cheimarrichthys fosteri, a new genus belonging to the New Zealand freshwater fishes. Transactions and Proceedings of the New Zealand Institute 6: HEFFORD, A. E New Zealand fish and fisheries. In Falla, R. A. (Ed.). Handbook for New Zealand, pp Australian and New Zealand Association for the Advancement of Science, Wellington. HIROA, TE R The Maori rraft of netting. Transactions and Proceedings of the New Zealand Institute 56: HOPKINS, C. L. and McDOWALL, R. M A review of present knowledge of fishes in New Zealand freshwaters. Proceedings of the New Zealand Water Conference 1970, part 1: HUTTON, F. W Index faunae Novae Zealandiae. Dulau, London. 372 pp. LANE, E. D The osteology of Cheimarrichthys fosteri Haast (Pisces, Percomorphi). Transactions of the Royal Society of New Zealand, Zoology 6(20): MCCULLOCH, A. R. and PHILLIPPS, W. J Notes on New Zealand fishes. Records of the Australian Museum 14(1): McDOWALL, R. M. 1964a. A bibliography of the indigenous freshwater fishes of New Zealand. Transactions of the Royal Society of New Zealand, Zoology 5(1): b. The affinities and derivation of the New Zealand freshwater fish fauna. Tuatara 12(2): Studies on the biology of the red-finned bully Gobiomorphus huttoni (Ogilby). II. Breeding and life history. Transactions of the Royal Society of New Zealand, Zoology 5 (14): A guide to the identification of New Zealand freshwater fishes. Tuatara 14(2): Galaxias maculatus (Jenyns), the New Zealand whitebait. Fisheries Research Bulletin, New Zealand Marine Department 2: McMILLAN, H. M An addition to the knowledge of the fish Retropinna anisodon Stokell (Retropinnidae). Transactions of the Royal Society of New Zeqland, Zoology 1 (10):

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