Memoirs OF THE Queensland Museum Brisbane June, 1978 Volume 18 Part 2 Volume 18 Part 2 Memoirs OF THE Queensland Museum Published by Order of the Board Printed in Australia by The Courier-Mail Printing Service, Campbell Street. Bowen Hills Q, 4006. THE QUEENSLAND MUSEUM Gregory Terrace Fortitude Valley Brisbane Qld 4006 Telephone: 52 2716-8 TRUSTEES J. C. H. Gill, B.A., LL.B. (Chairman) I. G. Morris, C.M.G. (Vice Chairman) Emeritus Professor S. A. Prentice, M.E.E., B.Sc. Professor J. M. Thomson. D.Sc. D. M. Traves, O.B.E., B.Sc. R. I. Harrison. M.B.E., E.D., B.Comm. A. F. Hinchy, A.A.U.Q. A. Bartholomai, M.Sc., Ph.D. {ex officio, Secretary) SCIENTIFIC STAFF Director: A. BARTHOLOMAI, M.Sc., Ph.D. Deputy Director: B. M. Campbell, M.Sc. (Editor) Senior Curators: E. C. Dahms, B.Agr.Sc., B.Sc., Curator of Insects Patricia Mather. Ph.D., D.Sc., Curator of Higher Invertebrates M. C. QUINNELL, B.A.(Hons), Curator of Anthropology/ Archaeology D. Robinson, B.Sc.(Hons), Ph.D., Curator of History and Technology Mary Wade, B.Sc.(Hons), Ph.D., Curator of Geology Curators: M. ARCHER, B.A., Ph.D., Curator of Mammals M. J. Bishop, M.A., Ph.D., Curator of Molluscs L. R. G. Cannon, B.Sc., Ph.D,, Curator of Lower Invertebrates Jeanette CovaCEVICH, B.A., Curator of Reptiles Valerie Davies, M.Sc., D.Phil., Curator of Arachnids R. G. Hardley, B.A., (Hons), Curator of Australian Ethnography G. J. Ingram, B.Sc., Curator of Amphibia R. J. McKay, B.A., Curator of Fishes R. W. Monroe, B.Sc., Curator of Crustacea I. G. Sanker, B.Sc., Dip. Ed., Curator of Industrial Technology P. P. Vernon, Ornithologist Mem. QdMus. 18(2): 127-43, pis. 23-7. [1978] THE MACROPODIDAE (MARSUPIALIA) FROM THE ALLINGHAM FORMATION, NORTHERN QUEENSLAND. RESULTS OF THE RAY E. LEMLEY EXPEDITIONS, PART 2. Alan Bartholomai Queensland Museum ABSTRACT The fossil Macropodidae from the Bluff Downs fauna of the Allingham Formation, northern Queensland, of lower Pliocene age, are described. Eight taxa have been recognized, although inconclusive evidence exists to suggest that additional taxa may be defined when broader collections become available. Potoroines are currently unknown and slhenurines are very poorly represented- Macropodines predominate. Four species are new. Troposodon bluffensis, Protemnodon snewini. Macropus (Prionotemnus) narada and M. iOsphranter) pavana. Two species, M. i?P.) dryas and T. minor are known also from the Chinchilla Sand, although the latter species has a wide temporal range being well represented in Pleistocene deposits. The occurrence none the less supports the previously ascribed late Pliocene age for the Chinchilla Sand. A preliminary statement on the fauna from the Allingham Formation and the definition of the Formation are provided in Archer and Wade (1976). The unit is regarded as being of lower Pliocene age because it is overlain in part by the Allensleigh ‘How' of the Nulla Basalt, dated radiometrically at between 4 and 4 5 M.y. This then represents a minimal age for the Formation, although from what is already known of the fauna, actual time of deposition was probably not much earlier. Continuing investigation of the deposits has been made possible through the support of Dr Ray E. Lemley as part of a joint Queensland Museum and South Dakota School of Mines project. Larger collections of macropodids have resulted, enabling the present study to be undertaken. Other groups will form the basis for additional studies. The deposits are considered to be of great importance and they represent only the second Pliocene, mammal-bearing site in Queensland. The reasonably precise minimal age and the known diversity of the fauna suggest that results will assume some significance in correlation of Australian lacustrine and terrestrial deposits. All measurements throughout are in milli- metres. STHENLRINAE Sthenurus sp. (Plate 23, fig. 1) Mati-.rim. Examined: F9104, partial, isolated right (breadth 12‘6). DISCUSSION; Only one fragmentary tooth of Stenurus Owen has been recovered from the Allingham Formation. Its morphology suggests that it is an ML Insufficient features are preserved for more than generic determination to be attempted. However, of species currently known, the tooth is believed to be from a form close to Sthenurus antiquus, described by Bartholomai (1963) from the late Pliocene Chinchilla Sand. Ornamentation and height of the lophs is similar, although in the Allingham specimen, the strength of the ridge which ascends into the median valley from the metaconc is considerably weaker than in the maxilla of S. antiquus figured by Bartholomai (1963, fig. 10). As in S. antiquus, the base of the crown is swollen posteriorly and the low midlink is positioned above the centre of the median valley. However, the strong ridge curving posterolingual- ly from the hypocone overlaps, rather than meets, the weaker ridge from the metacone. 128 MEMOIRS OF THE QUEENSLAND MUSEUM Although upper teeth are unknown in S. notabilis Bartholomai, also from the late Pliocene Chinchilla Sand (Bartholomai 1963). size and crown height would preclude this species from close relationships with the Allingham speci- men. Among Pleistocene species of Sthenurus reviewed by Bartholomai (1963) and Tedford (1966), or described by Merrilecs (1965, 1967), the Allingham specimen appears to be within the size range of S. areas De Vis, but has less pronounced accessory ridging. S. orientalis Tedford of comparable size, is known only from its lower dentition, while S. occidentalis Glauert, S. brownei Merrilees and S. gilli Merrilees are all smaller species. MACROPODINAE Troposodon minor (Owen, 1877) (Plate 23, figs. 2-4) Sthenurus minor Owen, 1877, pp. 352-61, pi. 37 , figs.' 1-3, pi. 38, figs. 1-4. Troposodon minor (Owen); Bartholomai, 1967, pp. 23-32, figs. 1-3 (synonymy). Matpriai. Examim-D: F9046, isolated left P^; F9047, isolated partial left M.; F9048. isolated partial left M 3 ; F9049, isolated left DP\ F9050. isolated right M,; F9051, isolated left DPh F9052, isolated partial right M’; F9053, isolated partial right M|. DISCUSSION: The genus Troposodon Bartholomai is currently represented in Australian Upper Cainozoic deposits by two species, T. minor (Owen) and T. kenti Campbell. Of these, T, minor is by far the most commonly recorded species, being present in sufficient numbers in collections from the Darling Downs deposits to enable Bartholomai (1967) to assess the populations statistically. Within the Darling Downs deposits, Barth- olomai (1967) has shown T. minor to be present in both the Pleistocene fluviatile deposits and the Chinchilla Sand ot late Pliocene age. Samples were compared using Student’s I Test and no significant size or morphological features were found to justify separation. Turnbull and Lundelius (1970) have also suggested that an isolated, partial lower molar. MNV-P26422, from the post-early Pliocene fossil soil deposits at Grange Burn, Hamilton, Victoria, may represent T. minor, a possibility which cannot be discounted on the basis of the description provided. T. kenti from the ?early Pleistocene Katipiri Sands, has been distinguished from T. minor by Campbell (1973) on the basis of its larger size, TABLE I: Measurements eor Troposodon minor (Owen, 1877) ' Specimen u Ml M. M 3 DP3 F9046 12-8x51 F9047 — — — x9-3 F9048 — — — — xlO-2 F9050 — 11-0x6-9 F9053 — — x7-3 — F9049 — — — — 10 - 2 x 8-1 F9051 — — — — 10-7x7-0 supported by the less ornamented nature of the apical shelves of the molar teeth and minor morphological differences in the premolars. Representation of material larger than T. minor in the Chinchilla Sand material, but insufficient for description, was noted by Bartholomai (1967). No additional material has as yet been recovered to support or refute the suggestions by Campbell (1973) that these specimens may represent T. kenti in the Chinchilla Sand, or that they may reflect extreme variation in T. minor. Although the sample available from the Allingham Formation is small, comprising only isolated teeth, a reasonable range of dental morphology is represented and some variation within the population is observable. Measur- ements for this material appear in Table 1. Slight size differences which are evident between teeth from the Allingham Formation and those from the Chinchilla Sand are attributable to the small sample size in the former and the fact that the Chinchilla Sand sample itself is a relatively small one for statistical purposes. Nevertheless, the Allingham sample falls within limits which one would expect for the Chinchilla Sand sample, based on large sample assessments for macropodines provided in Bartholomai (1971, 1973, 1975). Structurally, the Allingham sample generally presents coarser accessory ridging in anterior molars and in DP^ than is usual in Darling Downs specimens, but this feature is variably present in some individuals from the Darling Downs, including the Juvenile specimen figured by Batholomai (1967, fig. 2). No differences are therefore present which enable separation of the Allingham material from T. minor, although T. kenti is clearly excluded by both size and morphological considerations. This is of particular interest because of the age of the Allingham Formation, which indicates a minimal temporal spread for T. minor from at least the mid Pliocene to the late Pleistocene, the longest recognisable range for an Australian marsupial species. BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE 129 Troposodon bluffensis* sp. nov. (Plate 23, figs. 5, 7-10) Mattrim. Examined F9054. holotype, isolated right M**. adult, Allingham Formation of mid Pliocene age, Bluff Downs Station, Allingham Creek. In addition, F9055, an isolated left M 4 , F9056, a partial isolated left M‘*; F9057, an isolated right M 4 ; F9058, an isolated right P 2 and F9059, an isolated left P^ have been used in the description of the species. Specific Diagnosis: a small species of Troposodon. much smaller than T. minor: P^ small, with low longitudinal crest; upper molars lacking a forelink or with this present in very incipient form, and lacking a ridge from the paracone to the labial extremity of the anterior cingulum; midlink high with major contribution from near midpoint of melaloph crest. P^ small, low crowned, markedly L-shaped; lower molars with reduced accessory ridging on apical shelf and with base of crown swollen posteriorly without production of posterior cingulum. DESCRIPTION; Upper dentition known only from P^ and M**. P^ elongate, subcrcscentic in basal outline, with labial margin concave and lingual convex, broader posteriorly than anteriorly; longitudinal crest low, more elevated posteriorly than anteriorly, moderately concave iabially, transected by three, weak sets of vertical labial and lingual ridges, these becoming weaker posteriorly. Apex of paracone about one-fifth distance along crown from anterior limit. Hypocone moderately low, positioned posterolingual to metacone and connected to it by strong descending ridge; weak ridge curves posterolabially to meet extremely weak posterior ridge from metaconc near base of crown, without production of posterior fossette; strong anterior ridge from hypocone ascends to form well-defined lingual cingulum to point opposite paracone; cingulum sinuous in lingual view, converging only slightly anteriorly in occlusal view; lingual basin broad, smooth. Anterior ridge from paracone extending well anteriorly; base of crown slightly swollen anterolingually. small, subrectangular in basal outline, slightly constricted across median valley; lophs low, moderately bowed anteriorly, with melaloph narrower than prololoph. Anterior cingulum low. *Named for Bluff Downs Station, the property on which the Allingham Formation occurs. moderately broad, anteriorly broad convex, nearly flat in labial moiety, but ascending slightly iingually; forelink absent or present in very incipient form; labial margin of cingulum unconnected to paracone by accessory ridge; midlink high, strong, curving posterolabially from prolocone, then curving posteriorly abruptly to unite with strong ridge from near midpoint of crest of mctaloph; Junction marked by shallow cleft. Weak ridge ascends poslerolingually from paracone towards midlink, defining unornamented apical shelf; median valley shallowly U-shaped, transversely flaitcncd, with lingual extremity marked by low ridge; strong ridge curves posterolabially from hypocone towards base of crown; weaker ridge from metacone ascends to disappear below ridge from hypocone with production of posterolabial indentation; accessory ridges from hypocone and metacone ascend across posterior face of melaloph to unite near midline, defining unornamented apical shelf. Base of crown narrowly swollen posteriorly, this separated below by shallow groove. Lower dentition known only from Pt and M4. P 2 relatively small, subcrescentic in occlusal view, slightly broader posteriorly than anteriorly; longitudinal crest relatively low, markedly L-shaped, being flexed Iingually in its posterior extension. Anterior cuspid set posteriorly at about one-half distance along crown from anterior margin, with anterior ridge from cuspid angled anteriorly and marked by low cuspule about one-half distance to crown base; low, anterolabial cingulum developed from this cuspule to unite with labial ridge from anterior cuspid. Longitudin- al crest transected by two weak sets of vertical labial and lingual ridges; lingual extension of crest descends slightly to posterolingual cuspid, this having strong posterior ridge and stronger anterior ridge descending from it. Base of crown swollen Iingually and Iabially. M 4 small, subrectangular in basal outline, moderately strongly constricted across talonid basin; lophids low, moderately strongly convex posteriorly, with prololophid broader than hypolophid. Trigonid basin moderately broad, its length approximately equalling distance between lophids. Forclink high, strong, curving antero- lingually from protoconid across trigonid to point on anterior cingulum labiad to midline; anterior cingulum moderately high, broad, rounded anterolabially, more angular anterolingually, descending markedly Iabially and Iingually from forelink; strong accessory ridge curves anter- olabially from metaconid towards forelink. 130 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 2: Mhasuri MhNTS for Troposodon bluffensis SP. NOV. Specimen M'* P 2 M 4 F9054. holotype lI-7x8-5 F9056 — — x9-l — — F9055 — — — 12 - 0 x 8-2 F9057 — — — 12-7x8-2 F9058 1 0-6x5-] — — — F9059 — — 70x3-5 — defining apical shelf: weak accessory ridges present on apical shelf, descending anteriorly from near protolophid crest. Talonid basin narrowly U-shaped, descending slightly labially and lingually from midlink. Midlink strong, moderate- ly high, curving slightly anterolinguaily from hypoconid to near midpoint of posterior of protolophid; slight ridge curves towards midlink from enloconid without production of apical shelf. Base of crown swollen posteriorly but without production of posterior cingulum. DISCUSSION: Although at present known from a very small sample comprising only isolated cheek teeth, the material is so distinct both in its size and morphology as to conclusively justify its separation as a distinct species of Troposodon Bartholomai within the Allingham Formation. Compared with the Chinchilla Sand sample of T. minor (Owen) of late Pliocene age, the Allingham material falls well below the lower observed limits for size in equivalent teeth and, indeed, is well below the statistical limits for that population. The morphological distinction of molars in T. bluffensis lies in the greater simplicity of links and accessory ornamentation and in the more generalized basic structure overall. The Pt is more complex anteriorly and anterolabially than in T. minor, but it has more subdued ridging associated with the longitudinal crest. Ridges from the postcrolingual cuspid, especially the anterior ridge, are stronger and better-defined. The upper permanent premolar is less triangular in basal outline than in T. minor. That two species of Troposodon should be represented within the Bluff Downs deposits is not surprising, taking into account the macropodine representation within other fossil and recent faunas and the fact that the Allingham Formation rellects both Huviatile and lacustrine sedimenta- tion (Archer and Wade 1976) with attendant wide sampling of endemic and derived specimens. It is considered likely that in view of its more generalized nature. T. bluffensis stands closer to the ancestral form from which Troposodon was derived than do either 7. minor or 7. kenti. In addition to the material here referred to 7. bluffensis. there exists an isolated right P3, F9060, figured in Plate 23, fig. 8. which may belong with this species. The tooth is clearly of the Troposodon type, having the posterior extension of its longitudinal crest acutely angled lingually, giving the tooth a marked L-shaped appearance in occlusal view. It differs from P3 in 7. minor in being shorter (10- 1x5-2) and in having its anterior margin near vertical. Although the longitudinal crest is lower anteriorly than posteriorly, the crown is relatively high in comparison with 7. minor and with the P-, referred to 7. bluffensis. While it is considered highly unlikely that a further species of Troposodon is represented in the Allingham Formation, the specimen is maintained as doubtfully referred to 7. bluffensis until more complete mandibular material is collected to confirm its taxonomic position. Petrogale sp. (Plate 23, fig. 6) Matf.rial Examinrd: F7794, isolated left Mt; F7795, isolated right M 3 ; F9043, isolated left M 2 : F9044, isolated left M 4 ; F9045, isolated left M-,. DISCUSSION: The isolated molars here referred to Petrogale sp. were regarded by Archer in Archer and Wade (1976) as c.f. Thylogale sp., but were not figured. Petrogale and Thylogale are difficult to separate on the basis of isolated cheek teeth, particularly molars. However, it appears that Petrogale molars are more like those in Macropus than are those in Thylogale. In Thylogale, the lophids are very narrow longitudin- ally, even towards the crown base, while transversely, the lophid crests are more rectilinear. The midlinks are angled more anterolinguaily from the hypoconid in Thylogale. In size, the referred sample is similar to P. penicillata, but insufficient is known of the morphology of the Allingham sample to justify more discrete treatment at this tine. The isolated right DP\ F7785, mentioned by Archer in Archer and Wade (1976) as being difficult to distinguish from corresponding teeth in Thylogale (e.g. 7. stigmatica) does show greater morphological similarity to Thylogale than to Petrogale. Rather than promote additional speculation, its position is left doubtful until more BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE 131 TABLE 3: Measurements for Peirogale sp. Specimen M 2 M 3 M 4 F7794 6-2x41 F7795 — 7-lx4-l F9043 6-3x4- 1 F9044 — — 8-2x5-2 F9045 6-9x4 1 — — adequate samples become available. Thylogale is known from post-early Pliocene deposits at Grange Burn, Hamilton, Victoria (Turnbull and Lundelius 1970). Protemnodon snewini* sp. nov. (Plate 24. figs. 1-4; Plate 25, fig. 1) Protemnodon sp. Archer 1976 (partim), pp. 390-1, pi. 57a. Materiai Examined: F9061, holotype, right mandibular ramus with I|, P 3 -M 4 , adult, Allingham Formation of mid Pliocene age, Bluff Downs Station, Allingham Creek. This specimen is possibly associated with the partial skull, F9074, and a partial, disarticulated skeleton, F9075, all located close to one another at the same stratigraphic level within the deposits. In addition, F7810, a partial right maxilla with M'-M^ shattered M^-M"*, adult; F7809. an isolated left M^; F9067. an isolated, partial right M 4 ; F7788, an isolated left P^; F9068. an isolated right M 3 , F781 1, an isolated right M**; F9069, an isolated right M’; F9070, a partial, isolated left Pj; F9071, an isolated left M^ F9072, an isolated left P-; F9073, an i.solated right P-; F7786, a right P^, aged; F9062, an isolated right P^; F9063, an isolated partial right M 4 ; F9064, an isolated right M|; F9065, an isolated left M 3 : F9066, isolated partial left M^; F7824, an isolated right P^; have been used in the description of the species. Specific Diagnosis: A small species, generally smaller than P. anak. Mandible comparatively very shallow with an elongate symphysis ascending at about 10 ^ to base of mandible; lateral groove extends to below posterior root M 3 ; menial foramen close to diastemal crest about one-quarter distance along diastema from anterior root P 3 . Condyle reduced transversely. Ii shallow. Cheek tooth row slightly bowed labially in occlusal view. P 3 narrow, elongate, slightly longer than M 4 ; crown low, with straight longitudinal crest transected by four sets of slightly anteriorly directed labial and lingual. * Named for Mr W. Snewin who, with Mr J. Barrett, discovered the Allingham site. ridges. Lower molars slightly constricted across talonid basin, w'ith lophids low, links labiad to mid-line; lingual moiety of trigonid unswollen; anterior cingulum broad, extended anterolingually into distinct, subangular corner to crown; talonid broadly U-shaped; posterior cingulum poorly developed. P- elongate, with longitudinal crest low, transected by three or four sets of vertical labial and lingual ridges; lingual cingulum well developed, with shallow lingual basin narrowing slightly anteriorly. P^ narrow, elongate, slightly constricted basally at posterior one-third; longitudinal crest nearly straight, low, transected by three or four sets of vertical labial and lingual ridges; lingual cingulum very low, continuing to anterior of crown base, sinuous; lingual basin very narrow, shallow. Upper molars with low lophs, unconstricted, or slightly constricted across median valley in posterior molars; strong ridge ascends from paracone to limit of broad, flat, anterior cingulum; midlink very low; posterior ridge from paracone very weak; metaloph relatively broad in M*^- Description-. Mandible very shallow, relatively thick; base of symphysis ascending anteriorly at an angle of approximately 10 “ to base of mandible; symphysis elongate, shallow, not ankylosed, rugose; geniophyal pit shallow, near posterior symphysial limit; diastema elongate with diastema! crest angular posteriorly, more rounded anteriorly; ventral margin of ramus rounded between symphysis and extremely weak diagastric ridge and process. Mental foramen small, ovate, near diastemal crest, about one-quarter distance along crest from anterior root P 3 . Ramus with shallow lateral groove extending posteroventrally to below posterior root M 3 , somewhat disrupted by roots of P 3 : posterior of groove approximately mid-way between alveolar margin and base of ramus. Diagastric process separated from base of angle by very shallow post-diagastric sulcus, bounded above by shallow diagastric fossa. This fossa separated above from shallow, broad depression opening posteriorly into pterygoid fossa. Post-alveolar shelf short, with shelf angle reasonably well defined, leading to post-alveolar ridge ascending to disappear rapidly on medial wall of coronoid process, above large mandibular foramen. Masseteric crest low, approximately at level of alveolar margin; masseteric foramen moderately large with deep masseteric fossa. Angle of mandible broadly inflected, produced posteriorly and dorsally into crest. Anterior of coronoid process ascends posterodorsally at about 132 MEMOIRS OF THE QUEENSLAND MUSEUM 13 ° from vertical. Condyle very reduced, longer than broad, planar transversely and gently convex longitudinally, supported posteriorly by narrow shelf. I) elongate, deeply rooted; slightly curved in lateral view, markedly curved in occlusal view, developing subhorizontal transverse facet of wear, but gently convex wear facet longitudinally with upper incisors; mesial wear facet present along ventral margin by approximation with other lower incisor; root compressed, oval in section; crown subquadrantal in section, tapering and blade-like anteriorly, enamelled laterally, this produced dorsolabially and ventrolingually into flanges; crown slightly enamelled ventromcsially. P3 relatively elongate, robust, moderately broad; crown subovate in basal outline with longitudinal crest secant, nearly straight but strongly flexed lingually in its posterior extension; crest transected by two sets of near vertical labial and lingual ridges with production of cuspules at crest; strength of ridges and cuspules decreases posteriorly; anterior cuspid of crest well defined; anterior of crown straight, with ridge from anterior cuspid curving lingually towards base of crown; lingual base of crown swollen; broad, short, vertical, basal ridge present posterolabially. DP3 is not preserved. P3 elongate, narrow, deeply rooted; crown subovate in basal outline with longitudinal crest secant, slightly sinuous, and slightly flexed lingually in its posterior extension; crest transected by four sets of slightly anteriorly directed labial and lingual ridges, with production of cuspules at crest; strength of ridges and cuspules decreases posteriorly; anterior cuspid of crest well defined, but posterior cuspid less well developed; anterior of crown straight, near vertical extending well anterior to anterior root; labial base of crown slightly swollen, but not as strongly as posterior moiety of lingual base. Mi M 2 M 2 F9105, holotype F9106 F9107 9-lx7-2 90x7-1 10-lx7'9 — 12-5x9-0 Macropus (?Prionotemnus) dryas (De Vis, 1895) (Plate 26, fig. 6; Plate 27, figs 1-3) Halmaturus dryas De Vis, 1895, pp. 109-11, pi. 17, figs. 11-5. Macropus (Prionotemnus) dryas (De Vis): Barth- olomai, 1975, pp. 229-32, pi. 25, figs. 1-4 (synonymy). Macropus sp. c.f. M. dryas (De Vis): Archer in Archer and Wade, 1976, p.391, pi. 57, figs. c-d. Maiiriai Examined: F7823, partial right mandibular ramus with P^-Mj. P 3 excavated, juvenile (figd. Archer. 1976, pi. 57, figs. c-d). F9078, isolated right M^; F9079, isolated right M^; F9080, isolated partial left M 2 : F908I, isolated left Mt’ F9082, isolated right M^; F9083, partial isolated left M^ F9084. isolated left Ph F7828, isolated left Ph F7780, partial left maxilla with M--M^, juvenile; F7781, isolated left Ph F9085. partial left mandibular ramus with M 1 -M 3 , aged; F9086, partial right mandibular ramus with P 3 -M 4 ; adult; F9087. partial right mandibular ramus with M 2 , juvenile; F9088. partial left mandibular ramus with base of I), P 3 -M 1 , juvenile; F9089, partial left mandibular ramus with M 3 -M 4 , adult; F9090, isolated left M^; F909I, isolated right P 2 ; F9092, isolated partial right M^; F9093, isolated partial left M^; F9094, isolated left M 2 ; F9096, isolated right DP 3 ; F9097. isolated right Py. F9098, isolated partial right M 2 : F9103, isolated left M'; F9095, isolated partial left DP 3 ; F9099, isolated right M^: F9100, isolated partial left M'; F9I01, isolated partial right DP 3 ; F9102, isolated left P 3 ; F7790, isolated right P^; F7782, partial right mandibular ramus with M] M 3 , juvenile; F7783, isolated right P 3 ; F7791 isolated left P^. DISCUSSION: The sample of Macropus (? Prionotemnus) dryas (De Vis) from the 138 MEMOIRS OF THE QUEENSLAND MUSEUM Allingham Formation is numerically the largest among macropodids from the deposits, suggesting that this species was the most abundant grazing herbivore in the fauna. Bartholomai (1975) revised M. dryas, previously recorded only from the late Pliocene Chinchilla Sand in the western Darling Downs, where the species was well represented but not dominant within the assemblage present. Observed measurements for cheek teeth in the Allingham sample often fall towards the lower observed limits or even outside the lower limits of comparable teeth in the Chinchilla Sand material. No other morphological differences are evident which might be considered significant in the separation of the Allingham Formation sample from that in the Chinchilla Sand. With the larger sample now available, the possible structural distinctions noted by Archer in Archer and Wade (1976) arc considered to be within the range of variation for M. dryas. or are likely to be encountered in the species, bearing in mind the variation exhibited by modern species of Macropus (Bartholomai 1971). Two specimens, F7790 and F7791, have been referred to M. dryas. both of which arc P-. This tooth was not present in the available Chinchilla Sand material at the time of Bartholomai’s ( 1 975) revision. These possess a single vertical set of labial and lingual ridges transecting the TABLE 8: MFAStREMnNTS for Mandibles or Macropus {?Prionotemnus} drvas (De Vis. 1895) Specimen P2 DP 3 P 3 M, M 2 M 3 M 4 F7823 8-5x3-8 8-8x5-0 11-3x4-0 9-9x6-5 F9078 — — — — ■ 12-0x7-2 — — F9079 — — — — 11-6x7-0 — — F9080 — — — — — x8-3 — — F9081 — — — — 12-5x80 — — F9082 — — — — 11-9x7-1 — — F9085 — — — 8-8x6-3 10-0x7-6 12-2x8-2 — F9086 — — 11-3x3-9 9-lx5-7 1 1 -Ox— 12-7x8-0 13-8x8-1 F9087 — — — — 11 -5x— — — F9088 — — 10-5x3-6 8-7x5-9 — — — F9089 — — — — — 12-4x8-2 13-4x8-3 F9090 — — — — — 13-0x8-4 — F9091 7-6x3-9 — — — — — — F9092 — — — — — — x8-3 — F9093 — — — — — x7-9 — — F9094 — — — — 12-2x7-5 — — F9096 — 8-6x4-3 — — — — — F9097 — — 11-7x4-2 — — — — F9095 — 8-0x4-4 — — — — — F9099 — — — — 11-9x7-7 — — F9101 — — x4-5 — — — F9102 — — 11-4x3-9 — — — — F7782 — — — — 11 -7x— 13-6x7-7 — F7783 — 11-2x4-0 — — — — TABLE 9: Measurements for Maxillae of Macropus 1895) (?Prionolemnus} dryas (Dl: Vis. Specimen P2 DP3 P3 M' M2 M3 M4 F9084 _ 13-7x6-0 F7828 — — 13-8x6-4 — F7780 — — — — 11-1x9-3 11-8x9-4 F7781 ■ — — 13-3x6-3 — F9103 — — — 10-3x7-9 F9i00 — — — x8-2 F7790 8-2x5-4 — F77gi 8-3x5-l — __ — — — — BARTHOLOMAI: ALLINGHAM FORMATION MACROPODIDAE 139 longitudinal crest, have a low hypocone and a variably present lingual cingulum. This structure can be reasonably W'cll developed compared with the situation in where it is represented only by a series of low' tubercles closely adpressed to the base of the longitudinal crest. Macropus (Osphranter) pavana sp. nov. (Plate 26, figs. 1-5) Mathriai. Examined: F9108, holotype, partial left maxilla with DP-^-M', P-^ missing, juvenile, Allingham Formation of mid Pliocene age. Bluff Downs Station, Allingham Creek. In addition, F9109, a partial left mandibular ramus with M 2 -M 4 . adult; F9110, isolated right M-; F9111, isolated right M^; F9112, isolated right M'; F9113, isolated right Pb F9il4, isolated right P^; F9115 isolated right M-; F9116, isolated left P^; F9117, isolated left M^; F7773, isolated right M^; have been used in the description of the species. Specific Diagnosis-. A relatively small species with deep ramus, relatively narrow below anterior cheek teeth; P 3 relatively elongate with trenchant longitudinal crest transected medially by single set of vertical labial and lingual ridges and with crest slightly curving lingually posteriorly. Lower molars with midlink well labiad to midline; hypolophid with variable oblique groove. DP^ with high secant ridge from paracone to anterolabial limit of anterior cingulum; forelink strong; accessory link present mid-way between midlink and labial margin of median valley. P-^ with longitudinal crest slightly cleft at anterior one-third and w'ith single medial set of vertical labial and lingual ridges transecting crest; hypocone very low; posterolingual fossette and lingual cingulum very reduced, cingulum degener- ating to series of tubercles anteriorly. Molars with relatively strong forelinks; anterior molars with slight accessory link mid-way between midlink and labial margin of median valley; this is variably absent in posterior molars. Description: Upper incisors, P-, and not preserved. DP^ molariform, subovate in basal outline, slightly constricted across median valley, with metaloph much broader than protoioph. Lophs high, slightly convex anteriorly. Anterior cin- gulum moderately high, narrow, short, with well-defined forelink well linguad to mid-line from base of protoioph to cingulum; cingulum near planar, labiad to forelink, ascending at very high angle lingually. Strong high ridge ascends anteriorly from paracone to anterolabial margin of cingulum as secant crest. Well-defined, high, strong ridge ascends posterolabially from proloconc across median valley as midlink, uniting with ridge from near mid-point of metaloph, below median valley; relatively strong ridge ascends posteriorly from paracone towards median valley, producing slight accessory link across floor of valley mid-way between midlink and labial margin. Valley narrowly V-shaped labially and lingually, near planar transversely. Strong ridge ascends posteriorly from hypocone, curving labially to near poslerolabial margin of crown, uniting with slight posterior ridge from metacone; fossette present at midline. Base of crown swollen slightly labially and lingually. P^ relatively large, subovate in basal outline, broader posteriorly than anteriorly; high, relatively short, longitudinal crest transected by median set of vertical labial and lingual ridges with production of cuspule at crest; slight cleft present along crest about one-third distance from anterior cusp. Hypocone very low, connected to metacone by weak anterolabial ridge; stronger ridge curves posterolabially to meet posterolingual ridge from metaconc, defining shallow posterolin- gual fossette; anterior ridge from hypocone ascends abruptly forming low cingulum poster- olingually, this degenerating anteriorly into basal tubercles, lingual basin very restricted. M' 20 m dark stipple, 10-20 m pale stipple, 5 m thin contin. line, high water mark thick contin. line. STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 187 tidal currents in the area more easy access to the deeper waters to the south. The area initially selected for dredging lay on the eastern edge of the Middle Banks, abutting on the East Channel (see Fig. I). The effects of dredging this area are likely to be numerous. First, benthic organisms and their environment would be removed from a relatively small area, and there would be an interval before this was recolonised. Secondly, the local topography would alter as sediments filled the excavated area. Thirdly, if the excavation led to generally increased current velocities in the area as a whole, it could lead to general erosion. While such erosion could facilitate navigation it could affect the benthos more widely, and possibly could affect the adjacent shores of Moreton Island. Fourthly, material suspended in the water during dredging would be distributed by the tides. The most important effects could well be due to the slower flood tide which would carry material into the deeper waters to the south. Here there is known to be a rich benthos (Stephenson, Williams and Fig. 2: Location of sites within sampling area. Depths from 1972 survey by Queensland Dept, of Harbours and Marine: 10 fm broken line, 5 fm thin contin. line, high water mark thick contin. line. Sites as solid circles, beacons or buoys as crosses. MEMOIRS OF THE QUEENSLAND MUSEUM Lance 1970) and there is also a seasonal fishery for peneid prawns. After considering the above, we advised against dredging near the existing dredged break in the banks because it was fell this would give maximal erosion and most widespread biotic effects and it is gratifying to record that this advice was accepted. The position tentatively selected was in the vicinity of our sampling sites 22, 25, and 26, (see Fig. 2). A past survey of the area (Stephenson, Williams and Lance 1970) was made with a naturalists dredge and subsequent work at Peel Island in Moreton Bay (Stephenson, Williams and Cook 1974) has shown that this failed to reveal a substantial fraction of the benthos. Present experience showed that the van Veen grab used in near-concurrent studies in Bramble Bay (Stephenson. Raphael and Cook 1976, Stephen- son, Cook, and Raphael 1977) was ineffective at Middle Banks except in some areas during the brief periods of slack tide. Hence an 0- 1 m^ Smith-Mcintyre grab was employed and revealed a surprisingly rich and varied biota. Past experience at both Peel Island and Bramble Bay has shown appreciable time changes in the benthos and to make even approximately effective predictions of the likely effect of sand dredging at Middle Banks, it was felt that sampling should continue at three month intervals for at least two years. Sampling began in September 1972 and before the work was due to be completed two natural catastrophes had occurred. These, a major flooding of the Brisbane River in January 1974 followed by a cyclone in March 1974, produced very marked effects on the biota of Bramble Bay, but preliminary examina- tion of the present catches suggested only slight effects on the Middle Banks benthos, and so the project was concluded after the two years. Biotic Sampling Because of the rapid tidal currents, the effects of dredging will be greater to the north and south of the dredged area than to the east and west. Hence the sampled area was roughly in the form of a rectangle ca 9 km x U5 km. At both northern and southern extremities there were sufficient sighting marks for sites to be arranged on a grid ca 0-4 km apart but in the centre of the area due to a paucity of marks they lay on traverses ca 1 km apart. Fifty-eight sites were originally sampled but because one (site 39) was overlooked on one occasion, for present purposes we consider 57 sites, with original sites 40-58 renumbered 39-57. (Labelled specimens to be deposited in the Queensland Museum carry the original number- ing.) The positions of the final sites are shown in Fig. 2. Locations of sites were by horizontal sextant angles on conspicuous objects in the vicinity (landmarks or beacons) and inaccurary in relocating sites is believed to be not greater than 50 m. At each site on each occasion, duplicate catches were made with the O-I Smith-McIntyre grab. Catches were sieved through a final aperture of 1 mm and the two catches pooled, preserved and the biota separated from the residual sediment at base. The resultant biota is referred to as a sample. Samples were obtained on eight occasions from September 1972 at three month intervals until June 1974, and these are referred to below as times 1-8. Considerable difficulties were encountered in identifying species, this reflecting the paucity of work in similar environments in eastern Australia. Reference collections were established for what were adjudged to be species, and these were later referred either to named specimens collected in previous benthic surveys or to group experts (acknowledge later). In several cases the reference specimens belonged to more than one species, and these polyspecific taxa are indicated later. There were a few cases of ‘oversplitting’ and these wore not apparent until all the numerical analyses were completed. The Area of Sampling 1. General Description: The northern exit from Moreton Bay consists of alternate banks and channels which, south of the narrowest portion, run generally from a north-east to south-west direction. This portion has been called the ‘north-eastern corrugations’ by Stephenson, Williams and Lance (1970). The main features from west to east are Western Banks, Pearl Channel, Central Banks, Main Channel, Middle Banks, and East Channel. Larger vessels leave Moreton Bay through a dredged cutting in Middle Banks, then via the East Channel until it fuses with the Main Channel. North of the narrowest portion there are extensive banks to the north-west of Comboyuro Point, and the main navigational channel runs to the NNW. The southern part of the sampled area lies in the deepest portion of Moreton Bay, earlier called the ‘basin’ by Stephenson, Williams and Lance (1970). Depths are almost 30 m and here the tidal STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 189 current on the sea floor is likely to be negligible. The substratum contains obvious silt. The general topography of the northern exit from Moreton Bay is suggestive of a basin c. 30 m deep which has been partially filled by sand. Maxwell (1970) states that wind and stream erosion as well as undercutting of the shore line have been responsible for large quantities of sand being carried westwards into Moreton Bay. Presumably Moreton Island has supplied the sand for the north-eastern corrugations. While over a long period the Middle Banks area has become partially filled with sand, there is evidence that this has not been a continuous and progressive process. Three sets of soundings of the Middle Banks area have been made. The first made in 1865 show a considerable area of the Middle Banks shallower than 10 fm. The second soundings were made in 1946 and show an enlarged shallow area towards the south. A third set of soundings made by the Queensland Department of Harbours and Marine in relation to the Airport Scheme in 1972 show a reduction of the southern shallow area. This must be due to dredging the navigational channel. However the later soundings show that the portion of the East Channel deeper than 10 fm has become more extensive (sec Fig. 3), and it seems likely that sand has been removed to the north. Throughout tidal currents are strong and underwater observations showed sand ripples c. 0-5 m high on the bottom in the vicinity of site 12 and these conditions probably apply to most of the northern sites. Here the sediment surface will be partially mobile and is very ‘clean’. 2. SEDIMENTS: These were collected using the 01 m2 Smith-Mcintyre grab, with hand selection of an aliquot. (It was not appreciated at the times of collection that hand selection causes error, see Stephenson. Cook and Raphael 1977.) Samples were obtained in September 1972, September 1973 and March 1974. Analyses were by wet sieving as described in previous papers, using sieves of the following apertures in mm: 2-00, 1-00, 0-50, 0-25, 0-125, 0 063. The retained grades are described repectively as gravel = shell grit, very coarse sand, coarse sand, medium sand, fine sand, and very fine sand, with the material passing the finest sieve described as mud. The data comprise a three-dimensional matrix of percentages of 7 sediments grades x 57 sites X 3 times. This was classified by a hierarchical agglomerative method (Bray-Curtis, group average) differing from the methodology for biotic data (see later) only in that sedimentary data were not transformed (they are already standardised by Fig 3: Depths of southern % of sampling area on successive surveys; top 1865 survey, middle 1946, bottom 1972. Depths: > 10 fm, stipple; 6 fm, thin contin. line; 3 fm, dotted line; high water mark, thin contin. line. Arrow, true north; scale line 1 km. sample total). The method is identical with that used by Stephenson, Cook and Raphael (1977). Classification of sites by sediments is shown by dendrogram on Fig. 4a, and of times by sediments on Fig. 4b. The sites-classificalion gives an initial dicho- tomy at ca 70% dissimilarity and the main groups were accepted at 10-25% dissimilarity. In contrast the times-classification gives an initial dichotomy at ca 6% dissimilarity and the groups finally accepted were at ca 4% dissimilarity. In the sites classification of sediments three main groups were first accepted, and the subgroups of two of these explored for topographic coherence. Only one showed this and hence four sediment site-groups were finally accepted, and on Fig. 5 these groups are map-plotted. They form a general sequence from site-group 1 in the south-west, through site-group 2 as a transverse southern band to site-group 4 in the north. Site-group 3 consists of flanking sites mostly 190 MEMOIRS OF THE QUEENSLAND MUSEUM forming a western wedge between site-groups 2 and 4. Mean compositions of the sediments in the four site-groups are given in Table I. In site-group 1 the modal grade of sediment is fine sand, followed by mud; in site-group 2 it is fine sand followed distantly by medium sand while in site-group 3 it is again fine sand but more closely followed by medium sand. The northern sediment site-group (4) differs markedly from the remainder in having medium sand as the modal group. The mean composition of sediments in the three times arc given in Table I. This shows a progressive change towards coarseness over the period of investigation. It had been anticipated, following analyses on sediments in Bramble Bay, that the floods of January 1974 would have caused increased mudiness. By the time post-flood Middle Banks sediments could be collected there had been a cyclone. Wave action due to the latter could have been responsible for removal of mud and increased coarseness. Against this there is a distinct suggestion of progressive change. TABLE 1: Mean Percentages of Grades of Sediment in Sediment Site-groups and Sediment Times-groups. Sediment grade Sediment 1 2 Site-groups 3 4 1 (Sept 72) Times 5 (Sept 73) 7 (March 74) Mud 15-7 8-4 6-9 1-4 5-2 5-0 4-4 Very fine sand 4-4 1-9 1-7 0-1 I-O 1-2 1-0 Fine sand 76-0 78*7 59-8 13-4 40-6 36-9 33-8 Medium sand 3-6 12-9 31-7 83-6 52-0 56-1 59-6 Coarse sand O-I 0-2 0-3 0-7 0-5 0-6 0-5 Very coarse sand 01 0-1 01 0-3 0-2 0-2 0-3 Gravel/shell grit 01 0-2 0-2 0-6 0-5 0-3 0-4 38,41 39,40 20,28 1, 2, 3, 4, 5, 6 42,43 46,48 33,34 7, 8, 9.10,11,12 44,45 49,50 56 13,14,15,16,17,18 47 51,52 15,20,21,22,23,24 53,54 25,26,27,29,30,31 1 55 1 1 1 1 32,35,36,37,57 1 2 3 4 Fig 4: Dendrograms showing classifications using sediment compositions as attributes: A of 57 sites, B of 3 times. STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 191 Fig 5: Topographic distribution of the four sediment site-groups. Further analyses were conducted, to determine in which sediment site-group the chronological changes were greatest. First, grades of sediments were reduced to five by fusion of coarse sand, very coarse sand and gravel. Next, in each sediment site-group the variance due to the three times in each sediment grade was obtained, and variances were then summated over grades. Results for sediment site-groups 1-4 were sequentially as follows: 10-08, 8-58, 220-52 and 42-06. There is greater variation in the two northern sediment site-groups (3 and 4) than in the two southern ones (1 and 2) with extremely high variance in sediment site-group 3. 3. Hydrography: (a) Currents. No precise data are available on the extent of penetration of Pacific Ocean water into Moreton Bay and its mixation with the waters of the Bay. General observation suggests that surface water from outside the Bay penetrates to well beyond the sampled area on each tidal cycle. Also no precise data are available on tidal currents. It is known that they flood from and ebb to the north and that surface currents are generally in excess of 1 km/hr apart from periods of slack water. (b) Salinities. Data are available for Shark Spit (27°]7'S., 1 53^22 "E.) from a survey made by the CSIRO Division of Fisheries and Oceanography. The data from the deepest samples (c. 29 m) taken at Shark Spit have been reviewed by Stephenson, Cook and Raphael (1977). From August 1972 to July 1973 the salinity averaged 34-5°/oo and ranged from 33-l°/oo to 35-3°/oo. The variations showed a marked inverse relationship to rainfall in the catchment of the Brisbane River in the previous month. From October 1973 to September 1974 the average salinity was 33 ^“/oo and the range of 32-0°/oo to 35-l'^/oo. There was a period of relatively low salinity (c. 32-I®/oo) from February 1974 to August 1974 evidently due to thcieffects of the excessive runoff from the flood of January 1974 being prolonged by the rainfall associated with the cyclone of March 1974. It is clear that the effects of the extreme climatic disturbances on the salinities were prolonged but relatively minor. (c) Temperatures. Data equivalent to the above on water temperatures are: August 1972 to July 1973, mean 22-25°C, range 17-2-26-6'^C; October 1973 to September 1974, mean 21-79'^C, range I5-9~25*8°C. Comparison with data average over the period 1931-50 at the Old Pile Light near the mouth of the Brisbane River suggests that from December 1972 until about December 1973 temperatures at Shark Spit were consistently about half a degree warmer than average. Biotic Data — Analyses Involving all Species Many species were present in so few samples and in such small numbers that they are excluded from later analyses. These rarer species are important in aspects such as faunistics, species richness and diversity, and these and similar topics are dealt with in the present section. Faunistic Composition of the Samples: A total of 468 taxa were obtained originally, and this reduced to 463 because of ‘oversplitting’. Fifteen of the listed taxa are known to be polyspecific. All but the rare species are listed in the Appendix, together with the code numbers used throughout. 192 MEMOIRS OF THE QUEENSLAND MUSEUM The number 463 compares with 420 taxa (excluding amphipods) from grab sampling an area of c. 3 km- near Peel Island (Stephenson, Williams and Cook 1974), 182 from an area of 30 km- in Bramble Bay (Raphael 1974) and 98 from Serpentine Creek (Stephenson and Campbell 1977). Of the 463 taxa 31-8% were polychaetes, 27-9% crustaceans (10-6% decapods and 7-2% am- phipods), 23-5% molluscs (10-8% gastropods and 12-5% bivalves) and 6-3% echinoderms. The recording of 18 species of cumaceans and 14 species of fish was noteworthy. Amongst the latter were small specimens of dubiously benthic species, and several crustaceans (e.g. Lucifer) were also doubtfully benthic. Unless clearly non-benthic (e.g. some crustacean larvae, and some planktonic molluscs) all taxa which were caught are listed. Distribution Patterns of Rarer Species: Rarer species are here taken as those in which only one or two individuals were recorded in the survey. By taking recordings of each individual and summating over times we can determine whether the rare species were concentrated in any patterns of sites, and conversely by summating over sites we can search for patterns in times. The number of rarer recordings in sites ranged from zero to 10, and map plotting suggested a pattern in the data somewhat comparable to that obtained in the sediment site-groupings. Mean numbers in sediment site-groups were: site-group 1 (South-west) 3-57, group 2 (southern transverse band) 4 00, group 3 (flanking middle) 8*20 and group 4 (northern) 2-35. Overall the northern area has few rare recordings, the southern area an intermediate number, and the middle flanking portion (with the most unstable sediments) has the largest number. The number of rare recordings in times ranged from 18 to 29 with September values (mean 28-5) noticeably greater than the remainder. Sites x Times Analyses: The total data which were available for analysis formed a three dimensional matrix of 468 taxa (5 for species) x 57 sites {q for quadrats) x 8 limes (t). By summaling over species we derived q x Lmatricies with dimensions of 57 x 8 and two such summations were considered. The first involved the number of species {sj per sample; this is a simple measure of diversity and in Hulbert's (1971) terms it is the species density appropriate to the particular sampling parameters. The second summation was the total number of individuals (n) of ail species, and is the numerical equivalent of biomass. Numerous sample measures other than direct summations are available provided species become anonymous, and we used the standardized Shannon diversity (to log base 10) which we designate d. These three measures s, n, and d were previously used in work on Bramble Bay by Stephenson, Cook and Raphael (1976) (albiet with different symbolism) and Stephenson, Cook and Raphael (1977) and because the sampling procedures differed only in the type of grab employed comparisons can be effected. It is appreciated that with polyspecific and fractionat- ed taxa, the value of s and d are somewhat inaccurate. We are interested firstly in means over all samples. These are for s 3214, for n 170-23, and for d 1-143. They compare with 12-2, 158-4, and 0-74 respectively for preflood conditions in Bramble Bay (Stephenson, Raphael and Cook 1976). Clearly the Middle Banks grounds as a whole carried a richer and more diverse biota than did Bramble Bay in the preflood sampling. Secondly we are interested in the variation of s, n and d values between sites and between times. This requires first averaging by rows and by columns in each of the q x t tables, and secondly assessing the variability in row means and in column means. Variation was expressed as ranges and variances and results are given in Table 2 . These results show there is greater heterogenei- ty in sites than in times, with the ratios of variances being 4-72 for s, 2-94 for n and 8-67 for d. While a fraction of the differences may be due to greater ‘random’ variation in the sites data which are meaned over eight times compared with TABLE 2; Ranges and Variances of Mean Values of 5. n and d in Sites and in Times. Sites Range Means Variance Times Range Means Variance s 20-10-57-13 82-30 24-67-38-30 17-42 n 67-75-392-88 8393-4 85-44-217-65 2854-6 d 0-978-1-394 0-00702 1104-1-186 0-00081 STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 193 times data over 57 sites, the greater importance of sites in the overall heterogeneity is believed to be a real phenomenon. The Bramble Bay work showed a most marked time effect due to the flood of January 1974 (Stephenson, Cook and Raphael 1977) but this is not apparent in the present data. Table 2 shows reasonable constancy in diversities (d) from place to place and from time to time. This suggests that changes in numbers of species and number of individuals ‘balance out’ in the diversity formulation as noted previously in the postflood data from Bramble Bay and also in work on the Serpentine Creek benthos by Stephenson and Campbell (1977). Significant positive correlations were obtained between s. n and d for the different sites Spearman rank correlation coefficients were: between s and n +0-83, between s and d +0-68 and between n and d 4-0-33. Data on time trends are given in Table 3. Spearman rank correlation coefficients between the three sets of values were: s and n +0-98 (very highly significant), between s and d +0*24 and between n and d +0-13 (neither significant). This suggests that chronological changes in s and n are again sufficiently parallel almost to cancel out effects in diversity. Table 3 shows high values of n in each of the two Septembers, and this follows the pattern obtained in an earlier study at Peel Island (Stephenson, Williams and Cook 1974). The results are also similar in that lowest values occur in March of a non-flood year. Values of s follow the same tendency, but no general tendency is apparent in the diversity data. Classificatory Techniques General Techniques: The general approach is classificatory, beginning with a three dimensional matrix of s (species) x q (sites or quadrats) x t (times). By summation over times we drive an s X q matrix and by classification obtain site-groups and their related species-groups; TABLE 3: Ml A\ Valui s oi similarly by summation over sites we derive an 5 X t matrix and hence times-groups and their related species-groups. The classification of the two derived matrices (viz. s xq and sx t) followed the methods used by Stephenson, Raphael and Cook (1976) and this involved: transformation of data using iog,o (n+1), Bray-Curtis dissimilarity measure, group average sorting. For entity (i.e. site or time) classification data were not standardised; for species classification the transformed data were standardised by totals prior to deriving inter-species dissimilarities. Past experience has shown that the main objection to this method lies in the species classification where rarer species form isolated and ‘dominating’ species-groups. It was hoped that by operating in the first instance within narrow ranges of species abundances that this objection would be less serious (see later). In fact the species groupings which were obtained throughout still left much to be desired. Following earlier work (Stephenson, Raphael and Cook 1976; Stephenson and Campbell 1977; Stephenson, Cook and Raphael 1977) we take the view that species should be placed in the same species-group if they characterise groups of entities (sites or times) in a similar way. A satisfactory entity classification then becomes a necessary pre-requisite to a satisfactory species classification. Because of occasional misclassifica- tions, some re-allocation of entities may be necessary, involving either the occasional and obvious cases (e.g. Stephenson and Williams 1971) or by procedures such as REMUL (Lance and Williams 1975). In the present cases there were no obvious misclassifications and re- allocations have not been made. If the entities are sites then the next step is to replace the species x sites matrix by the much smaller species x site-group matrix — this contains the mean recordings of the species. By inspection we can immediately see which species may positively characterise a given site-group by occurring there in proportionally the largest numbers. n A\l) d A1 Dll I I RI N’T Tivii s (Sept 72) 2 (Dec 72) 3 (March 73) Times 4 (June 73) 5 (Sept 73) 6 (Dec 73) 7 (March 74) 8 (June 74) s 32-88 28-23 24-67 32-65 38-30 35-32 33-54 32-53 n 185-21 128-54 85-44 171-05 271-65 191-61 162-25 165-09 d 1-115 1-144 M04 1-127 1-139 1-186 1-172 1-163 194 MEMOIRS OF THE QUEENSLAND MUSEUM However the variability of recordings within the site-groups may be such that differences between means are misleading. To locate such cases, as in work of the recent past, we use a ‘pseudo-F test’ and employ such terms as ‘conformity’ and ‘noticeably different’ instead of ‘significant’, using the 0 05 probability appropriate to the F test as a cut-off point. For a given array of species recordings arranged in site-groups there are many possibilities for testing, for example each site-group set against all the remainder, each pair of site-group sets against the remainder etc. It has been found more economical to perform pseudo-F testing by a programmable hand calculator on those sets suggested by perusal of the data, than to test all by a computer-based programme. Pscudo-F testing reveals species which do not conform to the entity-groupings. When species with a range of abundances are considered we may expect non-conformers to include common species which are evenly distributed amongst the entity-groups and which must not be lost sight of. We may also expect them to include rarer species insufficiently concentrated in any entity group for the results to be meaningful. Data Reduction: In benthic work there are typically many rare species which add little to the ecological understanding of an area beyond the contributions they make to analyses like those of the previous section. There is a more pragmatic reason for their exclusion: they greatly increase computing costs and their inclusion may result in the computer being unable to accept the total data. In the present case, to achieve the limiting matrix of 199 x 57 x 8 would mean excluding species occurring 1 2 times or less, and it was felt this might be too severe. Because there are objections to most of the methods of data reduction previously employed we here attempted alternative approaches. The first consisted of dealing with the species in successive lots of 50 and compared the classifications obtained. This showed, inter al, that the sites were divisible into two markedly dissimilar groups. We then considered the two groups of sites separately — by reducing the q dimension in the matrix the critical s dimension becomes automaticallv reduced. Analyses on Lots of Fifty Species The potential advantages of this method are first that the ‘stopping rule’ as regards species is indicated by the data themselves. It occurs at that point in the successive analyses when either no patterns or ‘nonsensical’ patterns emerge. This involves only a minimum of subjectivity. Secondly it shows whether the patterns revealed by the abundant and less abundant species are similar, and thirdly the results should be less dependent on the data transformation which is employed. This is because there is a narrower range of abundances in a matrix; in practice this relative independence increased as we moved to the less abundant group of species. It was anticipated that the main disadvantage of the method would be in comparing the results of the different analyses but in the event this was only a minor problem. We deal first with ail j x ^ analyses, then all s "x. t analyses. Species x Sites Analyses: We first consider the site classifications, then the conforming species. (a) Sites — first 50 species: The dendrogram of the classification (Fig. 6A) shows remarkably clear separation into two site-groups which consist of .southern and northern sites. Apart from a single site (29) the boundary follows the topography of the area, and coincides with the 10 fm depth contour on the ‘lip’ of the Middle Banks. Three subordinate groupings are evident within the southern subarea, and although these have topographic coherence we do not discuss them further at this stage. (b) Sites — second 50 species: The dendrogram (Fig. 6B) again shows clear separation into northern and southern site-groups, although less clearly than in the previous dendrogram. The two main site-groupings are identical with the previous ones. Three subordinate groupings are again evident in the southern subarea but their constituent sites are not identical with the previous ones. (c) Sites — third 50 species: The northern and southern site-groups were identical with those above. (d) Sites — fourth to sixth lots of 50 species: In the fourth lot the results were only broadly similar to the above with one subgroup mostly of topographically northern sites (7, 20, 23, 24, 29, 35) in the main southern group which also included site 36. In the fifth lot excluding two vacant sites (9, 27), the original groups were revealed excepting that site 31 was in the southern group. In the sixth lot of species the south and north pattern was not evident. There were eight isolated sites and three groups at a 98% STEPFJENSON ET AL; MACROBENTHOS OF MIDDLEBANKS 195 1 ^ 1 75 B 1 55 _ r r--^ 35 1 29,35 38,41 28,33 1, 2, 3, 4, 5, 6 38,41 39,45 28,29 1, 2, 3, 4, 5, 6 50 42,43 34,39 7, 8, 9,10,11,12 42,43 48.51 33,34 7, 8, 9,10,11,12 44,45 40,46 13,14,15,16,17,18 44,47 52,53 35,40 13,14,15,16,17,18 47 48,49 19,20,21,22,23,24 54 46,49 19,20,21,22,23,24 51,52 25,26,27.30.31,32 50,55 25,26,27,30,31,32 53,54 36,37,57 56 36,37,57 55,56 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 a 1b Ic 2 1a lb 1c 2 Fi(i 6: Dendrograms of site classifications using species recordings as attributes: A first 50 speies, B second 50 species. dissimilarity level with only one group showing approximate topographic coherence. The stopping point is clearly at or about the fifth lot of species (species occurrence >6). The distinctiveness of the northern and southern subgroups decreased progressively from the first to the fifth lots of species. This is indicated in Table 4, TABLE 4: Levbls of Dhndrogram Fusions for Spi.c iis Lots with Dfci.ininc; Abundancfs Species lot Approx level of lowest fusions Approx level of highest dichotomy First 50 10 60 Second 50 20 70 Third 50 30 85 Fourth 50 40 90 Fifth 50 50 100 (e) Conformity of species to main site- groupings: For species in the first to third lots, the main site-groups are identical. For species in the fourth to sixth lots of 50 it seems more appropriate to test conformity to the general pattern than to the fragmenting remnants of it. In these cases the rationale of testing for conformity is altered. We obtain groups from elsewhere and test on the lines of a ‘construction set’ of data and a ‘validation set’. Instead of ‘noticeable differences’ of species distributions we might now approximate more closely to ‘sig- nificant differences' excepting that the problems of non-normal distribution with series of zero recordings became more acute. The sixth lot of 50 species takes in only some of the species in which three specimens were recorded and omits others so the present testing was extended to all occurring three times. Below three recordings there can be no conformity in the sense we have used the term. Results of conformity tests to the southern and northern groups of sites covering all species except the less abundant ones are given in the Appendix. It will be noted that more species positively characterise the southern site-group than the northern one. With decreasing abundance of species there are increasing proportions which fail to conform. Thus for 6 and 7 recordings the percentage not conforming is 29, for 5 recordings 36, for 4 it is 56 and for 3 it is 65. (f) The main ‘communities’: If we view a benthic community as an area and the species which characterise it positively, then we have two main communities. It is of interest that identical areas show up in analyses on the first, second and third sets of species and an almost identical one in the fifth set. Clearly the abundant and less abundant species follow the same broad area pattern. 196 MEMOIRS OF THE QUEENSLAND MUSEUM B Fig. 7: Dendrograms of time classifications: A first 50 species, B third 50 species. There are very many conforming species, 158 in the southern community and 80 in the northern one, and to assist comprehension these data must be compressed. If we list only the five commoner species which conform to each of the areas we arrive at a southern community of Prionospio, tanaid 1, Solemya, Callianassa and Poecilochaetus and a northern community of Urohaustorius, Amphiura octacantha, Con- cholestes, mysid 4 and amphipod 3. Four of the northern species are small crustaceans which are possibly mobile; mobility could be an advantage in this generally turbulent area. Conforming species are possibly different from numerically dominant species — a conforming and dominant species in the southern area may well be a dominant species (but clearly not a positively conforming one) in the northern area. Purely on a dominance basis the southern area is a community of Prionospio, tanaid 1, Solemya, Poecilochaetus and Discobotellina while the northern area is a Urohaustorius, Prionospio. Amphiura octacantha. Concholestes and mysid 4 community. Species x Times Analyses; Time classifica- tions of the first and third lots of 50 species are shown as dendrograms on Fig. 7, and these are representative of the remainder. No two classifications were identical and the only common grouping was of times 7 and 8. The times were grouped in a sequential or near-sequential order throughout, and there were never close groupings of the equivalent seasons of the two years. By comparing Figs. 6 and 7 it is evident that dissimilarities between times are much less than between sites. With the lack of constancy in time patterns, tests of conformity of species to these patterns are on a somewhat shifting basis. Taking three times-groups in each case gave only 75 conforming species compared with 238 conforming to two site-groups. Further consideration of times- groupings is deferred until the next section. Biotic Data — Separate Analyses OF Northern and Southern Areas GENERAL: For reasons given earlier it was decided to subdivide the study area into southern and northern subareas and to analyse each separately. Additional reasons are (a) by working within these respective communities heterogeneity is greatly reduced, (b) special attention can be given to the northern area which would be most immediately involved in human interference, (c) it is possible that time changes differ in the two areas. The problem about subdivision is that three different criteria can be used and each gives slightly different results. Using sediments site 20 is included in the southern group, but its biotic and topographic affinities are with the northern group. Using topography site 29 is in the northern group, but using biotic affinities it is in the southern. Site 36 alters affinities between one biotic classification and another. Eventually an overlapping division was decided upon with sites 29 and 36 in both groups. The northern group then consists of sites 1-27, 29-32, 36, 37 and 57 (34 in all) while the southern group consists of sites 28, 29, 33-36, 38-56 (25 in all). STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 197 A 35 _ 30 . B 38,41 29,35 39.45 28,33 29,36 15,16 42.43 50 48 51 34,40 17,20 44.47 1 1 ! 52,53 54 I I 46,49 55,56 I 1 1 1 1 1 4 2 3 6 10 1, 2 5,13 21,22 26,31 3. 4 14,19 23,24 32 7, 8 25,27 9,10 30 11,12 37,57 12 J L 11 J L Fig. 8: Dendrograms of site classifications in A southern and B northern sub-areas. (Site-group 12 is disregarded in the main analysis.) Fig. 9: Topographic distribution of the 11 main site-groups. Site-groups 4 and 6 overlap and their individual sites are shown by the symbols indicated. Species Considered: An arbitrary cut-off level was adopted, and species occurring less than 10 times in the reduced data were excluded. Because we now look for finer patterns and hence more site-groups, a higher cut-off level than previously was thought desirable. It was confirmed later as being approximately correct. The southern analyses involved 159 species and the northern ones 115. Species x Sites Analyses: (a) Sites classifica- tions. The dendrograms of site-classifications are given m Figure 8. We first consider the marginal sites 29 and 36 common to the two analyses. Site 36 joins the southern area dendrogram at a much higher dissimilarity level than for the northern area and is henceforth considered as a northern site. Site 29 is ambiguous in its affinities and is treated as both a northern and southern site. In Figure 8A four site-groups are readily recognisable at a dissimilarity level of c. 30% and these show topographical coherence. They are map plotted in Figure 9 and numbered 1-4 in a S-N direction. In Figure 8B at just over 30% dissimilarity there are five groups of very dissimilar sizes. The largest group divides at a slightly lower level to give groups of 11 and 14 sites and of these the former gives a chained series and is retained as a unit. The latter spans the area of greatest interest where excavation may occur, and divides into three topographically coherent sub groups at about 25% level. These subgroups were retained. In all eight northern groups were tentatively accepted and are map plotted as groups 5-12 (see Fig. 9). Site-group 12 consists of the single site 6 which is marginal to the area of 198 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 5: Species Conforming to the Four Southern Site-groups. Category Species Most in site-group 1 25, 16, 19, 90. Ill, 134, (-6) Approx coequal most site-groups 1 and 2 14, 29, 98, 96, 118, 124, 170 (-7) Most in site-group 2 156, 176 ( = 2) Approx coequal most site-groups 2 and 3 1, 9, 15, 37, 66, 75, 139, 146, 181, 212 (=10) Most in site-group 3 3, 4, 45, 65, 94. 108, 93, 172, 137, 173, 175, 219 (=12) Approx coequal most site-groups 3 and 4 52, 48, 68, 63. 64, 32, 61, 53, 34, 20, 84, 83, 88, 103, 22, 85, 126, 135, 128, 151, 113, 131, 49, 143, 147, 161, 168 ( = 27) Most in site-group 4 51, 8, 2, 7, 23, 21, 81, 46, 18, 43, 106, 58, 56, 150, 129, 6, 44, 57, 70 (=19) Most in site-groups I, 2, 3 (least in 4) 24, 27, 33, 36, 38, 60, 95, 127, 159 ( = 9) Most in site-groups 2, 3. 4 (least in 1) 26, 30, 39, 35, 97, 74, 107, 117, 77, 144, 141 (=11) Species arc ranked from those with highest to those with lowest recordings in each group. TABLE 6: Species in the Seven Northern Site-groups. Category Species Approx coequal most site-groups 5 and 6 31, 3 ( = 2) Most in site-group 6 1, 10, 4, 52, 9, 30, 15, 63, 26, 13, 11, 40, 83, 37, 41, 91, 17 ( = 17) Most in site-groups 5, 6, 7 138 (=1) Most in site-groups 6 and 7 5 (=1) Most in site-group 7 49, 199 ( = 2) TABLE 6: cont Category Species Most in site-groups 7 and 8 160 (=1) Most in site-group 8 54, 86, 104 ( = 3) Most in site-groups 7, 8, 9 7, 57, 70, 91, 163, 93, 109 ( = 7) Most in site-groups 7, 8, 9, 11 20, 59 ( = 2) Most in site-groups 6, 7. 8, 9 23, 77 ( = 2) Most in site-group 9 6, 8, 102, 89, 116, 203 (=6) Most in site-group 10 21, 47, 55, 155, 182 ( = 5) Most in site-group 1 1 43, 99, 136, 140 ( = 4) Most in site-groups 10 and 1 1 2 (=1) Most in site-groups 9, 10, 1 1 32 (=1) Most in site-groups 9 and 1 1 105 (=1) Most in site-groups 5, 6, 7, 8 (least in 9, 10, 11) 39 (=1) Most in site-groups 5, 6, 7, 8, 9 (least in 10, 11) 34 (=1) Most in site-groups 5, 6, 7, 8, 9, 10 (least in 11) 113 ( = 1) Most in site-groups 6, 7, 8, 9, 10, II (least in 5) 18 (=1) Most in site-groups 5, 7, 8, 9, 10, 11 (least in 6) 22 (=1) Species are ranked from those with highest to those with lowest recordings in each group. STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 199 interest and is disregarded in subsequent analyses. (b) Species-groupings, southern sub-area. Of the 159 species considered, 42 did not conform to any rational arrangement of site-groups. Of the six species occurring 11 and 10 times, only one conformed indicating a correct choice of cut-off point. The conforming species are listed in Table 5 which shows that in general more species characterise two site-groups than one. If we include species characterising one, two or three, site-groups, then 22 species characterise site-group I, 39 site-group 2, 69 site-group 3 and 57 site-group 4. To describe communities in terms of these numbers of characterising species is not helpful. As stated earlier an alternative method is not to consider conforming species but instead a few of the numerically dominant species in each site-group and this we follow. (c) Species-groupings, northern sub-area. Of the 115 species considered, 54 did not conform to any rational arrangement of site-groups. Of the four species occurring less than 12 times, only one conformed, again confirming the choice of cut-off level. The conforming species are listed in Table 6, which shows that site-group 6 has the largest number of uniquely characterising species (17); this agrees with the isolated position of site-group 6 on the dendrogram (Fig. 8B). Many of the species conform to combinations of site-groups, with the largest number (17) involving site-groups 7 and 8 followed by site-groups 7, 8 and 9 (16spp.). This confirms the similarity of these site-groups, again as shown in the dendrogram. Counting species which characterise more than one site-group, then 7 characterise site-group 5, 27 site-group 6, 21 site-group 7, 20 site-group 8, 23 site-group 9, 10 site-group 10 and 1 1 site-group II. (d) Overall view of site-groups. Table 7 shows the site composition of each of the eleven main site-groups considered, and their more abundant characterising species. Data from the previous q X / analyses are added to this table. They comprise the mean number of individuals of all species per site in a site-group per time (n), the mean number of species (s) and mean standardised Shannon diversity (to log base 10) {d}. Site-group 1 occupies the extreme SW. of the sampled area where sediments are predominantly fine sand (76%) with the highest percentage of mud (ca 16%), and in relatively deep water (ca 31 m). It may be described in terms of dominants as a Schizaster — Nucula — Prionospio community and within the southern area has the lowest average population (197), lowest species density (36), and lowest diversity (M2). Site-group 2 forms an arc to the north of site-group 1 in slightly shallower water (mean depth c. 30 m). It was not clearly distinguished by its sediments but these are predominantly of fine sand (79%) followed by medium sand (10%). It may be described as a Prionospio - Aglaophamus — Callianassa community, has a high average population (278) and for the southern sub-area a relatively low species density (38) and diversity (1T7). Site-group 3 (depth c. 22 m) is parallel to and to the north of site-group 2. It does not precisely follow the topography of the area, lying at a slight angle to the 20 m contour and is mostly deeper than this. It is not clearly distinguished by its sediments. These are predominantly fine sand (69%) followed by medium sand (24%). It may be described as a Prionospio — tanaid -- Solemya community. It has the highest recorded population (320) and number of species (48), and the average of the diversity in the total area (1-20). Site-group 4 (depth ca 16 m) consists of three sites straddling and somewhat askew to the 20 m contour just south of the southern lip’ of the sampled area. Its sediments are highly variable, site 50 having predominantly fine sand (78%), and both sites 29 and 35 having predominantly medium sand (76% and 66% respectively). It could be described as a Rhizammina — Discobotellina — Prionospio — Solemya community but for the fact that the first named species is markedly seasonal. The population (215) is below the average for the southern site-groups but much in excess of any northern groups. The species density (46) is almost that of site-group 3, and the diversity (1-29) is the highest of the southern site-groups. Almost identically high diversity also occurs in site-group 6 (depth c. 9 m) which again contains site-group 29. It can be described as a Prionospio — Urohaustorius community, has the lowest population (79) of any site-group, the lowest species density (24) but the highest diversity (1-56). Site-group 5 is at the western edge of the Middle Banks lip, with a modal sediment of medium sand (82%). It is a Prionospio — Urohauslorius — tanaid communi- ty and if reckoned as within the northern sub-area has its highest population (143), and species density (38), and has a high diversity (1-30), second only to site-group 6 (depth c. 1 1 m). 200 MEMOIRS OF THE QUEENSLAND MUSEUM TABLE 7: Drtah s OF Site-groups. Site-group I 2 3 4 5 Constituent sites 38, 41, 42 43, 44, 47 39, 45, 48, 51, 52, 53, 54 28, 33, 34, 40, 46, 49, 55, 56 29, 35, 50 37, 57 Most abundant species (occurrences*) 25(22), 12(22), 1(21), 10(18), 16(17), 19(14), 14(14), 11(11) 1(72), 11(17), 5(14), 9(14), 3(13), 4(12), 3(11) 1(77), 3(32), 4(30), 9(16), 13(13), 15(12) 28(27), 10(20), 1(16), 4(14), 3(10) 1(11), 2(9) h* 197 278 320 215 79 }* 36 38 48 46 24 d* 1-12 M7 1-20 1-29 1-56 Main sed. grades 76% fine sand 16% mud 79% fine sand 10% med. sand 69% fine sand 24% med. sand Variable 87% med. sand Mean depth (m) 30-5 29-9 22-3 16-2 1 1-0 * Rounded None of the remaining site-groups appear to be distinguished by their sediments, ail having predominantly medium sand present as in the two previous cases. Site-group 7 (depth c. 15 m) includes two sites (31, 32) in or close to the northern end of the dredged navigational channel. It can be described as a Concholestes — Callianassa — Prionospio community, and has almost the lowest population of any site-group (81), and also low species richness (25) and diversity (Ml). Site-group 8 (depth c. 14 m) lies on the eastern slopes of the Middle Banks in the area from which sand might be dredged and may be described as a Concholestes — Urohaustorius — mysid 4 community. It has a relatively low population density (91) but somewhat higher species richness (30) and diversity (M8) than adjacent site-groups. Site-group 9 lies in the NE. of the sampled area, in generally deeper water than adjacent sites (depth c. 19 m) and may be described as a Concholestes — Amphiura octacantha — mysid 4 community. It has a higher population than adjacent site-groups (125); species richness is low (25) as is diversity (109). Site-group 10 lies in the W. of the sampled area in a generally shallow and turbulent area (depth c. 8 m). It forms a Urohaustorius — Platyischnopus — Prionospio community with low population (113), species density (25) and diversity (103). Site-group 1 1 is the northernmost area sampled. It lies in a turbulent area adjacent to but deeper than that of site-group 10 (depth c. 14 m). It forms a Urohaustorius — Amphiura octacantha — Prionospio community, again with low population (96), species richness (25) and diversity (Ml). SPECifs X Times Analyses: The dendrograms of time classification for the southern sub-area (Fig. 10A)and northern sub-area (Fig. lOB) are alike in showing groupings which are predomin- antly of sequential times and in not following seasonal patterns. The sequential pattern is slightly better developed in the northern sub-area and also the inter-time dissimilarities are generally lower. By dealing with sub-areas the dissimilarity levels in the times dendrograms have become roughly equated to those of the sites dendrograms and hence times now become relatively more important. Perusal of the two-way coincidence table of time-groups with species showed that less than a third of the species conformed noticeably to the time-groups of the dendrograms. As an alternative to the dendrogram groupings of times, these were dissected into chronologically meaningful groupings and the conformity of species to these groupings was investigated. Approximately half the species analysed con- formed to these groupings. Some of the remaining species occurred at random (see later) and it was concluded here there were no chronological STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 201 TABLE 7: Cont. 6 7 8 9 10 11 29, 36 26. 31, 32 21, 22, 23 24, 25, 27, 30 5, 13, 14, 19 15, 16, 17, 20 1, 2, 3, 4, 7, 8, 10, 11, 12, 18 1(24). 7(10), 5(10), 7(19), 2(8), 7(27), 6(21), 2(34), 21(12), 2(16), 6(14), 2(6), 1(8), 6(6), 8(8), 6(6), 8(14), 1(7), 1(10), 18(7), 1(10), 8(9) 3(5) 20(6), 1(6) 23(6) 8(5) 143 81 91 125 113 96 38 25 30 25 25 25 1-30 11 1 1-18 1-09 1-03 Ml 82% med. sand — > 80% med. sand — — 9-1 14-6 14-0 19-2 7-6 14-0 changes. For the residue while recordings were too heterogeneous to be statistically at random, the groupings of times to which the species conformed did not make obvious chronological sense. Finally the species were divided into four broad categories, and each of these further subdivided. Throughout the categories were non-overlapping and sometimes this involved difficult decisions regarding allocation. The categories were: A — those occurring at random, and hence showing no chronological pattern. For the commoner species randomness was tested by using summed recordings in each of the eight times. For species occurring with fewer than 16 individuals, the limit of formal use of is exceeded. We have still applied the mechanics of the X “ test, which might now be designated a ‘pseudo- X 2 test’ and describe the species as ‘pseudo-random’; B — those with Fig. 10: Dendrograms of time classifications within A sequential patterns of various types. One sequence of times was dissected from the remainder and the pseudo F test applied to the Iog,o (« + 1) transformations of the two sets of recordings. The first dissection point separated times 1-6 from 7 and 8, and we thus determine the species with noticeably decreased numbers after the flood and those with noticeably increased numbers. The second point was between times 1-4 and 5-8, separating the two years of study, and we thus derive species occuring in high numbers in the second year and in the first year respectively. The remaining sequential species followed a variety of time combinations and we first separated those with three sequential noticeably high recordings ('9 month species’) from those with two in sequence ('6 month species’). Finally we listed species with a sequence of two noticeably low B southern and B northern sub-areas. 202 MEMOIRS OF THE QUEENSLAND MUSEUM recordings; C - those with seasonal patterns with noticeably high recoi dings in the same season of both years (mostly in September) and those with noticeably low recordings, D — non-random species giving apparently 'nonsense' chronological patterns; these are presented in order of decreasing likelihood that they give conceptual sense. The first group contains species with a single outstandingly low value, the second species Tabu- 8; Conkormity or Specics to Prescribed Time-groups. Random Random South: 29, 53, 125, 137, 156 ( = 5 = 3%) North: 15, 72, 73, 83, 87, 116, 130, 134, 136, 142 (=10=9%) Pseudo-random South: no, 171, 180, 190, 191, 192, 219, 224 (=8=5%) North: 17, 38, 109, 117, 149, 169, 189 ( = 7 = 6%) Sequential Fewer after flood South; 25. 30, 41 ( = 3 = 2%) North: 22, 42, 110 ( = 3 = 3%) More after Jlood South: 4. 56, 58. 66, 105, 127, 146, 149, 161, 184 (=10=6%) North: 31, 59, 86, 146, 203 ( = 5 = 4%) More in 1st year South: 96, 131 ( = 2=1%) North: 89. 199 ( = 2=2%) More in 2nd year South: 9, li 19, 20, 23, 26, 38, 49, 55, 65, 75, 109, ill. 133, 147, 212 ( = 6=12%) North: 49, 66. 125, 162, 182 ( = 5=4%) 9-month species Soulh; 6, 59, 83, 95, 128 ( = 5 = 3%) North: 18, 23, 54, 55, 74 ( = 5=4%) 6-month species South: 1, 2, 7, 8, 10, 13, 14, 16, 21, 24, 27, 33, 39, 44, 50, 64, 71, 74, 106, 107, 120, 154, 166, 197 (=24=15%) North: 2, 6, 7, 1 1, 13, 24, 37, 67, 69, 70. 71, 82, 92 (=13 = 11%) Sequential low recordings South: 3, 31, 32, 48 (=4=3%) North; 9, 20, 26. 32, 57 ( = 5=4%) Seasonai. Seasonal high recordings (mostly Septembers) South; 12, 28. 45, 54, 61, 81, 97, 101, 108, 118, 126, 129, 135 (=13 = 8%) North: 21, 63. 77, 121 (=4=3%) Seasonal low recordings (various) South, nil (=0=0%) North: 5, 8, 56, 58 ( = 4 = 3%) O BSC l IRE Single outstanding low South: 36, 37, 40, 69 (=4) North; 20, 35, 39, 43, 44, 46, 50 ( = 7) Oscillating South; 11, 18, 35.. 42, 43. 46, 47, 52, 57, 60, 62, 67, 70, 76, 80, 84, 85, 88. 90, 91. 92, 93, 94, 98, 104, 113, 115, 117, 122, 132, 139, 141, 143, 152, 153, 168, 170, 172, 176, 204, 209, 216 ( = 42) North- 1, 4, 10, 12, 19, 40, 41, 45, 47, 48, 53, 62, 65. 75, 78, 79, 85, 91, 93, 99, 102, 104, 105, 112, 113, 114, 132, 140, 141, 147, 155. 158, 160, 163, 185, 187 ( = 36) Single outstanding high South: 5, 17, 22, 34, 51, 68, 77, 8?, 103„ 124, 138, 144, 145, 150, 151, 159, 173, 175, 178, 181, 186, 196, 211 ( = 23) North: 3, 27, 43, 52, 100, 119, 138, 148, 164 (-9) with oscillating values and typically with alternate high and low recordings, while the last group contains species with a single outstandingly high value. Details of the species in each of these categories in southern and northern sub-areas are given in Table 8. Summated percentages of random and pseudorandom species are 8% S., 15% N.; of sequential species 40% S., 33% N.; of seasonal species 8% S., 7% N., and of obscure time patterns 43% S., and 45% N None of the differences between the two sub-areas were significant. The flood of 1974 produced noteworthy changes in less than 10% of the species, in marked contradistinction to Bramble Bay (Stephenson, Cook and Raphael 1977) More species occurred in noticeably high numbers after the flood (15) than in noticeably low numbers. Apart from flood-affected species, 21 species occurred in higher numbers in the second year of study compared with four in the first year. DISCUSSION Site PATTERNS Four groups of sites were clearly recognisable in the southern sub-area and seven groups were less clearly recognisible in the northern sub-area. There is no accepted level at which these site-groups can be designated as communities and subcommunities. Characteristics of each of these site-groups have been given in the text (especially in Table 6) and we now discuss all except the most northerly ones (site-groups 10 and 11). The southern groupings (site-group 1-4) are in deeper areas of fine sand furthest from oceanic water and have relatively STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 203 high populations, species densities and diversities. The individual groups within the southern area do not precisely follow either the topography or sedimentology of the area and one can only assume that a combination of hydrographic parameters is responsible for their delineation. A likely combination would be distance from oceanic water and current velocity. The highest popula- tions and species densities occur on the relatively steep slopes just south of the Middle Banks, and further south populations and species densities are reduced and other ‘communities’ can be recognised. Within the confines of our study area the areas of highest populations and species densities do not form an ecotone between south and north; there is no doubt the populous area belongs to a southern site-group. One suspects on the basis of a study of the dredged biota of Moreton Bay by Stephenson, Williams and Lance (1970), that our most south-western site-group 1 may extend considerably beyond our study area, in which case the populous area of site-group 3 (and to a lesser extent site-group 2) become something of especial local interest. Taking site-groups 2 and 3 together, in terms of population density we have an average of c. 1500 animals per square metre contrasting with r. 1000 further south and c. 600 further north. These are not high values within either a Moreton Bay or a more global context. For example inshore at Bramble Bay, Stephenson, Raphael and Cook (1976) give preflood mean values of up to and 3000 organisms per square metre, and quote both higher and lower figures from the literature. On the other hand both species density (c. 43) and diversity (r. 1-2) are much higher than were obtained with a van Veen grab at Bramble Bay (c. 12 and 0-8 respectively). The dense macrobenthos in site-groups 2 and 3 corresponds with an area in which prawn trawling is concentrated. Presumably the factors responsible for the dense benthos, plus the benthos itself, produce a concentration of prawns. If so the prawns and the trawling could both be expected to take their loll of the benthos. It is undesirable that future engineering works should have any permanent deleterious effects on the areas of site-groups 2 and 3. We can obtain some insight on the likelihood of these occurring by attempting an evaluation of the effects of engineering works now in operation — viz. dredging of the navigational channel (between the beacons in Fig. 2). Sites 40, 50 and 56 lie immediately south of this channel, and they do not appear to show any biotic differences compared with neighbouring sites. In fact two of these sites (40 and 56) are in the area of greatest population density (site-group 3). Site-group 5 is of especial interest because the two sites lie in (site 37) and in immediate proximity with (site 57) the southern end of the dredged channel. They are characterised by the lowest population density (79) and species density (24) obtained in the entire survey, and it is difficult to believe other than that dredging has been responsible. It is of interest to note that this site-group has the highest standardi.sed Shannon diversity (mean value 1*56), and this provides a good example of how misleading diversity measures can be. Site-groups 4 and 6 lie respectively just south of the topographic boundary between southern and northern areas, and on or just north of the sedimentary boundary. They are intermediate as regards population density (ranging from c. 1050 to c. 750 individuals per square metre) but have higher species densities and diversities than northern sites. Site-group 4 is of especial interest because two of its characterising species {Rhizammina and Discobotellina) are arenaceous Foramenifera, although one of these iRhizammina) is markedly seasonal. Discobotellina is widespread in Moreton Bay (Stephenson, Williams and Lance 1970) and in different areas is in association with a great variety of other species. There is a reasonably close parallel with dominance of other species of areaceous Foramenifera off the W. African coast (Buchanan 1960). In site-group 7 one site (32) lies in the northern end of the dredged navigational channel, one lies very near this end (31) and the third (26) c. 1 ‘/2 km away. Populations densities and species densities are approximately as low as those of site-group 5, and again it seems that dredging is responsible. Site-groups 7, 8 and 9 show considerable biotic similarity and form a series running northwards from the dredged area on the western side of the Eastern Channel. There is a progressive increase in population density from 7 to 9, and it is possible this is due to distance from the dredging. Timh-PatternS: Analyses which involved summations over all species indicated seasonally high populations and species densities in each of the Septembers and further that values were higher in the second year than the first one. Analyses on separate species gave obscure sequential groupings of times rather than seasonal groupings, and suggested a progressive shift in the biota. 204 MEMOIRS OF THE QUEENSLAND MUSEUM The latter analyses, while generally more informative, run into a major problem in interpretation which has been raised in an earlier study (Stephenson, Raphael and Cook 1976). Because of inaccurate relocation of sites on successive samplings different local concentrations of species are likely to be sampled on different occasions. It could be argued that because the values considered involve summated recordings at many sites (25 and 34 in the main analyses) ‘random’ microtopographical variations should cancel out. Against this species may be concentrated in certain areas, and a single exceptional patch within these areas could bias the results. The importance of microtopographical patterns has been demonstrated by Poiner (1977) working north of Peel Island. In the present study we can only estimate the relative importance of chronology and microtopography. We assume that when species conform to apparently meaningful chronological changes these changes are real, and that when the chronological picture is decreasingly meaningful there is an increasing possibility of a microtopographical effect. Almost 10% of the species considered conformed to seasonal patterns and in most cases showed noticeably high populations in the two Septembers. High September values have been noted in two previous local studies — near Peel Island (Stephenson, Williams and Cook 1974) and at Bramble Bay before the 1974 flood (Stephen- son, Raphael and Cook 1976) and can now be regarded as an established feature in the benthic calendar. Sequential patterns of one sort or another occurred in about 40% of the species analysed. Because there is a grow'ing interest in non-seasonal changes in benthos (e.g. Stephenson, Williams and Cook 1974; Buchanan, Kingston and Sheader 1974; Eagle 1975; Stephenson, Raphael and Cook 1976; and Stephenson, Cook and Raphael 1977) and because this was the largest category of species with recognisable patterns, special attention was devoted to it. The general approach we followed was to look for changes in biota w'hich are paralleled by abiotic changes and to assume in the first instance that the former are due to the latter. Where no such parallels occur we must fall back upon biotic interactions either between benthic species (Rhoads and Young 1970; Eagle 1975) or between benthic organisms and nektonic predators (Stephenson and Searles 1960; Levings 1972, 1974; Mills 1975). In the present case the most severe abiotic changes for nearly a century were probably the floods of January 1974 and resultant prolonged dilution of the Middle Banks area (Stephenson, Cook and Raphael 1977). Almost 10% of the species showed noticeable changes between the prefiood times (times 1-6) and the postflood ones (times 7-8). While this is more than for any other sequential period, it is a miniscule effect compared with that on Bramble Bay (Stephenson, Cook and Raphael 1977). Moreover at Middle Banks the Hoods had a ‘beneficial’ effect with 15 species occurring in noticeably higher numbers after the Hood and only six before. The effect was most noticeable in the southern area, where ten species increased in number after the Hood. The simplest explanation, that this is due to a supply of sedimentary food material, as MacGinitie (1939) suggested after a Californian Hood, appears unlikely. There was less small particulate sediment in the area after the Hood than before. The remaining sequentially occurring species mostly involved noticeably high numbers for only two consecutive sampling periods i.e. for 6 months or slightly longer. It appears that a sequence of species occupied (in quantity) the different areas of sampling each for tolerably brief periods and were then replaced by others. This is suggestive of rapid recruitment and rapid replacement, as Stephenson, Raphael and Cook (1976) suggested at Bramble Bay. It implies that biotic interactions, whose nature is as yet unknown, are an underlying phenomenon. Superimposed on the transitory aspects of the data there is an underlying trend from lime 1 to time 8 (as shown by the dendrograms) suggesting an overall biotic shift. The somewhat scanty sedimentary data suggest a parallel shift towards coarser sediments and it is tempting to interrelate the two. The largest single category of species revealed by the times analyses (40-45% of the species considered) fail to give immediate chronological sense. There are two possible explanations, microtopographical patchiness or real chron- ological changes on a 3 monthly lime bases. For example species with a single outstandingly low value could reHect a single occasion of severe predation followed by replacement during the next three months; species with oscillating values could reflect irregular and repeated recruitment; while species with a single outstandingly high value would be showing settlement in 3 months and disappearance during the next three months. Present data do not permit an easy choice between the alternative microtopographic or rapid chronological changes; current work suggests both are about equally applicable on an 8 week time basis. STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 205 Continual Biotic Flux: Wc have noted the likelihood of tolerably rapid changes in at least 40% of the species — the sequential ones. We have suggested that rapid changes may be a feature of a greater proportion of the species. If the bulk of the biota consisted of perennial species we would not expect the biota to show rapid changes, and what might appear as such would be due to topographical patchiness. If the bulk of the species were annuals, rapid changes would be inevitable. Thus in theory we could argue from life-spans to the reality of the chronological changes. Unfortunately we have no direct information upon the life-spans of species from the area apart from some data on Discobotellina (Stephenson and Rees 1965a, b) which suggests that larger specimens may be three years old. In general we can only assume that relatively large size equates to an age exceeding one year, and amongst the 100 most abundant species the only cases of relatively large sized species are: spp. 5, 10, 12, 14, 24, 25, 29, 39, 41, 42, 49, 69, 87, 95 and 98 (i.e. 15 in all). Amongst the less abundant species the proportion of large species is much lower, and that of annuals or subannuais must be very high. As Eagle (1975) has noted when populations consist of single year classes we can expect instability. He relates this to low diversity, but as indicated by Stephenson, Williams and Cook (1974) it could go a long way to explaining high diversity in a time context. The overall impression was of a biota dominated by small individuals, of which very few species would attain sexual maturity within the study area. This was particularly so in the northern sub-area; of the 15 larger species listed only one characterised the northern sub-area. If species disappear from within the area before attaining sexual maturity, predation by fish seems a likely cause. Levings (1972, 1974) has shown how seasonal variations in predation by a single species of fish can produce complex changes in a relatively simple benthic biota off the Atlantic coast of Canada. In the present study area there are probably some 20 species of benthic feeding fish (personal communication from Professor J. M. Thomson) and their predation could account for the various and sometimes rapid quasi- seasonal declines in population which were observed. Non-seasonal increases in population are more difficult to explain. One possibility is that many species have prolonged breeding seasons (which is not unlikely in a subtropical situation) and can occupy vacant space at any period within a prolonged settlement period. A balance between prolonged settlement and variable predation could well explain the cases of oscillation noted above. It is clear that much more data are required before this suggestion can be confirmed. Meanwhile the evidence that the benthic population is in an overall state of flux seems tolerably strong. It carries some important consequences as regards a temporary human disturbance of the area. Excavation in the proposed area is unlikely to cause major destruction of breeding populations because few species remain there long enough under natural circumstances to breed. Also, if the normal method of maintenance of the biota is by settlement of one of a great variety of species, it seems probable that this process will only be briefly interrupted, and that a biota roughly comparable with the original should soon be re-established. We might postulate that the settling biota will be more simplified than originally because a larger area and a more vacant area than that provided after fish predation will be available. After some predator-prey oscillations possibly of a few months duration a situation very comparable with the original could be expected. ACKNOWLEDGMENTS We are deeply grateful to the (then) Commonwealth Department of Housing and Construction and to the Queensland Department of the Co-ordinator General for sponsoring the study and providing financial support. We are also most grateful to (a) the CSIRO Division of Fisheries and Oceanography for access to their currently unpublished hydrographic data, (b) the survey branch of the Queensland Department of Harbours and Marine for access to their 1972 soundings of the area and for helpful advice on chartings, and (c) to the Hydrographer of the Royal Australian Navy for survey datings and copies of older charts of the area. Especial thanks are due to the numerous individuals who have assisted in species identifica- tions. In addition to those who named past reference collections, acknowledged in previous papers, these comprise: Dr A. N. Baker, National Museum of New Zealand (ophiuroids and echinoids); Prof. J. H. Day, lately of Zoology Dept., University of Cape Town (cirratulid and orbiniid polychaetes); Dr S. Edmonds, Zoology Dept., University of Adelaide (sipunculids); Dr J. Haig, University of S. California (pagurids); Dr P. Hutchings, Australian Museum (nephtyid and terebellid polychaetes); Mr M. G. Johnston, Zoology Dept., University of Queensland 206 MEMOIRS OF THE QUEENSLAND MUSEUM (cumaceans); Dr J. D. Kudenov, Dept, of Fisheries and Wildlife, Victoria (aphroditid and spionid polychaetes); Dr J. G. B. Poore, same address, (Callianassa and anthurid isopods); Dr C. R. Smalley, Zoology Dept., University of W. Australia (alpheids); Prof. J. M. Thomson, Zoology Dept., University of Queensland (fish); and Miss V. A. Wadley, CSIRO Division of Fisheries and Oceanography (peneids). Not all material in these groups was identified by these experts and the authors accept responsibility for any errors which may have occurred. REFERENCES CITED Buchanan. J. B., 1960. On Julienella and Schizammina two genera of arenaceous foramin- ifera from the tropical Atlantic, with a description of a new species. J. Linn. Soc. (Zool.) 44: 270-7- Buchanan. j. B., Kingston. P. F. and Sheadkr. M., 1974. Long-term population trends of the benthic macrofauna in the offshore mud of the Northum- berland Coast. J. mar. Biol. U.K. 54: 785-95. Eagle, R. A., 1975. Natural fluctuations in a soft bottom benthic community. J. mar. Biol. V.K. 55: 865-78. Hurlbert. S. H., 1971. The nonconcept of species diversity: a critique and alternative parameters. Ecology 52: 577-86. Lance, G. N. and Williams W. T., \915.REMIJL: a new diversity polythetic classificatory program. The Aust. Comp. J. 7: 109-12. Levings. C. D., 1972. ‘A Study of Temporal Change in a Marine Benthic Community, with Particular Reference to Predation by Pseudopleuronectes americanus (Walbaum). (Pisces: Pleuronectidae). Ph.D. Thesis, Dalhousie University, Halifax, Nova Scotia. pp.201. 1974. Seasonal changes in feeding and particle selection by winter flounder (Pleuronectes amer- icanus). Trans Am. Fish Soc. 103: 828-32. MacGinitie, G. E., 1939. Some effects of fresh water on the fauna of a marine harbor. Amer. Midi. Nat. 21: 681-6. Maxwell, W. G. H., 1970. The sedimentary framework of Moreton Bay, Queensland. Aust. J. mar. Freshw. Res. 21: 71-88. Mills. E. L. 1975. Benthic organisms and the structure of marine ecosystems. J. Fish. Res. Bd. Canada. 32: 1657-63. PoiNER. 1. R., 1977. Microvariation in the fauna of a sublittoral sand bank, Moreton Bay, Queensland. Aust. J. Ecol. 2: 297-308. Raphael, Y. L, 1974. ‘The Macrobenthic Fauna of Bramble Bay, Moreton Bay, Queensland." M.Sc. thesis (Unpublished), Zoology Department, Univer- sity of Queensland. Rhoades. D. C. and Young. D. K., 1970. The influence of deposit feeding organisms on sediment stability and community trophic structure. J. mar. Res. 28: 150-78. Stephenson. W., and Campbell, B. M., 1977. The macrobenthos of Serpentine Creek. Mem. Qd Mus 18(1): 75-93. Stephenson. W., Cook. S. D. and Raphael, Y. L, 1977. The effect of a major flood on the macrobenthos of Bramble Bay, Queensland. Mem. Qd Mus. 18(1): 95-118. Stephenson. W., Raphael. Y. 1. and Cook.S. D., 1976. The macrobenthos of Bramble Bay, Queensland. Mem. Qd Mus 17(3): 425-47. Stephenson. W. and Rees. M., 1965a. Ecological and life history studies on a large foraminiferan (Discobotellina biperforata Collins, 1958), from Moreton Bay, Queensland. I. Life cycle and nature of the test. Pap. Dep. Zool. Univ. Qd 2: 207-23. 1965b. Ecological and life history studies on a large foraminiferan (Discobotellina biperforata Collins, 1958), from Moreton Bay. Queensland. 11. Aquarium observations. Pap. Dep. Zool. Univ. Qd 2: 239-58. Stephenson. W. and Searles, R. B., 1960. Experimen- tal studies of the cology of intertidal environments at Heron Island. L Exclusion of fish from beach rock. Aust. J. mar. Freshw. Res. 11: 241-67. Stephenson. W. and Williams, W. T., 1971. A study of the benthos of soft bottoms, Sek Harbour, New Guinea, using numerical analysis. Aust. J. mar. Freshw. Res. 22: 11-34. Stephenson. W., Williams, W. T. and Cook. S. D., 1974. The benthic fauna of soft bottoms. Southern Moreton Bay. Mem. Qd Mus. 17: 73-123. Stephenson. W., Williams. W. T. and Lance, G. N., 1970, The macrobenthos of Moreton Bay. Ecol. Monogr. 40: 459-94. STEPHENSON ET AL; MACROBENTHOS OF MIDDLEBANKS 207 APPENDIX Species from Middle Banks benthos with indications of their systematic positions. Code numbers are in hierarchical order of abundance. Code No. Species Systematic position Number Main Collected site group 1 Prionospio sp. 1 Polychaeta: Spionidae 12802 S 2 Vrohaustorius sp. — polyspecific Amphipoda: Haustoriidae 3394 N 3 tanaid 1 Tanaidacea 3094 S 4 Solemya terraereginae (1 redale) Bivalvia: Solemyidae 3075 S 5 Callianassa arenosa Poore Crustacea: Callianassidae 2776 S 6 Amphiura octacantha H. L. Clark Ophiuroidea: Amphiuridae 2741 N 7 Concholesies sp. polyspecific Amphipoda: Corophiidae 2674 N 8 mysid 4 Mysidacca 2390 N 9 Poecilochaetus sp. Polychaeta: Trochochaetidae 2258 S 10 Discobotellina biperforata Collins Foraminifera: Aslrorhizidae 2131 s 11 Aglaophamiis verrilli McIntosh Polychaeta: Nephtyidae 2040 s 12 Nucula spp. — polyspecific Bivalvia: Nuculidae 1922 s 13 Prionospio sp.4 Polychaeta: Spionidae 1837 s 14 Brissopsis luzonica Gray Echinoidea: Spalangidae 1343 s 15 Magelona sp.2 (‘cincta’) Polychaeta: Magelonidae 1341 s 16 aloidid Bivalvia: Aloididae 1274 s 17 Magelona sp.3 (‘capensis’) Polychaeta; Magelonidae 1217 s 18 amphipod 3 Amphipoda 993 N 19 Golfingia trichocephala (Sluiter) Sipuncula: Golfingiidae 927 s 20 Eunice cf. indica Kinberg Polychaeta: Eunicidae 886 N 21 Platyischnopus sp. — polyspccific Amphipoda: Haustoriidae 852 N 22 gastropod (red and white) Gastropoda 819 N 23 amphipod 4 Amphipoda 792 N 24 bivalve 1 Bivalvia 775 S 25 Schizaster lacunosus (Linnaeus) Echinoidea: Hemiasteridae 703 S 26 Goniada emerita Audouin and Milne Edwards Polychaeta: Glyceridae 688 s 27 Leptomya pura Angas Bivalvia: Semelidae 673 s 28 Rhiiammina cf. algaeformis H. B. Brady Foraminifera: Aslrorhizidae 668 N 29 Xenophrhalmoides dolichophallus Tesch Decapoda; Goneplacidae 654 s 30 carid 2 Decapoda: Caridae 648 s 31 Dispio sp.2 Polychaeta: Spionidae 643 N 32 juv. fibularid Echinoidea: Fibularidae 632 N 33 ‘Dentaliuni’ sp. Scaphopoda 564 S 34 Tharvx sp.A (undescr.) Polychaeta: Cirratulidae 466 N 35 amphipod 6 Amphipoda 433 S 36 Paralacydonia paradoxa Fauvel Polychaeta: Lacydoniidae 432 s 37 Terebellides sfroemi Sars Polychaeta: Terebellidae 428 s 38 Haploscoloplos hifurcatus Hartman Polychaeta: Orbiniidae 423 s 39 Branchiostoma nioretonensis Kelly Cephalochordata 412 s 40 Sthenelais sp. Polychaeta: Aphroditidae 409 s 41 Echinocardium cordatum (Pennant) Echinoidea: Spatangidae 405 s 42 Neosolen vaginoides (Lamarck) Bivalvia: Culiellidae 403 N 43 Mesanthura sp.l Isopoda: Anlhuridae 396 N 44 amphipod 5 Amphipoda 380 N 45 orbiniid spp. — polyspccific Polychaeta: Orbiniidae 360 s 46 Caulleriella sp.A + Tharyx sp.B — polyspccific (both undcscr.) Polychaeta: Cirratulidae 349 N 47 Lucifer hanseni Nobili Decapoda: Sergestidae 344 N 48 lumbrinerid 1 Polychaeta: Lumbrineridae 322 N 49 Epigonichthys cultellus Peters Cephalochordata 298 N 50 Glycera prashadi Fauvel Polychaeta: Glyceridae 293 N 5! Ophiura kinbergi Ljungman Ophiuroidea: Ophiolepidae 290 S 52 Amphipholis loripes Koehler Ophiuroidea: Amphiuridae 269 s 208 MEMOIRS OF THE QUEENSLAND MUSEUM Code No. Species 53 nemertean (pink) — polyspecific 54 Micronephlhys sphaerocirrata Wesenberg-Lund 55 nemertean (pale) — polyspecific 56 Marphysa sp. 57 Mesanthura sp.2 58 Magelona cf. papillicornis Muller 59 polycirrinae — polyspecific 60 bivalve 2 61 Amphioplus depressus (Ljungman) 62 anthurid (undescr. genus 2, sp.2) 63 amphipod 15 — polyspecific 64 bivalve 8 65 ?Notomastus cf. aberans Day 66 = species 34 (Tharyx sp.A) 67 ?Noiomastus latericeus Sars 68 polyzoan 1 69 Sipunculiis aequabilis Sluiter 70 terebellid (undescr. genus) 71 amphipod 14 72 anthurid (undescr. genus 2, sp. 1) 73 Thalenessa sp. 74 Poly dor a sp. 1 75 Owenia fusiformis Delle Chiaje 76 Sthenelepis cf. japonica (McIntosh) 77 Aricidea sp. 78 bivalve 4 79 bivalve 1 2 80 bivalve 14 8 1 bivalve 6 82 Chaetozone sp.A (undescr.) 83 Scalibregma injlatum Rathke 84 amphipod 8 85 nemertean (orange banded) — ?polyspecific 86 Eunice sp.2 87 Polycarpa tinefor (Quoy and Gaimard) 88 Chaetozone sp.B (undescr.) 89 Paraonides sp. 90 Paphia sp. 91 phoronid — ?polyspecific 92 amphipod 9 93 Cyclaspis iribulis (Hale) complex 94 Euclymene sp. 95 Pectinaria antipoda Schmarda 96 Alpheus distinguendus de Man 97 Naticu cf. colliei Recluz 98 Raphidopus ciliaius Stimpson 99 Lumbrineris sp.l 100 gastropod 10 101 Eocuma agrion (Zimmer) 102 bivalve 7 103 gastropod 5 104 Loimia medusa (Savigny) 105 Apseudes sp. 106 Aspidosiphon inquilinus Sluiter 107 Fulvia sp. 108 amphipod 20 109 nereid 1 110 isopod 5 — polyspecific Systematic Number Main position Collected site group Nemertea 264 S Polychaeta: Nephtyidae 262 Nemertea 256 Polychaeta: Eunicidae 227 N Isopoda: Anthuridae 219 N Polychaeta: Magelonidae 212 N Polychaeta: Terebellidae 208 S Bivalvia 188 s Ophiuroidea: Amphiuridae 184 s Isopoda: Anthuridae 180 N Amphipoda 179 s Bivalvia 171 s Polychaeta: Capitellidae 168 s Polychaeta: Cirratulidae 165 s Polychaeta: Capitellidae 162 s Polyzoa 161 s Sipuncula: Sipunculidae 157 s Polychaeta: Terebellidae 151 N Amphipoda 137 s Isolda: Anthuridae 134 N Polychaeta: Aphroditidae 133 N Polychaeta: Spionidae 129 Polychaeta: Oweniidae 117 S Polychaeta: Aphroditidae 109 s Polychaeta: Paraonidae 109 s Bivalvia 106 N Bivalvia 102 N Bivalvia 95 S Bivalvia 94 s Polychaeta: Cirratulidae 93 N Polychaeta: Scalibregmidae 90 s Amphipoda 90 s Nemertea 85 s Polychaeta: Eunicidae 84 N Tunicata: Styelidae 84 s Polychaeta: Cirratulidae 82 s Polychaeta: Paraonidae 80 N Bivalvia: Veneridae 80 s Phoronidae 80 s Amphipoda 78 Cumacea: Bodotriidae 78 s Polychaeta: Maldanidae 75 s Polychaeta: Pectinariidae 75 s Decapoda: Alpheidae 75 s Gastropoda: Naticidae 74 s Decapoda: Porcellanidae 73 s Polychaeta: Eunicidae 71 N Mollusca: Gastropoda 70 Cumacea; Bodotriidae 69 s Bivalvia 69 N Gastropoda 68 s Polychaeta: Terebellidae 66 s Tanaidacea 66 Sipuncula: Aspidosiphonidae 66 s Bivalvia: Cardiidae 65 s Amphipoda 64 s Polychaeta: Nereidae 63 s Crustacea: Isopoda 63 N STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 209 Code No. Species 1 1 1 bivalve 29 1 1 2 Scoloplos johnstonei Day 113 Leptanthura cf. australis Haswell 114 Matuta inermis Miers 115 amphipod 28 1 16 Thermiste sp. 117 Conuber conica (Lamarck) 1 18 Tellina sp.l 119 Diogenes cf. rectimanus Miers 120 Natica sp.l 121 bivalve 16 122 amphipod 13 1 23 gastropod 8 1 24 Protankyra sp. 1 25 Sigalion sp. 126 Dimorphostylis australis (Foxon) 127 aphrodilid 3 1 28 Harmothoe sp. 129 mysid I 130 Drilonereis sp. 131 capitellid spp. — polyspecific 132 carid 4 133 bivalve 22 134 Lumbrineris sp.2 135 amphipod 18 1 36 Syllidia sp. 137 Nereis jacksoni Kinberg 138 amphipod 17 139 Chloeia flava (Pallas) 140 onuphid 1 141 carid 3 142 Veletuceta hedleyi (Lamy) 143 amphipod 21 144 gastropod 3 145 bivalve 3 146 phyllodocid 3 147 sabellid 2 1 48 isopod 3 149 amphipod 1 1 150 amphipod 16 151 gastropod 2 152 Leonnates stephensoni Ruillier 153 Muraenicthys godeffroyi Regan 1 54 isopod 2 155 bivalve 5 156 Tellina texturata Sowerby- 157 Theora lata Hinds 158 bivalve 17 1 59 Spiochaetopterus sp. 160 Pomacuma cognata (Hale) 161 nereid 2 162 cumacean 6 163 Austrolepidoda schmitti Efford and Haig 164 platyhelminth 1 165 amphipod 10 166 isopod 1 167 anemone 1 168 bivalve 9 Systematic Number Main position Collected site group Bivalvia 63 S Polychaeta: Orbiniidae 57 N Isopoda: Anthuridae 56 Crustacea; Calappidae 56 N Amphipoda 51 S Sipuncula: Golfingiidae 51 N Gastropoda: Naticidae 50 S Bivalvia: Tellinidae 50 S Decapoda: Paguridae 49 N Gastropoda: Naticidae 49 S Bivalvia 49 N Amphipoda 48 S Gastropoda 47 Holothuriidae: Synaptidae 47 s Polychaeta: Aphroditidae 46 Cumacea 46 s Polychaeta: Aphroditidae 44 s Polychaeta: Aphroditidae 44 s Mysidacea 44 s Polychaeta: Eunicidae 43 N Polychaeta: CapitelHdae 40 s Decapoda: Caridae 40 Bivalvia 40 s Polychaeta: Eunicidae 39 N Amphipoda 39 s Polychaeta: Hesionidae 36 N Polychaeta: Nereidae 36 S Amphipoda 36 Polychaeta: Amphinomidae 35 s Polychaeta: Eunicidae 35 N Decapoda: Caridae 35 Bivalvia: Glycymeridae 35 N Amphipoda 34 S Gastropoda 34 s Bivalvia 34 s Polychaeta: Phyllodocidae 33 Polychaeta: Sabellidae 33 s Isopoda 33 N Amphipoda 33 Amphipoda 33 s Gastropoda 33 s Polychaeta; Nereidae 32 s Pisces: Myridae 32 s Isopoda 31 s Bivalvia 31 N Bivalvia: Tellinidae 29 s Bivalvia: Semelidae 29 Bivalvia 29 N Polychaeta: Chaetopteridae 28 S Cumacea: Bodotriidae 26 N Polychaeta: Nereidae 25 S Cumacea 25 N Decapoda: Albuneidae 25 N Platyhelminthes: Polycladida 24 N Amphipoda 24 Isopoda 23 S Actiniaria 22 Bivalvia 22 210 MEMOIRS OF THE QUEENSLAND MUSEUM Code No. Species 169 amphipod 24 170 bivalve 11 171 Spiochaetopterus cf. vitrarius (Ehlers) 172 Glyphocurna sp. 173 Penaeus plehejus (Hesse) 174 Polyonyx iransversus (Haswcll) 175 phyllodocid 7 176 Sternaspis scutaia (Renier) 177 Metapenaeus hennettae Racek and Dali 178 gastropod 16 179 phyllodocid 1 180 Glyphocurna halei Greenwood and Johnston 181 Chaetopterus variopedatus Renier 182 Moira lelhe Lutken 183 capitellid 1 184 tanaid 6 185 Trachypenaeus fulvus Dali 186 Philine angasi Crosse and Fisher 187 gastropod 27 188 Cyclaspis mawsonae (Hale) 1 89 isopod 4 190 Dosinia cf. sculpta Hanley 191 bivalve 10 192 Ophiocentrus sp. 193 phyllodocid 2 194 amphipod 26 195 juv. carid 1 196 bivalve 13 197 bivalve 18 198 gastropod 20 199 gastropod 21 200 Ophiodromus sp. 201 Dorvillea sp. 202 Travisia cf. forbesii Johnston 203 maldanid 1 204 maldanid 2 205 tanaid 5 206 Apanthura sp. 207 amphipod 29 208 Hexapus granulijerus Campbell and Stephenson 209 Modiolus sp. 210 bivalve 21 21 1 bivalve 30 212 Natica sp. 4 213 gastropod 25 214 Pista spp. — polyspecific 215 Laonice sp. 216 Merinides sp. 217 Cyclaspis ? crelala (Hale) ? granulos (Hale) 218 anlhurid (undescr. genus 1 sp.l) 219 Doxander viiiaius Linnaeus 220 nemertean (green) 221 aphroditid 1 222 Diplocirrus sp. 223 ^Pontophilus sp.l (doubtfully benthic) 224 Phos sculptilis Watson 225 phyllodocid 6 Systematic Number Main position Collected site group Amphipoda 21 Bivalvia 21 S Polychaeta: Chaetopteridae 20 S Cumacea 20 s Decapoda: Peneidae 20 s Decapoda: Porcelianidae 20 Polychaeta: Phyllodocidac 19 Polychaeta: Slernaspidae 19 s Decapoda: Peneidae 19 Gastropoda 19 s Polychaeta: Phyllodocidae 17 Cumacca: Bodotriidae 17 s Polychaeta: Chaetopteridae 16 s Echinoidea: Hemiasteridae 16 N Polychaeta: Capitcllidae 15 Tanaidacca 15 s Decapoda: Peneidae 15 Gastropoda: Philinidae 15 s Gastropoda 15 N Cumacea: Bodotriidae 14 Isopoda 14 N Bivalvia: Veneridae 14 S Bivalvia 14 s Ophiuroidea: Amphiuridae 14 Phyllodocidac 13 Amphipoda 13 Decapoda: Caridae 13 Bivalvia 13 s Bivalvia 13 s Gastropoda 13 N Gastropoda 13 N Polychaeta: Hesionidae 12 Polychaeta: Runicidae 12 Polychaeta: Opheliidae 12 Polychaeta: Maldaiiidae 12 N Polychaeta: Maldanidae 12 S Tanaidacea 12 s Isopoda: Anthuridae 12 N Amphipoda 12 N Decapoda: Goneplacidae 12 S Bivalvia: Mylilidae 12 s Bivalvia 12 Bivalvia 12 s Gastropoda; Naticidae 12 s Gastropoda 12 Polychaeta: Tercbellidae 11 s Polychaeta: Spionidae 11 s Polychaeta: Spionidae 11 s Cumacea: Bodotriidae 1 1 N Isopoda; Anthuridae 1 1 Gastropoda: Strombidae 1 1 s Nemertca 10 Polychaeta: Aphroditidae 10 s Polychaeta: Flabelligeridae 10 s Crustacea: Crangonidae 9 N Gastropoda; Photidae 10 s Polychaeta: Phyllodocidae 9 N STEPHENSON ET AL: MACROBENTHOS OF MIDDLEBANKS 211 Code No. Species 226 = species 216 (Nerinides sp.) 227 Matuta spp. juv. not M. inermis} — polyspecific 228 bivalve 1 5 229 bivalve 24 230 Sthenolepis sp. 231 Pseudeurythoe microcephala (Fauvel) 232 nereid 4 233 eunicid 1 234 Travisia sp.2 235 bivalve 19 236 Luidia sp. 237 Rhynobrissus pyramidalis Agassiz 238 Glossobalanus sp. 239 polyzoa 2 240 nemertean (brown) 241 Eupanthalis sp. 242 Etenone sp. 243 Nephtys vikingensis Paxton 244 spionid 7 — polyspecific 245 Ophelina cf. acuminata Oersted 246 tanaid 3 247 amphipod 22 248 amphipod 27 249 Betaeus spp. — polyspecific 250 oyster 1 251 Amphiura catephes H. L. Clark 252 Nematonereis sp. 253 Polydora sp.2 254 bivalve 20 255 blenny I (doubtfully benthic) 256 phyllodocid 15 257 Arabella sp. 258 maldanid 3 259 sabellid 3 260 amphipod 19 261 Eucrassatella cumingii (Adams) 262 Mactra sp. 263 Tellina sp.2 264 Natica sp.3 265 gastropod 33 266 Mogula mollis Herdmann 267 Euleanira sp. 268 phyllodocid 5 269 phyllodocid 8 270 phyllodocid 10 271 nereid 3 272 Prionospio sp.3 273 amphipod 25 274 Alpheus spp. (larva) — polyspecific 275 Albunea cf. microps Miers 276 Limaria sp. 277 bivalve 25 278 bivalve 26 279 bivalve 31 280 gastropod 13 281 gastropod 15 282 Pentaceraster australis (Luther) 283 enteropneust Systematic position Number Main Collected site group Polychaeta: Spionidae 9 S Decapoda: Calappidae 9 N Bivalvia 9 N Bivalvia 9 S Polychaeta: Aphroditidae 8 S Polychaeta: Amphinomidae 8 S Polychaeta: Nereidae 8 S Polychaeta: Eunicidae 8 N Polychaeta: Opheliidae 8 N Bivalvia 8 N Asleroidea: Luididae 8 S Echinoidea: Spatangidae 8 N Balanogiossidae 8 S Bryozoa 7 Nemertea 7 s Polychaeta: Aphroditidae 7 s Polychaeta: Phyllodocidae 7 Polychaeta: Nephtyidae 7 N Polychaeta: Spionidae 7 s Polychaeta: Opheliidae 7 Tanaidacea 7 s Amphipoda 7 Amphipoda 7 s Decapoda: Alpheidae 7 s Bivalvia: Ostreidae 7 Ophiuroidea: Amphiuridae 7 S Polychaeta: Eunicidae 6 s Polychaeta: Spionidae 6 S Bivalvia 6 s Pisces: Blenniidae 7 N Polychaeta: Phyllodocidae 5 N Polychaeta: Eunicidae 5 Polychaeta: Maldanidae 5 s Polychaeta: Sabellidae 5 Amphipoda 5 S Bivalvia: Eucrassatellidae 5 N Bivalvia: Mactridae 5 N Bivalvia: Tellinidae 5 N Gastropoda: Naticidae 5 S Gastropoda 5 Tunicata: Molgulidae 5 Polychaeta: Aphroditidae 4 s Polychaeta: Phyllodocidae 4 Polychaeta: Phyllodocidae 4 s Polychaeta: Phyllodocidae 4 s Polychaeta: Nereidae 4 Polychaeta: Spionidae 4 Amphipoda 4 Decapoda: Alpheidae 4 s Decapoda: Albuneidae 4 Bivalvia: Limidae 4 s Bivalvia 4 s Bivalvia 4 s Bivalvia 4 Gastropoda 4 Gastropoda 4 Asteroidea: Oreasteridae 4 Enteropneusta 4 212 MEMOIRS OF THE QUEENSLAND MUSEUM Code No. Species Systematic position Number Main Collected site group 284 Paraplagusia unicolor (Macleay) Pisces: Cynoglossidae 4 S 285 Leanira yhleni Malmgren Polychaeta: Aphroditidae 3 s 286 aphroditid 7 Polychaeta: Aphroditidae 3 287 phyllodocid 4 Polychaeta: Phyllodocidae 3 288 phyllodocid 12 Polychaeta: Phyllodocidae 3 289 phyllodocid 14 Polychaeta: Phyllodocidae 3 290 phyllodocid 19 Polychaeta: Phyllodocidae 3 s 291 syllid 1 Polychaeta: Syllidac 3 292 spionid 6 — polyspecific Polychaeta: Spionidae 3 N 293 = species 216 (Nerinides sp.) Polychaeta: Spionidae 3 294 Ophelina gigaritea Rullier Polychaeta: Opheliidae 3 295 capiteilid 2 Polychaeta: Capitellidae 3 296 sabcllid 6 Polychaeta: Sabellidae 3 s 297 unknown polychaete Polychaeta 3 298 Gynodiastylis sp. Cumacea: Diastylidae 3 299 tanaid 7 Tanaidacea 3 s 300 mysid 2 Mysidacea 3 301 mysid 3 Mysidacea 3 302 Squilla laevis Hess Crustacea: Stomatopoda 3 s 303 Lysiosquilla perpasta (Hale) Crustacea: Stomatopoda 3 304 ?Pontophilus sp.3 Decapoda: Crangonidae 3 N 305 Conchocetes artificiosus (Fabricius) Decapoda: Dromiidae 3 306 Dorippe australiensis Miers Decapoda: Dorippidae 3 307 Actumnus squamosus de Haan Decapoda: Xanthidae 3 s 308 Achaeus lacertosus Stimpson Decapoda: Majidae 3 s 309 Sanguinolaria sp. Bivalvia: Sanguinolariidae 3 310 gastropod 1 Gastropoda 3 3 1 1 gastropod 7 Gastropoda 3 312 gastropod 19 Gastropoda 3 N 3 1 3 gastropod 22 Gastropoda 3 N 314 gastropod 26 Gastropoda 3 315 gastropod 29 Gastropoda 3 316 Artiphipholis squamata (Delle Chiaje) Ophuiroidae: Amphiuridae 3 S 317 Lovenia camorata H. L. Clark Echinoidea: Spatangidae 3 318 holothurian 5 Holothuroidea 3 Mem. QdMus. 18(2): 213-7. [1978] A NEW SPECIES OF PONTELLOPSIS (COPEPODA, CALANOIDA) FROM MORETON BAY, QUEENSLAND J. G. Greenwood University of Queensland ABSTRACT A new species, Pontellopsis tasmanensis sp. nov., found in coastal waters of eastern Australia is described and figured, and the relationship of this to allied species is discussed. A study of the zooplankton of Moreton Bay, Queensland (Greenwood 1976) revealed the presence of males and females of a species of Pontellopsis which could not be allocated to any known species and which is described here. All specimens were collected with either Clarke-Bumpus or 40 cm diameter conical nets of nylon mesh with an aperture size of 195 microns. Dissections were made in polyvinyl alcohol lightly stained with chlorazol black, and drawings made with the aid of a Wild M20 drawing tube. Pontellopsis tasmanensis sp. nov. (Figs 1 a-j, 2 a-f) Materiai. Examined HolotypE: Queensland Museum, W520I, J, Rainbow Channel, Moreton Bay (27° 20’S., 153° 15’E.), SE.Q., coll. 8. vii. 1964, J. G. Greenwood (Station 1 of Greenwood 1976). ALLOTYPE: W5202, same data as Holotype. Paratypes: W5203, 5c3c3, Sjj, same data as Holotype. P. tasmanensis was taken on 45 occasions during 1963-66 from Rainbow Channel, central Moreton Bay, and Cleveland Channel (Stations. I, II, and 111 of Greenwood 1976). Males were taken on 25 occasions, females on 29 occasions, males with females on 9 occasions. Material other than the type series was examined in less detail and discarded. Description of Female (Fig. 1) Body very robust. Total body length varying between 142-1 *69 mm (av. 10 individuals 1-56 mm). Lenth ratio of metasome to urosome 2-98:1; metasome length to width ratio 1-62:1. Metasome with fourth and fifth thoracic segments fused; posterior margins of fifth segment produced symmetrically into small rounded posterolateral processes. (Fig. 1 a, b, c). Urosome two-segmented, genital segment approx. 1-47 limes length of anal segment. Genital segment asymmetrical, right postero-dorsal region produced into rounded or broadly triangular lobe (Fig. 1 d, e) which extends laterally to maximum distance at level of posterior third of segment. This lobe truncated abruptly posteriorly, at right angles to body axis. Degree of development of this lobe shows some variability. Most individuals as in Fig. 1 d, but others with larger lobes, to the maximum found as in Fig. 1 e. Forms with larger lobes sometimes with a small seta at tip of lobe and similar small seta on mid-ventro-lateral surface. No other spines or projections present on genital segment. Anal segment produced postero-dorsally into large, slightly asymmetrical triangular supra-anal plate reaching to posterior level of furcal rami (Fig. 1 d, e). Furcal rami symmetrical, length c. twice width. Fifth legs asymmetrical (Fig 1 f, g). Right exopod with stout spine developed from inner border slightly distal to midpoint, not reaching end of ramus; corresponding spine absent from left exopod. Both exopods terminate in single point. Right exopod (Fig 1 h) with two subterminal spines on outer border, the more distal the larger; two minute spinules spaced more proximally along outer border. Left exopod (Fig I i) with cluster of three subterminal spines, two forming lateral borders of small plate on which third is mounted; 214 MEMOIRS OF THE QUEENSLAND MUSEUM GREENWOOD: A NEW SPECIES OF PONTELLOPSIS 215 two minute spinules more proximally along outer border, first being just proximal to the ‘plate'. Endopods similar, bifid terminally, but left endopod narrower and longer, about 1-3 times length of right endopod. Description of Male (Fig. 2) Body very robust. Total body length varying between 1-28-1 -54 mm (av. 10 individuals 1-36 mm). Length ratio of metasome to urosome 2-58:1, metasome length to width ratio 1-96:1. Metasome with fourth and fifth thoracic segments fused; posterior margins of fifth segment rounded on left side, right side produced as elongate spine-like process reaching to mid or posterior border of fourth urosomal segment (Fig. 2 a, b). Urosome five-segmented, segments 1-5 in proportions 40:15:17:12:16 (total 100). Genital segment symmetrical, with single small seta on right postero-lalcral margin (Fig. 2 c). Second segment with patch of minute tubercles on right postero-lateral surface. Third segment slightly asymmetrical, slightly swollen on right side, apex of swelling with minute tubercles. Fourth and fifth segment without spines or tubercles. Fifth legs as figured (Fig 2 d, e, f). Right leg with thumb-like process from base of proximal exopod segment (Rel) similar in length to finger-like distal exopod segment (Re2) thumb slightly expanded or spathulate distally; large seta on Rel distal to base of thumb. Re2 with excavated lateral surface bearing one larger, one smaller seta; small terminal, longer subterminal seta; minimum distance between origins of thumb and finger similar to length of segment. Left leg with distal protopod segment (B2) and Rel of similar length. Disto-lateral margin of Rel with stout spine extending c. two-thirds length Re2. Re2 short, c. half length Rel; length twice width; lobe on medial face clothed with fine setae; one long one short seta terminally. Remarks During much of the survey in which these specimens were taken, the male and female forms described here were regarded as belonging to different species, the male being provisionally related to P. tenuicauda and the female regarded as a new form related, by structure of the fifth legs particularly, to P. macronyx, P. scotti and P. tenuicauda. Subsequent realisation of the coincidence in frequency and occurrence pattern of these two strongly suggested conspecificity. There were, for example, no female P. tenuicauda yet the apparent males occurred on 25 occasions. More detailed examination of the two forms showed some differences between these males and those of P. tenuicauda as described by Mori (1937), and considerable differences between the present females and those of P. tenuicauda sensu Mori (1937). and particularly sensu Giesbrechl 1892 (from which Mori’s also show differences). Designation of the new species was therefore necessary. Both male and female share some characteris- tics with several related species, but are distinguished from them on combinations of features. These distinguishing features are discussed below, firstly for females, subsequently for males. FEMALES: The asymmetrical fifth legs of female P. tasmanensis show some resemblance to those of the following species: P. pexa Scott, P. yamadae Mori, P. macronyx Scott, P. scotti Sewell, P. tenuicauda (Giesbrecht): P. pexa differs in that the right exopod there terminates in three small spines, not a single point, and although a spine is developed from the inner border it arises distally and extends well beyond end of the ramus (Scott 1909; p 173, pi. 54 fig. General body form is similar to that of P. yamadae. but although the female fifth legs of that species are asymmetrical they differ from the present form in having one (or more) large inner spines from both exopods (Mori 1937, p. 98, pi. 47 fig. 6; pi. 48 fig. 13. Brodsky 1962, p. 147, pl. 48). P. yamadae is also a much larger calanoid (2-8 mm, Mori 1937; 2-45 mm, Brodsky 1962). Form of the fifth leg is very close to that of P. macronyx and P. scotti. However in those species the endopods are of similar length and the exopods are not (reverse of P. tasmanensis). the right exopod being shorter than the left (Scott 1909, p. 173, pi. 54 fig. 6; Sewell 1932, p. 388, fig. e). P. macronyx further differs in armature of the genital segment, which has paired dorso-lateral spines both anteriorly and posterior- ly, in having a smaller anal plate, and it is a slightly larger copepod (1-73-1-97 mm), (Scott Fig 1 Pontellopsis tasmanensis sp. nov., female, a, lateral view; b, dorsal view; c, Th5 and urosome with P5 in situ lateral view d, urosome with typical development of genital segment; e, urosome with maximal, development of genital segment (as in ‘e’ above); h, detail of terminal region right P5 exopod; i, detail terminal region left P5 exopod, j, proximal portion of right antennule. 216 MEMOIRS OF THE QUEENSLAND MUSEUM 1909; Sewell 1932; Kasturirangan 1963). P. scotti differs additionally in having: an elongate anal segment without extended anal plate, and on which the furcae are set asymmetrically; paired lateral spines anteriorly on the genital segment, the posterior end variously developed but typically with a swelling on the left side (elongate in some), some with rod-like spine on right side (Sewell 1932, fig 129a)- Closcst similarity in fifth leg structure is with P. tenuicauda. However in that species the genital segment does not develop a bulbous postcro-lateral projection, having instead two lateral spines on the right side (some variability is apparent in development of these spines, cf. Giesbrecht 1892, pi. 41 fig. 43, and Mori 1937, pi. 46 fig. 8). P. tenuicauda also has a small anal plate (not reaching mid-region of furcae), and very small Fi(.. 2. Pontellopsis tasmanensis sp. nov., male, a, dorsal view; b, lateral view; c, urosomal segments 1-4; d, P5; e, terminal portion left P5; f, anterior view, right P5. GREENWOOD: A NEW SPECIES OF PONTELLOPSIS 217 fifth thoracic segment lobes (see espec. Giesbrecht 1892). Sherman (1964) took females of a Pontellopsis sp. from the central South-Pacific which could not be identified with published descriptions but which were similar to those of P. macronyx. No further details of that species were given by Sherman, but his females must be similar also to those of P. tasmanensis. MALES: P. tasmanensis males bear close resemblance to those of P. scotti, P. yamadae and P. tenuicauda. They differ from P. scotti in relative lengths of the 'thumb and finger' (Re2) of the right fifth leg; absence of the central swelling found in the right fifth thoracic segment process of P. scotti (Sewell 1932, p.390, fig. 129 b, f). P. yamadae males are almost twice the size of the present forms (2-5 mm, Mori 1937, p. 99) and differ in some further details: Right fifth leg ‘thumb’ shorter than ‘finger' in P. yamadae (slightly longer in P. tasmanensis) and without spathulate tip; distal spine on Rel of left fifth leg very short in P. yamadae. P. tenuicauda is only slightly larger than P. tasmanensis (c. ! -65 mm, Mori 1937, p. 98) and, according to Mori’s description and figure, closely similar. It differs in that: The right fifth thoracic segment process is longer, reaching to posterior border of anal segment; the second urosomal segment is asymmetrical with pronounced swelling on right side similar to that on the third segment; a large seta is borne on the base of the thumb; the thumb is not spathulate distally. Mori (1937, pi. 46 fig. 10) did not show any spine distally on the left Rel, but this may have been an omission. LITERATURE CITED Brodsky, K. A., 1962. On the fauna and distribution of Calanoida of surface waters in the north-western part of the Pacific Ocean. Issled. dalnevost. Morei SSSR 8: 91-166. GiLSBRtCHT. W., 1892. Systemalik und Faunistik dcr Pelagischen Copepoden dcr Golfes von Neapei. Fauna Flora Golf. Neapei, Mongr. 19: 1-831. Grei-nwood. J. G., 1976. Calanoid copepods of Moreton Bay (Queensland) I. Families Calanidae, Eucalan- idae and Paracalanidae. Proc. R. Soc. Qd 87: 1-28. Kasturirangan, L. R., 1963. A key for the identification of the more common planktonic copepoda of Indian coastal waters. Ind. Nat. comm, on Oceanic Res. Publ. 2: 1-87. Mori. T., 1937. The pelagic Copepoda from the neighbouring waters of Japan, pp. 1-150. (Reprinted 1964 by Soyo Co. Inc; Tokyo). Scott. A., 1909. The copepoda of the Siboga Expedition, Part 1. Free-swimming, littoral and semi-parasitic Copepoda. Sibioga Exped. Mongr. 29a: 1-323. Sewell. R. B. Seymour. 1932. The Copepoda of the Indian Seas. Mem. Indian Mus. 10: 223-407. Sherman. K., 1964. Pontellid copepod occurrence in the central South Pacific. Limnol. Oceanogr. 9(4); 4 ' 76 - 84 . Mem. Qd Mus. 18(2); 219-31, pis. 31-8. [1978] ASTHENOPSIS WHITEHOUSE, 1939 (TRIEOBITA, MIDDLE CAMBRIAN) IN NORTHERN AUSTRALIA Peter A. Jell University of Queensland ABSTRACT Comparisons of the Trilobite Asthenopsis with other genera, previously confused with it, show it to be a distinct taxon containing six Australian species. Apart from the type species, Asthenopsis levior Whilehouse, 1939, five are described as new {queenslandica. rhinostrongyla. unquinsepta. butorosa, and opalensis). Another new species is recognized but not named and another population is compared with A. rhinostrongyla. Some remarks are also made on muscle insertions. Since Whilehouse erected the genus in 1939 and introduced Asthenopsis levior as type, no new species have been assigned and those European species placed there by Whilehouse may now be more correctly placed in other genera. Of the species from China referred by Walcott (1913) to Solenopleura, by Kobayashi (1935) to Solenoparia. and by Whilehouse (1939) to Asthenopsis. none belong to Asthenopsis. No references to Asthenopsis have appeared since 1939 other than the short note (Opik 1967, p.l84) indicating its ptychoparioid affinities and several cursory identifications at the generic level. Terms used in the systematics are those defined by Harrington, Moore, and Stubblefield {in Moore 1959), glabella being used to include the occipital ring. Notation for muscle areas of glabella and points on facial suture follow Richter and Richter (1940). 1 have referred to any sagittal or exsagiltal dimension in terms of length and any transverse dimension in terms of width. Figured material is housed in the Common- wealth Palaeontological Collection (CPC), Department of Geology, Australian National University (ANU), or Department of Geology, University of Queensland (UQF). Locality numbers referred to are in the registers of the Bureau of Mineral Resources, Canberra (M, H, or MNF), Department of Geology, Australian National University (ANU), or Department of Geology, University of Queensland (UQL). Genus Asthenopsis Whilehouse, 1939 Type Species; Asthenopsis levior Whilehouse, 1939 by original designation. Generic Diagnosis Ptychoparioid with glabella of slightly variable length, three or four pairs of lateral glabellar furrows, an anteromedial glabellar muscle scar, slightly tapering, straight to very slightly waisted lateral glabellar margins. Furrow Ip long and forked adaxially. Fixed cheeks w'idc (0-7 - 0-9 of basal glabellar width). Eye ridges paired or tripled. Anterior axial furrow weakly or not impressed, always much weaker than lateral axial furrow, and shallowing through ontogeny. Fossulae deep. Brim 0-2 - 0-5 of glabellar length. Border furrow with wide pits anterior to the fossulae. Palpebral lobes short and near midlength of cephalon or slightly posterior to it. Free cheek with high eye socle, with strong anterior doublural projection, and genal spine that is commonly deflected slightly laterally to break the cephalic marginal curve at the level of the posterior of the palpebral lobe. Thorax of 14 segments. Pleural lips either rounded, broadly squared or spinose. Pygidium relatively small (c. 0-3 of cephalic length), transverse, and moderately vaulted to flat in pleural areas. Border furrow poorly to not impressed. Doublure no wider than border, usually 220 MEMOIRS OF THE QUEENSLAND MUSEUM very much less. Ornament highly variable from species to species including smooth, finely papillose, and tuberculate (both fine and coarse). Comparison Solenopleura Angelin, 1854 with holometopa as type is a distinct genus characterized by the lack of preglabellar field, glabella occupying a larger percentage of the cranidial surface, fixed cheek 0-6 or less of basal glabellar width (canaliculata is probably not congeneric), almost aspinose cheek, and more quadrangular pygidium. Jincella Snajdr, 1957 with S. prantli Ruzicka, 1944 as type is distinguished from Asthenopsis by the short convex border, short preglabellar field, well impressed anterior axial furrow, rather conical glabellar shape, narrow fixed cheeks, well impressed glabellar furrows, and long narrow posterior limb. Solenoparia Kobayashi, 1935 with Ptychoparia toxeus Walcott, 1905 (Walcott 1913, pi. 19, fig. 10) as type is distinguished by its short convex border, narrow fixed cheeks, longer palpebral lobes, glabella occupying larger part of cranidial area, and less transverse pygidium. Kounamkites Lermontova in Chernysheva, 1956 based on virgatus Chernysheva, 1956 is distin- guished by the short border relative to brim, strongly diverging anterior branches of suture, well impressed anterior axial furrow, undeflected genal spines, and less transverse pygidium. The content and validity of the several genera mentioned above are not commented on herein as 1 have had no first hand experience with them. However, they are sufficiently known to be sure of their distinction from Asthenopsis. Eosoptychoparia kochihei (Walcott, 1911) from Manchuria represents a very similar genus even to the deflected genal spine but distinctions are evident. I shall describe E. kochibei more fully in a forthcoming paper. Musculature As illustrated for Triarthrus (Cisne, 1974, fig. 2) the ventral longitudinal muscles of Asthenopsis are interpreted as running forward to the two wide slightly deeper areas of the anterior border furrow situated directly in front of the fossulac. These muscles that run back to the major ventral endoskeletal bars to which attach many of the muscles of the mouth parts and other appendages, are probably strongly digitate anteriorly (by homology with the cephalocarid Hutchinsonieila, see Hessler 1964) with many small attachment sites on the cuticle. In passing it should be noted that a large percentage of trilobites have some structures in or near the anterior border furrow (i.e. low anteriorly on the sagittal or exsagittal glabellar profile) that can be readily interpreted as ventral longitudinal muscle attachment sites. Those lacking such structures are not precluded from having muscle insertions in this position but are simply interpreted as not showing them in any distinctive way. In the Crepicephalidae the two or three wide distinctive pits in the border furrow are formed by several smaller circular pits providing apodemes for the digitations of the ventral longitudinal muscles. This structure also suggests that broad depressions and discrete pits may be analogous. Many post-Cambrian trilobites exhibit strong apodemes or muscle scars well forward on the cephalon that would be ideally placed for attachment of the ventral longitudinal muscles. The dorsal longitudinal muscles of Asthenopsis are interpreted as anchoring anteriorly to the glabellar furrows and posteriorly to the apodemes of the transaxial furrows. While these muscles form single sheaths the full length of the animal, they are also attached to each segmental apodemal area to allow intersegmental mechan- isms to function. A very small muscle scar is situated anteromedially on the glabella. I am unable to find the description of any other ptychoparioid referring to a similarly situated muscle scar. Jell (1975, p, 23) interpreted a muscle scar in this position on a species of Pagetia and discussed its possible function as a hypostomal diductor muscle by direct homology with the living notostracan Triops. In many dalmanitaceans a pit is present anteromedially on the glabella (Campbell in press) which may be homologous with that of Pagetia and indeed with that of Asthenopsis. Affinities The taxonomic position of Asthenopsis is not clear. Originally placed in the Solenopleuridae it has been regarded as correctly assigned except for the statement by Opik (1967, p. 184) that it may be regarded as a ptychopariid. I agree with Opik and believe that in the light of new species described herein the genus can be confidently divorced from any close relationship with the group of genera that have come to be known as the Solenopleuridae. I shall not make any assignment at this stage as I am at present engaged in a numerical taxonomic study of ptychoparioids and do not wish to pre-empt those results. JELL; MIDDLE CAMBRAIN TRILOBITA 221 Asthenopsis levior Whitehouse, 1939 (Fig. la; Plate 31, figs. 1-9; Plate 32, figs. 1-5) Asthenopsis levior Whitehouse, 1939, p. 214, pi. 22, figs. 17-20. MaTI RIAI Ewminfd Hol.oTVPi' UQF3337, an external mould of a complete exoskeleton the counterpart of which is missing from the University of Queensland collections, from V Creek at the Undilla to Camooweal Road crossing. Otiiir Mati RIAI Four cranidia (C), I free cheek (FC), 2 pygidia (P), and 1 thorax (T) from M41 at the base of an isolated hill of Split Rock Sandstone 4-8 km east of Douglas Creek on the Old Burketown Road (lat. 19° 27-5 'S., long. 138° 37 'E); 3C, 2P. and one complete from M247 at Barkly No. 7 Bore west of Yelvertoft; 2C from the left bank of Douglas Creek halfway from the Old Burketown Road to the O’Shannassy River; 3C from 12 km south of Douglas Spring on the ‘Morslone’ to ‘Undilla’ Road; IC and 2T from left bank of Douglas Creek at ‘Morstone’; 2C from M52 1-3 km west of M41; IFC from Ml 39 just west of Top Hands Waterhole on the upper reaches of Harris Creek; 5C. I P, and 3T from UQL469 on top of hill immediately east of the crossing of Harris Creek by the Camooweal to Thorntonia Road; 3C and IFC from UQL3510 15 km south-west of the Thorntonia airstrip on the Kangaroo Flat road. This includes the figured specimens CPC 17064-8 and UQF68711-4 (Pis. 31, 32). Whitehouse's originally figured specimens were also examined and are refigured herein (PI. 31 figs. 1, 4, 6, 7). All localities are in the V Creek Limestone except M52 which is in the Mail Change Limestone. The age of the material is the Zone of Ptychagnostus nathorsti. Diagnosis Outline regularly oval. Surface ornament lacking or of minute papillae with rare slightly larger tubercles. Glabella extending 0-7 of cephalic length, with tapering straight or convex lateral margins, with truncated anterior and angular anterior corners. Three pairs of lateral glabellar furrows, smooth to very poorly impressed, with furrow Ip long and wide, furrows 2p and 3p successively shorter and narrower. Occipital node at midlength of ring. Border flat to weakly convex long and tapering laterally. Facial suture with slightly convex anterior branch in exsagittal line or slightly convergent. Eye ridges variable in prominence, paired. Palpebral lobe short, just behind midlength of glabella. Caecal ornament on extra-glabellar parts of cephalon. Thorax of 14 segments. Pleural furrows long, well impressed, running almost to pleural tips. Pleurae downturned in fulcral line, with spinose tips. Pygidium about 0T5 of total length. Axis with three rings and terminus. Pleural areas highly vaulted with four pairs of ribs. Posterior margin concave. Description The holotype designated by Whitehouse (1939, p. 278) has a cephalon 9 mm long but other specimens have been collected with cephala up to 16 mm long so the type is not a fully mature individual. This description is based on the more mature material and its variations from the morphology of the holotype are discussed below under DISCUSSION OF Holotype. Exoskeleton almost twice as long as wide, moderate overall convexity with steeper slopes towards the margin. Cranidium two or three times length of pygidium. Overall shape oval but somewhat more rounded anteriorly than posteriorly. Cranidium moderately vaulted, twice as wide as long, with moderately steep slope to border furrow in lateral profile. Glabella tapering forward to half its basal width, with straight or slightly convex sides, truncated almost transverse anterior. Anterior axial furrow very poorly impressed (represented almost entirely by a change of slope in lateral profile), never as well impressed as lateral axial furrow. Axial furrow very broadly V shaped in section, with deep fossulae anteriorly, floor raised adjacent to lobe Ip, running directly past occipital ring to margin. Occipital ring five times as wide as long, with prominent sagittal node at the midleigth, posterior margin convex, tapering laterally into a low ridge that crosses the axial furrow into the posteroproximal corners of the fixed cheek. Occipital furrow poorly impressed, slightly deeper laterally, with smooth surface of apodemal pit extending up onto lobe Ip and occipital ring. Glabellar furrows three pairs in number, defined mainly as smooth areas on an otherwise weakly ornamented glabella. Furrow Ip long, directed at 30° to transverse line out of axial furrow then posteriorly at 60° to transverse line in a shorter adaxial terminally expanded section, with medial swelling just out of axial furrow. Furrow 2p shorter and narrower than Ip, of same general shape but with much narrower unexpanded adaxial part and directed less posteriorly. Anteromedian glabellar muscle scar present. Brim downsloping much more steeply and twice as long at facial suture as axially, convex becoming more so laterally, very slightly depressed from fossulae forward to border furrow, with prominent anastamosing caeca originating out of the eye ridge and running forward into the 222 MEMOIRS OF THE QUEENSLAND MUSEUM Fkj. 1 A: Reconstruction of Asthenopsis levior Whitehouse, 1939 based on holotypc and UQF44314. border. Border furrow poorly to moderately impressed, with a pair of very wide indistinct pits directly anterior to the fossulae (ventral longitudinal muscle attachment sites), transverse except for slight convexity between wide pits. Anterior border moderately convex often accen- tuated by sharp posterior slope but always with gently sloping anterior, highest point at midlength. Eye ridges double, prominent, continuing through the fossulae and around front of glabella as low parafrontal band, directed posteriorly at approx- imately 20° to transverse line. Palpebral lobe 0-25 of cephalic length, abaxially upturned, and posteriorly downturned, moderately arcuate, opposite furrow Ip and lobe 2p. Palpebral furrow moderately impressed, shallowing at midlength. Fixed cheeks 0*82 basal glabellar width, moderately convex, rising gently to palpebral lobe. Posterior limb wide and long, almost rectangular in shape except for slightly convex and posteriorly directed anterior margin, sloping steeply abaxial- ly, with faint caecal system posterolaterally from palpebral lobe. Posterior border furrow lengthen- ing and deepening laterally, turned slightly forward at extremity. Posterior border lengthening and flattening laterally, with marked ridge forming high point for most of its width but meeting the posterior margin two thirds of distance from fulcrum to u'. Posterior margin straight to fulcral line (used in sense of Opik, 1967, p.57) then convex. Facial suture with o- n only 0-6 of /?- v variable from convex and converging to straight and exsagittal, (S at midlcngth of palpebral lobe, e at posterior of furrow Ip. Free cheek with long anterior extension where facial suture cuts obliquely across border, y^at posterior of border furrow. Margin evenly curved to base of rounded gcnal spine, then deflected slightly abaxially down spine. Border wide, flat to weakly convex. Eye socle low, of JELL: MIDDLE CAMBRAIN TRILOBITA 223 B Fig. IB: Reconstruction of Asthenopsis queenslandica sp. nov. based on hoiotype and CPC 17072. constant height. Genai field exhibiting very low anastamosing caeca. Rostral plate and hypostoma unknown at present. Thorax of 14 segments each 12 times as wide as long. Articulating half ring slightly lower than axial lobe, shorter than furrow or lobe, with prominent transverse ridge near posterior forming high point and dividing smooth relatively flat anterior part from steeper granulose posterior part. Articulating furrow with posterior wall steeper and higher, with fine papillose ornament anteriorly. Axial lobe expanding slightly laterally to occupy full segmental length, flat (sag.) with transverse posterior margin. Accessory apodemal pits low abaxially, joining articulating furrow and posterior margin. Pleural furrow occupying 40-50% of fixed pleura, swinging forward and shallowing adaxially, meeting axial furrow a short distance from margin, with very shallow posterior arm adaxially giving a forked appearance. deepening and shortening on free pleura before finishing at the midlength near the base of the pleural spine. Pleural strips of equal length except on free pleura where anterior strip shortens as it runs obliquely behind wide moderately long smooth facet. Pleural spines stout, curving slightly posteriorly. Segments not overlapping at all on fixed pleura (i.e. margins abutting) but overlap- ping almost all facet on free pleura. Segments transverse throughout. Pygidium almost three times as wide as long, well vaulted, with rounded margin except for slightly concave section across axis posteriorly. Articulating half ring extending in front of otherwise transverse margin, with fine papillose ornament posteriorly. First axial ring highest part of pygidium, with posterior excavation on axial three quarters cutting out most of segmental length (sag.) with weak accessory apodemal pits in exsagittal line close to axial furrow. Second 224 MEMOIRS OF THE QUEENSLAND MUSEUM axial ring with almost imperceptible accessory apodeme and smaller posterior excavation. First and second iransaxial furrows moderately impressed with wide undepressed apodemal areas. Posterior transaxial furrows possibly three in number, progressively weaker, and continuous. Axis parallel sided with rounded posterior, with moderately steep posterior slope to border and margin. Pleural areas with three well impressed pleural furrows and one poorly impressed interpleural furrow anteriorly, furrows deepest in fulcral line. Caecal ornament running out from axial furrow along pleural strips, through border furrow and into the border. Facets wide, relatively short, triangular, smooth and steeply sloping. Anterior pleural strip of first segment short and convex to fulcrum then longer, less convex and oblique beyond. Border furrow poorly impressed more so behind axis. Border flat and of uniform width. Ornament of fine papillae on axial rings, terminus, pleural ribs and border of some specimens, furrows smooth. Discussion of Holotype As already mentioned the holotype specimen represents an immature individual so it varies in several minor respects from the description given above. With the recognition of these morphogene- tic changes further outlined under Morphogeny below comes the realization that the species has considerable intraspecific variation between individuals of the same and only slightly different size. The holotype is crushed with only slight displacement of the free cheeks that do however, obscure the course of the facial suture. It has fine, close-spaced papillae with an occasional very widely spaced tubercle. Details of occipital ring and glabellar posterior are lost as the exoskeleton of that part has remained in the counterpart. Fossulae are deeper so the frontal area is more convex (exsag.) but less steeply sloping. The caecal network on the brim and eye ridges is less prominent. The palpebral furrow is abaxially convex and the lobe more strongly upturned. Fixed cheek width and basal glabellar width almost the same. On the thorax the ornament is coarser axially but less apparent on the pleura. On the pygidium the posterior margin is straight, the interpleural furrow is very weak, the axis does not reach so close to the margin, and the pleural areas are not so vaulted. Morphogeny The smallest individual with a complete thorax (Plate 31, fig. 2) has 14 segments and a cephalon 4 mm long so that it must be a very early holaspide. A slightly smaller cranidium (Plate 31, fig. 3) 3-5 mm long is similar to that of the complete individual. While the cepahlon is shorter than 5 mm the fixed cheeks are more convex, the axial furrow is deeper, the ornament is coarser, the furrows forward from the fossulae are more evident, the palpebral lobes are relatively long extending from lobe Ip to furrow 3p or half glabellar length, pleural tips are not spinose but simply pointed. Individuals with cranidia 5-10 mm long are characterized by the holotype described above but it should be noted that most individuals of this size range have coarser papillae than the holotype. Affinities Asthenopsis levior is quite variable from locality to locality but only in minor details of glabellar shape, ornament, relief, and anterior course of facial suture. As no uniform variation of these characters can be observed and as the variation is not as great as between this and other species of the genus described below no other species have been separated from the relatively distinct (within the genus) levior. Asthenopsis queenslandica* sp. nov. (Fig. lb; Plate 33, figs. 1-8; Plate 34, fig. 1) Material Examined Holotypf: CPC 1 7074, a complete specimen from M243 on the flood plain of Whistler Creek, 0-8 km from its junction with the Buckley River, 42 km west of Yelvertofl Homestead (Lat. 20° 04' S.. long. 138° 30' E): V Creek Limestone. Zone of Ptychagnostus nathorsti. Other Material; Three complete specimens, 14C, 2FC and 3P including CPC17070-3 and 17075-8 (Pis. 33 and 34) from M243; 2C from UQL467 4-8 km west of Redbank Creek, just west of laterite ridge between Redbank and Harris Creeks; 20C, IFC, 8P, and several thoracic segments from UQL463 top of hill 6 km south of Thorntonia Homestead on left bank of West Thornton River. Both UQL463 and 467 are in the V Creek Limestone with 463 belonging to the Euagnostus opimus Zone and 467 to the Ptychagnostus punctuosus Zone. Diagnosis Asthenopsid with relatively square anterolateral glabellar corners, low cephalic convexity generally and in the anterior margin, four pairs of smooth ♦Named for the state of Queensland. JELL: MIDDLE CAMBRAIN TRILOBITA 225 to very poorly impressed lateral glabellar furrows, anteriorly placed occipital node, relatively long brim, doubled eye ridges, relatively narrow fixed cheeks, straight slightly converging anterior branches of facial suture, fine close spaced papillose ornament with sparse small tubercles superimposed. Pleural tips squared. Transverse pygidium having flat pleural areas, no border furrow, and an evenly curved margin. Description Cranidium four times as long as pygidium, very weakly vaulted. Glabella standing above flat to slightly abaxially rising fixed cheeks in anterior profile, subquadrale tapering only slightly forward from furrow Ip, with angular anterolateral corners and only slightly convex anterior margin. Occipital ring with small sagittal node anteriorly. Occipial furrow well impressed, short, and smooth (apodemal part) with steeper anterior than posterior wall laterally and shallow, slightly longer, with papillose ornament and steeper posterior than anterior wall axially. Glabellar furrow 4p may be present low in the axial furrow. Glabellar ornamented with fine papillae except on the apodemal areas. Brim downsloping slightly more steeply at the facial suture than axially. Eye ridges only slightly elevated, divided into two parallel ridges. Fixed cheeks 0-73 of basal glabellar width, rising gently to palpebral lobe with fine papillose ornament and a very few slightly larger tubercles interspersed, with very faint caecal network radiating from lobe Ip and from the rear of the eye. Posterior limb elongate triangular in shape, sloping steeply down abaxially. Facial suture with ji - 7 straight and very slightly converging, tS behind midlength of palpebral lobe. Free cheek relatively flat, not vaulted, ji at posterior of border. Border convex anteriorly, flatter posteriorly, with one or two continuous parallel terrace lines near the margin, with papillose ornament. Genal spine with flat dorsal and slightly rounded ventral surfaces. Thorax of 14 segments each 15 times as wide as long. Each pleuron with fine papillae and single row of tubercles (up to 7). Free pleura with rounded anterolateral and semisquared posterolateral corners. Anterior fixed pleural margin just under posterior of next anterior segment but with margins directly opposed in same plane for short distance at fulcrum then overlapped on the free pleural facets. Segments becoming slightly anteriorly convex near the pygidium. Pygidium with an unbroken marginal curve, and a low profile with axis standing only slightly above flat pleural areas. Furrows all shallower than in A. levior, decreasing in depth posteriorly and laterally. Border furrow not impressed. Border may be defined as area beyond pleural furrows. Fine papillae on areas out of furrows and off muscle scars. Morphogeny Although no small specimens with a full complement of segments are available some comments on the morphological changes with increased cranidial size are pertinent. The adults described above all have cranidial length of 10-12 mm whereas two other groups exist with cranidial lengths of 3 mm and 5-6 mm respectively. The 3 mm cranidia (Plate 33, figs. 2, 4) have a second order of larger tubercles .scattered over the entire exoskeleton giving the same ornament as in mature individuals of A. rhinostrongyla. They have an anterolaterally rounded glabella, almost no glabellar furrows, better impressed anterior axial furrow, convex fixed cheeks Just slightly more than half basal glabellar width and with no apparent division of the eye ridge. The 5-6 mm cranidia (Plate 33, fig. 7) have the same ornament as the 3 mm cranidia, the anterolaterally rounded glabella, very poorly impressed glabellar furrows, virtually no anterior axial furrow, convex fixed cheeks 0-6 of basal glabellar width, and paired eye ridge. Astbenopsis rhinostrongyla* sp. nov. (Fig. 2A; Plate 34, figs. 2-7; Plate 35, figs. 1, 2) Matf-rial Examined Holo TYPE CPC 1 7080. a cranidium from H 1 38 on the Huckitla 1:250,000 Geological map at 22°35’ south latitude, 136° 02’ east longitude on Arthur Creek, Northern Territory; Arthur Creek Beds, Euagnostus opimus Zone. Other MATF.RtAL: Seven C, 3 FC, 1 T, and 3 P including CPC 17079, 17081-6, (Pis. 34 and 35) from H138. Diagnosis Asthenopsid with rounded anterolateral glabel- lar corners, flat unarched anterior border, moderate convexity, anterior branches of facial suture diverging forward and abaxially convex, relatively long pelpebral lobes, ornament of very close spaced papillae over entire surface except apodemal areas, palpebral lobes and furrows, and coarse perforated tubercles interspersed sparsely ^Rhinos (Gr.) nose, strongylus (Gr.) rounded, refers to the rounded glabellar anterior. 226 MEMOIRS OF THE QUEENSLAND MUSEUM Fig. 2: Reconstruction of cephalon and pygidium of Asthenopsis species. A: Asthenopsis rhinostrongyla sp. nov. based on the holotype, CPC 17085, and CPC 17086. C: Asthenopsis butorasa sp. nov. based on the holotype, ANU30646, and ANU30647. B; Asthenopsis opalensis sp. nov. based on the holotype, CPC17100, and CPC17102. D: Asthenopsis unquinsepta sp. nov. based on the holotype, CPC17087, and CPC17088. JELL; MIDDLE CAMBRAIN TRILOBITA 227 over the same areas. Pleural tips rounded. Transverse pygidium with flat pleural areas. Description Cranidium with moderately steep anterior slope in lateral profile. Glabella subquadrate tapering only very slightly forward to 0-8 of its basal width. Occipital furrow with markedly deeper apodemal pits, almost dumb-bell shaped apodemal areas extending onto lobe Ip and occipital ring. Four pairs of lateral glabellar furrows smooth, not depressed. Furrow Ip with adaxial part not expanded. Furrow 3p separated from axial furrow. Furrow 4p very indistinct, narrow and smooth. Occipital node situated anteriorly. Axial furrow distinctly shallower adjacent to lobe Ip and occipital ring. Eye ridge quite low and formed of two trunks. Fixed cheeks 0-8 of basal glabellar width, slightly convex and horizontal. Posterior limb steeply downsloping. Facial suture with (> - ji convex and diverging. Posterior border furrow lengthened slightly laterally. Posterior border only moderately convex and slightly longer laterally, with its ridge not markedly distinct and not obvious beyond fulcrum. Ornament of papillae and sparse perforated tubercles over cranidium except on furrows, palpebral lobe and distal parts of posterior limb. Free cheek doublure without ornament, extending only slightly adaxial to long anterior dorsal projection. Thorax of at least 12 segments (exact number unknown). Anterior half of half ring smooth but posterior half sloping into furrow, moderately papillose. Up to 14 large perforated tubercles on each pleural strip. Posterolateral pleural corner well rounded. Facet more than half segmental length. Pygidium transverse with second transaxial furrow poorly impressed. Axis with relatively gentle posterior slope to flat border area. Pleural areas almost flat with one well and one poorly impressed pleural furrow separated by long flat rib bearing a poorly impressed interpleural furrow. Anterior strip of first segment lengthening markedly at and beyond fulcrum. Ornament of papillae everywhere except in furrows and close packed on high parts with large sparsely scattered tubercles over the same areas. Asthenopsis sp. cf. A. rhinostrongyla sp. nov. (Plates 38, figs. 3-7) Material Examined Two C with FC, IFC, and 3T including CPC17103 to 17105 and UQF68715 to 68716 from Ml 56 (located on Camooweal 1:250,000 Geological map) on the middle reaches of Opal Creek in the Age Creek Formation, Ptychagnostus punctuosus Zone, of the Currant Bush Limestone. Description Those features not consistent with A. rhinos- trongyla are referred to comparatively in this description. Glabella with better impressed anterior axial furrow producing a distinct break in lateral profile. Axial furrow poorly impressed throughout. Palpebral lobes narrower, markedly upturned laterally. Fixed cheeks almost as wide as the base of the glabella. Facial suture with o much closer to /i . Ornament, especially the papillae, more subdued, larger tubercles fewer in number. Free cheek with terrace lines prominent near margin. Border slightly narrower. Genal spine slightly more abaxially deflected. Pygidium unknown at present. Remarks The nature of the glabella, anterior border and palpebral lobes along with the slightly different ornament make reference to A. rhinostrongyla difficult. However, when a larger population of that Northern Territory species is studied the limits of variation may be such that this material will be assignable. It should be pointed out that the cephalic convexity, anterior border arch, subtle ornament, and in one instance (Plate 38, fig. 6) short frontal area also ally this species to A. opalensis. However, shape of the glabella and its furrows, eye ridges, shorter palpebral lobes, and straight anterior parts of facial suture distinguish that species. Asthenopsis unquinsepta* sp. nov. (Fig. 2D; Plate 35, figs. 3-8) Material Examined HolotypE; CPC17089, a cranidium from M157 on a sharp bend near the second creek junction on Opal Creek, latitude 19° 25' S, longitude 138° 35' E.; Age Creek Formation, Zone of Ptychagnostus punctuosus. Other Material: Five C, 2 FC, and 2P including CPC 1 7087, 17088, and 17090-2 (PL 35) from M157. Diagnosis Asthenopsid with glabella tapering forward to half its basal width, glabellar anterior truncated, anterolateral glabellar corners quite angular, furrow Ip with prominent domed area at its *From the latin numerals 1, 5, and 7, the material being from M157. 228 MEMOIRS OF THE QUEENSLAND MUSEUM midiength abaxially, frontal area relatively long and fixed cheeks of moderate width, anterior branch of facial suture straight and converging slightly forward, eye ridges doubled, ornament on border and glabellar of fine tubercles but on frontal area and fixed cheeks of coarser close spaced tubercles. Pygidium moderately vaulted. Description Glabella tapering anteriorly to half its basal width, sharply truncated anteriorly by change in ornament, anterior axial furrow not impressed even by a change of slope. Occipital ring less than four times as wide as long with medially placed node. Occipital furrow poorly impressed on axial half, with very steep anterior wall into pronounced apodemal pits laterally. Furrow Ip with domed abaxial elevation prominent. Glabellar ornament of fine tubercles. Axial furrow very poorly impressed throughout, with diverticulum from lobe Ip conspicuous. Ventral longitudinal muscle pits poorly impressed. Frontal area with prominent caecal network standing well above surface. Eye ridges doubled. Fixed cheeks almost flat. Palpebral lobe flat, only very slightly upturned and opposite furrow and lobe 2p. t - lc directed posteriorly at moderate angle to transverse line, crossing posterior border oblique- ly. Posterior limb relatively long and subtriangular in shape. Posterior border furrow increasing in length beyond fulcral point. Ornament on anterior border, on glabella except furrows of fine tubercles closely spaced, absent on palpebral lobes, posterior border furrow, and border furrow; on fixed cheeks, posterior limb, and frontal area ornament of close packed coarser perforated tubercles, with occasional ones larger than the rest. Free cheek with ornament on genal field similar to that of frontal area. Doublure extending adaxially, with terrace lines only on marginal curve, absent on ventral dorsally sloping part. Thorax with each articulating half ring having distinct transverse ridge near its midlength, anterior wall of articulating furrow rising very steeply to this ridge. Ornament of coarse tubercles on pleural strips not arranged in a single row. Pleural extremity squared, with rounded anterior corner and right angled to slightly pointed posterior corner. Pygidium with pleural areas moderately vaulted, anteriorly almost as high as axis that is sunken between pleural areas, axis well above pleural areas posteriorly. Axis lightly tapering to a well rounded posterior, almost flat transversely between apodemal pits, with apodemal pits becoming more adaxially placed posteriorly. Facets narrow and short. Ornament of coarse close packed tubercles on prominences and border. Morphogeny An individual with cranidium 6-5 mm long (Plate 35, fig. 6, lower) has 14 segments and can be interpreted as an holaspide. Another with cranidium 3-5 mm long (Plate 35, fig. 6, upper) has 1 1 segments of an incomplete thorax and may also be an holaspide. However, some mor- phological differences between these individuals and the mature ones (1 1 mm cranidia) are worth noting. At 3-5 mm ornament is fine with a few larger tubercles, the axial furrow is better impressed (possibly from compaction), anteriorly rounded glabella, fi— converges more strongly forward, and fixed cheeks arc narrower. At 6-5 mm the ornament is of moderately coarse close spaced tubercles, axial furrow is well impressed, fixed cheeks are narrower, and more convex, caecal ornament is subdued, /; - (S converges forward, and thoracic pleura have a line of up to seven large tubercles. Asthenopsis butorosa* sp. nov. (Fig. 2C; Plate 36, figs. 1-9) MaT[-RIA1. EXAMINFI) Hoi.gtvpE: ANU30649, a cranidium from ANU 10325, 5-6 km west of Chummy Bore on the Thorntonia to Camooweal Road; Currant Bush Limestone, Ptychagnostus punctuosus Zone. Other Material: One complete, 1 T, and 3 P including ANU30645-8 (PI. 36) from ANU10325; 6C, 3 FC, and 2 P including CPC 17093-6 (PI. 36) from Mi 61 at the base of the V Creek Limestone 12-8 km east of Morstone Homestead (marked on Camooweal 1:250,000 Geological Map), Ptychagnostus punctuosus Zone. Diagnosis Asthenopsid with highly vaulted cephalon, rounded anterolateral glabellar corners, poorly impressed anterior axial furrow represented by considerable change in slope, markedly laterally tapering anterior border, anterior branch of facial suture convex diverging forward, with very steeply sloping frontal area more than twice as long at facial suture as sagittally, with relatively close packed ornament of coarse tubercles on border. * Bu (Lat.) large, torus (Lat.) rounded protuberance, refers to the coarse tubercular ornament. JELL: MIDDLE CAMBRAIN TRILOBITA 229 frontal area, fixed cheeks, glabella, occipital ring, thoracic and pygidial axes and pleura. Description Cranidium (up to 16 mm long) with highly vaulted cheeks and glabella, with steeply sloping frontal area (especially anterolaterally) and posterior limb, and with glabella standing well above convex cheeks in anterior profile. Glabella tapering slightly forward to 0-75 basal width, with straight to very slightly waisted (at furrow Ip) lateral margins, with anterior axial furrow represented by a considerable change of slope (up to 40°) and hence more distinct than in any other species of the genus. Occipital furrow poorly impressed over axial half, apodemal area dumb-bell shaped extending up onto the rear of lobe Ip and occipital ring. Glabellar furrows smooth, Ip not expanded at adaxial end and without abaxial dome shaped elevation. Anteromedial muscle scar almost in anterior axial furrow. Anterior border most convex of any in genus, tapering strongly laterally. Border furrow well impressed with prominent axial and lateral anterior convexities. Frontal area moderately convex, almost vertically downsloping at facial suture, with depressions extending forward from axial to border furrows. Fixed cheek width almost equal to basal glabellar width, moderately convex. Facial suture with ji - (S abaxially convex and diverging forward. Or- nament on glabella (except furrows), frontal area, fixed cheeks (except palpebral lobe and furrow), posterior limb, and posterior border of coarse (up to 0-3 mm diameter) high tubercles, on anterior border and palpebral furrow of slightly finer tubercles, with tubercles on fixed cheeks becoming more widely separated towards the axial furrow. Free cheek moderately vaulted, with quite high eye socle. Genal spine with convex dorsal surface, and slight adaxial curve in distal portion. Ornament on genal field of close spaced tubercles superimposed on caecal ornament. Thorax of at least 14 segments (no complete mature thorax known). Axial lobe with coarse tuberculate ornament. Pleural furrow unforked adaxially. Pleural strips with single row of tubercles becoming less distinct on free pleurae. Free pleura with squared to slightly pointed posterolateral corner, with finer tuberculate ornament than elsewhere. Pygidium just over twice as wide as long. Pleural area with three or four pleural furrows and one interpleural furrow. Facets short, narrow, and directed strongly posteriorly. Fulcra prominently raised. Marked posterior excavation on first axial ring. Morphogeny Cranidium 3-5 mm long (Plate 36, fig. 1) much less vaulted, with coarse but sparse ornament, anterior part of facial suture converging forward. Cranidium 5*5 mm long (Plate 36, fig. 3) vaulted, with coarse, closer spaced ornament, anterior part of facial suture convex and diverging laterally, fixed cheeks higher than glabella. Asthenopsis opalensis* sp. nov. (Fig. 2B; Plate 37, figs. 1-4; Plate 38, figs. 1, 2) Material Examined Holotype: CPC17098, a cranidium from MNFI5, 1.6 km south of the junction of Opal Creek and the O’Shanassy River; Mailchange Limestone, Ptychagnostus punctuosus Zone. Other Material: Six C, 1 FC, and 1 P including CPC17097, and 17099-102 (Pis. 37 and 38) from MNF15. Diagnosis Asthenopsid with anterolaterally rounded glabella standing well above fixed cheeks, moderate cephalic convexity, four pairs of lateral glabellar furrows with smooth or punctate surfaces, distinctive occipital apodemal pits, anteriorly placed occipital node, eye ridges of three parallel trunks, straight strongly convergent anterior branches of facial suture, very fine close packed papillose ornament overlain by sparse larger tubercles. Pygidium transverse, weakly vaulted, with papillose ornament. Description Cranidium moderately convex, with glabella standing above convex fixed cheeks, a moderately steep frontal area, and a moderately arched anterior border. Glabella with rounded anter- olateral corners, convex anterior, with straight only slightly tapering lateral margins, with four pairs of lateral glabellar furrows and an anteromedial muscle scar. Occipital furrow shallow over axial half. Occipital apodemal pit distinctive, with wavy slightly raised smooth areas both anteriorly and posteriorly giving an hour glass shape with long axis transverse. Furrow 3p slightly arcuate, directed very slightly to the *Named for Opal Creek on which it is found. 230 MEMOIRS OF THE QUEENSLAND MUSEUM posterior adaxially and not connected to axial furrow. Furrow 4p short narrow, directed slightly anteriorly out of fossula very close to glabellar anterior. Glabellar furrows either smooth or finely punctate. Axial furrow crossed by low distinct diverticula at occipital ring and lobe Ip, 2p and 3p. Fossulae not at all distinct. Frontal area flat short and downsloping axially but convex laterally, with both slope and length (to twice sagittal length) increasing laterally. Eye ridges consisting of three parallel trunks (not visible in all specimens) with the median one highest and joining the anterior one just abaxial to the fossula. Parafrontal band low, arising out of fossula, not visible on all specimens. Palpebral lobe flat only slightly upturned laterally, anterior opposite anterior of lobe 2p, posterior level with anterior of lobe Ip. Palpebral furrow straight to slightly abaxially convex, deepending posteriorly. Fixed cheeks 0-9 of basal glabellar width, slightly convex and horizontal, with sparse tubercles over an extremely faint anastomosing caecal network radiating from the axial furrow posteriorly. Facial suture with I'i - ji straight, converging strongly anteriorly, t - concave adaxially then straight. Posterior border much longer and flatter beyond fulcrum. Free cheek moderately vaulted, with ji in border furrow. Border flat throughout. Genal field with sparse, coarse tubercles superimposed on faint caecal network. Pygidium transverse, with well rounded margin, and flat pleural areas. Two pleural furrows and an interpleural furrow present. Anterior pleural strip of first segment lengthening well inside fulcrum, weakly convex throughout. Marked posterior excavation in first axial ring. Asthenopsis sp. nov. (Plate 32, fig. 6) Material Examined One complete, but damaged, exoskeleton (CPC 17069) from 12 km south of Douglas Spring on the Morstone to Undilla Road. It occurs in the V Creek Limestone in the Ptychagnostus nathorsti Zone in association with Asthenopsis levior and Papyriaspis lanceola. Description This specimen, being somewhat damaged anteriorly and distorted by several cracks, cannot be satisfactorily compared with other species on cranidial features. Only those features that are at variance with the description of A. levior above, are referred to here. It has a flatter border on the free cheek. The genal spine is not deflected. The pygidium has only a weakly convex pleural area and a poorly defined border. The thorax has 16 segments, and a tuberculate pleural ornament. The segments are individually shorter than their corresponding number in A. levior whether matching them (i.e. 14 against 14) from the anterior or posterior. Remarks Although this specimen could be included in A. levior as representing a population well away from the topotype population 1 believe that the undcficcted genal spine alone, is sufficient to separate it at the species level. In the absence of other material I am reluctant to erect a new name but am confident further collecting will necessitate the assignment of one. Distinguishing features of the five new northern Australian species described herein are outlined in Table 1. LITERATURE CITED Campbell. K. S. W., (in press). Trilobiles of the Haragan, Bois d'Arc and Frisco Formations. Bull. Okla geol. Surv. CiSNE. J. L., 1974 Trilobites and the origin of the arthropods. Science 186: 13-18. Hfssler, R. R., 1964. The Cephalocarida. Comparative skeletomusculature. Mem. Conn. Acad. Arts Sci. 16: 1-97. Jell, P. A., 1975, Australian Middle Cambrian codiscoids with a review of the superfamily. Palaontographica A 150: 1-97. Kobayashi. T., 1935. The Cambro-Ordovician forma- tions and faunas of south Chosen. Palaeontology. Part 3. Cambrian faunas of south Chosen with a special study on Cambrian trilobite genera and families. J. Fac. Sci. Tokyo Univ. (sect. 2) 4: 49-344. Moore, R. C., Editor. 1959. ‘Treatise on Invertebrate Paleontology’. Part O, Arthropoda 1 (Geological Society of American and University of Kansas: Boulder and Lawrence). Opik, a. A., 1967. The Mindyallan fauna of north-western Queensland. Bull. Bur. Miner. Resour. Geol. Geophys. Aust. 74: 1-404. Richter, R. and Richter, E., 1940. Die Sankianda — Stufe von Andalusien, eine fremde Fauna im europaischen Ober-Kambrium. Abh. Senkenberg Naturf. Ges. 450: 1-88. Wai ( on . C. D., 191 1. The Cambrian faunas of China. Smithson misc. Colins 57: 69-109. Whitehouse, F. W., 1939. The Cambrian faunas of north-eastern Australia. Mem. Qd Mus 11* 179-282. TABLE 1: Distinguishing Features of Species of Asthenupsis JELL: MIDDLE CAMBRAIN TRILOBITA 73 is (X "O M u 73 '5 J3 — OiD o CQ s ffi 231 MEMOIRS OF THE QUEENSLAND MUSEUM Platf 31 Asthenopsis levior Whitehouse, 1939 (1, 4, 6, 7, 8 from the type locality on V Creek) Fig. 1: Thorax and pygidium, UQF3340, (figured by Whitehouse, 1939, pi. 22, fig. 20), x 1-2. Fig. 2: Latex cast of damaged early holaspid exoskeleton, CPC 17064, X 6, from M4L Fig. 3: Early holaspid cranidium, UQF68711, x 10, from UQL3510. Fig 4: Pygidium, UQF3339, (figured by Whitehouse, 1939, pi. 22, fig. 19), X 2. Fig. 5; Damaged cranidium, UQF68712, x 4-5; b, anterior oblique view, from UQL3510. Fig 6: Damaged cranidium, UQF3338. (figured by Whitehouse, 1939, pi. 22, fig. 18), X 2. Fig. 7: Latex cast of slightly damaged Holotype exoskeleton, UQF3337, (figured by Whitehouse, 1939, pi. 22, fig. 17), x 2-5; a, lateral oblique view, b, dorsal view. Fig. 8: Large slightly damaged complete exoskeleton, UQF44314, x 1-5. Fig. 9: Ventral view of left free check, CPC17065, x 3 from M139. Illustrations are dorsal views unless otherwise stated. JELL: MIDDLE CAMBRAIN TRILOBITA Plate 31 MEMOIRS OF THE QUEENSLAND MUSEUM Pi ATI- 32 Asthenopsis levior Whitehouse, 1939 Fig. 1: Damaged cranidium, CPC 17066, x 2-5, from M247. Fig. 2: Cranidium, UQF68713, x 1-5, from UQL469. Fig. 3: Damaged pygidium exhibiting well developed caeca passing into the border, UQF68714, a, x 2-5, b, x 7, from UQL469. Fig. 4; Cranidium, CPC 1 7067, x 3, from Douglas Creek halfway from the Old Burketown Road to the O'Shanassy River. Fig. 5: Cranidium, CPC 17068, x 3, same locality as Fig. 4. Asthenopsis sp. nov. Fig. 6; Damaged exoskeleton, CPC 1 7069, x 2-2, from 12 km south of Douglas Creek on the Morstone to Undilla Road. JELL: MIDDLE CAMBRAIN TRILOBITA Plate 32 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 33 Asthenopsis queenslandica sp. nov. (All material is from M243) Fig. 1: Cranidiiim, CPC17070, a, x 4. b, x 6-5. Fig 2; Small cranidium, CPCI7071, x 9. Fig. 3: Right free cheek, CPC17072, x 3. Fig. 4: Small cranidium, CPC17073. x 7. Fig. 5: Holotype exoskeleton, CPC 17074, x 3. Fig. 6: Large damaged exoskeleton without free cheeks, CPC 17075, X 2-5. Fig. 7: Large damaged exoskeleton without free cheeks, CPC 17076, X 5. Fig. 8: Pygidium, CPC17077, x 6-5. JELL; MIDDLE CAMBRAIN TRILOBITA Plate 33 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 34 Asihenopsis queenslandica sp. nov. Fig. 1: Cranidium, CPC17078, x 4-5, a, anterior view, from M243. Asthenopsis rhinostrongyla sp. nov. (All material is from HI 38) Fici. 2: Small cranidium, CPC17079, x 5-5. Fig. 3: Hololype cranidium, CPC 1 7080, x 3-5, a, anterior view. Fig. 4: Ventral view of right free check, CPC19081, x 4-5. Fig. 5: Thoracic fragment, CPC17082, x 2-5, b, lateral view. Fig. 6: Latex cast of cranidium, CPC17083, a, x 4-2, b, lateral oblique view, X 4. Fig 7: Left free cheek, CPC17084, x 4. JELL: MIDDLE CAMBRAIN TRILOBITA Plate 34 MEMOIRS OF THE QUEENSLAND MUSEUM Pl.ATH 35 Asthenopsis rhinostrongyla sp. nov. Fi(i, 1; Left free cheek, CPC 17085, x 5, b, lateral view, from HI 38. FKi, 2; Pygidium, CPC17086, x 6, from H138. Asihenopsis unquinsepta sp. nov. (All material is from Ml 57) Fig. 3: Pygidium, CPC17087, x 3. Fki. 4: Left free check, CPC 17088, x 4. Fig. 5: Holotype cranidium, CPC 17089. x 3, a, lateral view. Fig. 6: Two small individuals slightly damaged, CPC 17090, x 3. Fig. 7: Latex cast of left free cheek, CPC 17091, x 4. Fig. 8: Cranidium and thorax, CPC17092, x 2-5. JELL: MIDDLE CAMBRAIN TRILOBITA Plate 35 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 36 Asthenopsis butorosa sp. nov. Fig. 1: Latex cast of complete (less free cheeks) early holaspide, ANU30645, x 5. from ANUI0325. Fig 2: Pygidium, CPC17093, x 3-5, from M161. Fig, 3: Small cranidium, CPC17094, x 7, b, lateral oblique view, from from Ml 61. Fig. 4: Latex cast of pygidium and damaged thorax, ANU30646, x 3, from ANU 10325. Fig. 5; Cranidium, CPC 17095, a, lateral oblique view, x 4, b, dorsal view, X 4, c, anterior oblique view, x 5, from M161. Fig. 6; Right free cheek, ANU30647, x 5, from ANU10325. Fig. 7: Latex cast of pygidium, ANU30648, x 5, from ANU10325. Fig. 8; Holotype cranidium, ANU30649, x 2-8, from ANU10325. Fig. 9: Cranidium, CPC17096, x 3, from M161. JELL: MIDDLE CAMBRAIN TRILOBITA Plate 36 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 37 Asthenopsis opalensis sp. nov. (All material is from MNF15) Fig. 1: Cranidium. CPC17097, x 3. Fig, 2: Holotype cranidium, CPC17098, a, anterior view, x 3, b, 4. Fig. 3: Cranidium, CPC17099, x 4, a, anterior oblique view. Fig. 4: Cranidium, CPC17100, a, x 5-5, b, x 6, c, x 2*5. JELL: MIDDLE CAMBRAIN TRILOBITA Plate 37 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 38 Asthenopsis opalensis sp. nov. Fig. 1: Cranidium. CPCI7101, x 3, from MNF15. Fig. 2; Pygidium, CPC17102, x 4-5, from MNF15. Asthenopsis sp. cf. A. rhinostrongyla sp. nov. Fig, 3: Cranidium (mostly exfoliated), CPC17103, x 3, a, lateral view, b, from MI 56. Fig. 4: Right free cheek, CPC17104, x 4-5, from M156. Fig. 5: Pygidium, UQF68715, x 5, from UQL447. Fig. 6: Cranidium, CPC17105, x 3, a, anterior view. Fig. 7; Cranidium, UQF68716, x 4-5, b, anterior oblique view. JELL: MIDDLE CAMBRAIN TRILOBITA Plate 38 Mem. QdMus. 18(2): 233-43, pi. 39. [1978] PARANCHISTUS PYCNODONTAE SP. NOV., A NEW PONTONIINE SHRIMP ASSOCIATED WITH AN OSTREID BIVALVE HOST A. J. Bruce Heron Island Research Station, Gladstone ABSTRACT Paranchistus pycnodontae (Crustacea, Decapoda, Pontoniinae), a new species of shrimp from Heron Island, Queensland, is described and illustrated. The species is most closely related to P. spondylis Suzuki, from which it is distinguished. It is the second species of this genus to be found in Australian waters and the first to be found in association with an ostreid host, Pycnodonta hyotis L. The genus Paranchistus was first designated by Holthuis (1952), and at present includes four species, all of which are now known to be associates of large marine bivalve molluscs. Of these four species, only one, Paranchistus armatus (H. Milne-Edwards), has so far been recorded from Australian waters. This species is an associate of the giant clam, Tridacna gigas (L.), and has been reported from Undine Reef, Cape Tribulation, and from Chapman Island, Queensland (McNeill 1968, Bruce 1975). Recently a single example of another species of this genus was found in association with an ostreid bivalve, and the specimen is now described here as new species. Paranchistus pycnodontae sp. nov. (Figs. 1-5, Plate 39) Material Examined HolotypE; Queensland Museum W7337, 5 , non- ovigerous; Heron Island, Capricorn Group, Queensland; No. 2442, 3 m, central lagoon, 17 August 1976, coll. D. Fisk. Description A medium sized pontoniine of moderately slender build. Carapace smooth. Rostrum well developed, strongly compressed, with feebly developed carina and extending anteriorly to middle of intermediate segment of antennular peduncle; rostrum horizontal, dorsal and ventral margins straight and subparallel, tapering to an acute tip distally; distal third of upper border with five small subequal, acute teeth, with numerous setae in interspaces; ventral border sparsely setose with a single very small acute tooth below third dorsal tooth. Orbit feebly developed, supraorbital spines absent; inferior orbital angle slightly produced, broad in dorsal view. Antennal spine small, slender, marginal, not exceeding inferior orbital angle. Hepatic spine distinctly projecting, more slender than antennal spine and lower and more posterior, in a small fossa and mobile. Anterolateral angle of carapace bluntly rectangular. Abdominal segments smooth. Third segment not produced posteriorly in dorsal midline. Length of sixth segment 013 x depth, 015 x length of fifth segment. Pleura 1-3 only slightly expanded, 4-5 smaller, bluntly rounded posteriorly, 6 with posteroventral angle blunt, posterolateral angle slightly more acute. Telson c. 1-5 x length of sixth abdominal segment, 2-2 x longer than broad; lateral margins almost straight, converging posteriorly; anterior width c. 2-7 X width at level of lateral pair of posterior spines. Two pairs of small, subequal, submarginal dorsal spines at 0-6 and 0-7 of telson length. Posterior telson margin rounded, without median point, with three pairs of spines; lateral pair small, similar to dorsal spines, situated on dorsal surface of telson, slightly in advance of posterior margin; intermediate pair very stout, c. 0-14 X telson length, with attenuated distal ends, submedian pair uniformly tapering, setulose, c. 0-75 X intermediate spine length. 234 MEMOIRS OF THE QUEENSLAND MUSEUM Eyes normal, cornea globular, hemispherical, slightly oblique, with distinct accessory spot; podophthalmite slightly flattened, c. 1-2 x longer than broad, distinctly wider than diameter of cornea. Antennules normal; peduncle exceeds rostrum by most of intermediate segment; proximal segment c. 1-3 x longer than proximal width, medial border straight, without ventral spine; styiocerite slender, acute, exceeding half segment length; lateral border feebly convex, medially convergent, with very short subrectangular distolateral spine; anterolateral margin strongly produced, distinctly angulated; intermediate and distal segments subequal, obliquely articulated, c. 0-7 X length of proximal segment. Upper flagellum short, biramous, proximal three segments fused; shorter free ramus with two stout segments, longer with nine slender segments; four groups of aesthetascs. Lower flagellum slender, sixteen segments. Antenna with robust basicerite, with feeble distolateral tooth; ischiocerite, merocerite normal; carpocerite slender, exceeding middle of scaphocerite, c. 3-2 x longer than wide. Sca- phocerite well developed, extending well beyond antennular peduncle: lateral border very feebly convex, with acute distolateral spine, not exceeding anterior margin of lamina. Lamina broad, c. 2-2 x longer than width, anterior margin broad, slightly truncate. Flagellum short, slender, c 2 x postorbital carapace length. Mouthparts generally similar to other species of Paranchistus. Mandibles moderately robust, without palp; molar process stout, with several large blunt distal teeth, incisor process slender, three acute teeth distally, central tooth smallest. Maxillula normal; palp with feebly developed lateral lobe, medial lobe normal with small hooked terminal seta; upper and lower lacinia broad, densely setose with strongly setulose seta; short, stout, feebly dentate spines distally on upper lacinia, spiniform setae on lower. Maxilla with stout, proximally swollen palp, medial and lateral borders setose, distally slender, non-setose. Basal endite well developed, broad, deeply cleft, each lobe with numerous long, slender, setulose setae. Scaphognathite broad, c. 2 6 X longer than width; anterior lobe broad. Fig. 1: Paranchistus pycnodontae sp. nov., holotype female, Heron Island, Queensland. Scale in mm. BRUCE: A NEW PONTONIINE SHRIMP 235 Fig 2’ PcirQfichistus pycnodontQ^ sp. nov., holotype: A, anterior carapace, rostrum, and antennae, dorsal aspect, B, anterior carapace and rostrum, lateral; C, orbital region, dorsal; D, distal rostrum; E, inferior orbital angle and antennal spine; F, eye, dorsal; G, antennular peduncle; I, antenna; J, distolateral tooth of scaphocerite; K, telson; L, posterior telson spines; M, uropod; N, distolateral spine of exopod of uropod, 236 MEMOIRS OF THE QUEENSLAND MUSEUM Fig. 3: Paranchistus pycnodontae sp. nov., holotype: A, mandible; B, maxillula; C, maxilla; D, first maxilliped- h, second maxilliped; F, third maxilliped; ^ BRUCE: A NEW PONTONIINE SHRIMP 237 20mm C 0-5m m BF 0-25 mm HI Fig. 4: Paranchistus pycnodontae sp. nov., holotype: A, first pereiopod; B, chela of first pereiopod; C, second pereiopod; D, chela of second pereiopod; E, fingers of chela; F, fixed finger of chela; G, third pereiopod; H, distal propod and dactyl of third pereiopod, lateral; I, idem, dorsal aspect of dactylus; J, accessory dactylar tooth of fourth pereiopod; 238 MEMOIRS OF THE QUEENSLAND MUSEUM First maxilliped with subcylindrical palp, distomcdially with setulose setae, exceeding anterior margin of endite but not caridean lobe; basal endite broad, rounded, medial border straight, confluent with coxal endite, junction indicated by minute protuberance, with numerous slender finely setulose setae; dorsal surface with oblique row of long, coarsely setulose setae; coxal endite sparsely setose. Exopod well developed, flagellum with four plumose distal setae only, caridean lobe small, broad. Epipod triangular, deeply bilobed. Second maxilliped normal; dactylar segment c. 3 X longer than broad, numerous slender, finely setulose setae medially. Exopod with four plumose setae distally; epipod small, irregularly subrectangular; podobranch absent. Third maxilliped moderately slender, reaching proximal end of carpocerite; ischiomerus feebly separated from basis, junction indicated medially by small notch: ischiomerus slightly tapering distally, moderately bowed, r. 4 x longer than proximal width, medial border feebly setose, short simple setae; penultimate segment c. 0-6 x ischiomerus, 3-2 x longer than width, with longer stouter setae medially and laterally; distal segment similarly setose, c. 0-5 x ischiomerus, tapering, c. 3-5 X width. Basis broad, sparsely setose medially, as long as wide: coxa stout, medially rounded, well developed epipod laterally. Exopod slightly exceeds ischiomerus, four plumose distal setae. Arthrobranch rudimentary. Coxae of third maxillipeds widely separated; fourth thoracic slernite broad, unarmed. First pereiopods slender proximally, stouter distally, exceeding carpocerite by carpus and chela; chela robust, palm subcylindrical, slightly compressed; fingers subequal to palm, broadly subspatulate, distally rounded with fine denticula- tions along distal third of cutting edge medially and two thirds laterally, slightly gaping proximally on medial side, densely setose. Palm r. 1-7 X longer than width, transverse rows of cleaning setae proximally. Carpus proximally slender, width increasing x 3 distally, r. 5 x longer than distal width, cleaning setae dis- tomedialiy; mcrus subequal to carpus, slightly bowed, c. 6-4 x longer than width; ischium compressed, c. 2-6 x width, c. 6-5 x merus, medial border with row of short setae; basis 0-33 x merus, sparsely setose along medial margin, coxa robust, small setose ventromedian process present. Second pereiopods well developed, slender, subequal, similar. Palm of chela subcylindrical, smooth, slightly compressed distally, c. 3-4 x width; fingers well developed, c. 0-6 of palm length, dactylus strongly curved, overreaching fixed finger, tip acutely hooked, cutting edge sharp, entire except for small tooth at 0-3 of length, outer surface with numerous short, erect setae: fixed finger similar, proximal half of cutting edge with eight small teeth, distal four subacute, proximal four rounded, distal part of cutting edge feebly denticulate. Carpus c. 0-47 x palm length, 2 X longer than distal width; distal margins feebly excavate, unarmed. Merus c. 0*75 x palm length, 4 X longer than width, uniform, with small distovenlral tooth. Basis and coxa normal. Ambulatory pereiopods slender, third extending beyond scaphocerite by 0-3 of carpus and chela; dactyl about 0-27 x propod length; unguis distinct from corpus, dorsoventrally compressed, hastate, surface covered with dense felt of short processes giving furry appearance; corpus w'ith single well developed, slender acute accessory spine; short distal sensory setae present. Propod c. 6-6 x longer than width, uniform, with pair of distoventral spines and single preterminal ventral spine. Carpus 0-63 propod length, c. 3-2 x longer than distal width, unarmed; merus c. 1-27 x propod length; ischium c. 1-0 x propod length, c. 5 x longer than distal width; basis and coxa robust, all unarmed. Pereiopods 4-5 similar: 3, 5 subequal, 4 slightly shorter. Accessory spines on dactyls of pereiopods 4. 5, with small additional denticle on posterior margin. Uropods slightly exceed telson tip, protopodite with large acute posterolateral tooth; exopod broad, c\ 2-3 x longer than wide, broadly rounded distally, with small mobile spine only at posterior end of convex lateral border; endopod c. 3 x longer than width. MEASUREMENTS: (In millemetres) Total length (approx.) 18.7 Rostrum and carapace 6.2 Postorbital carapace 4.5 Second pereiopod chelae 3.8 (left and right) COLOURATION: Semi-lransparenl with numer- ous small round evenly distributed red chromato- phores, all over body, ambulatory pereiopods, antennae and caudal fan. Similar but slightly larger chromatophores over second pereiopods. Chromatophores absent from antennal flagella and propods of ambulatory pereiopods. HOST; Pycnodonta hyotis L. (Ostreidea) ASSOCIATES: One speeimen of Onuxoden parvibrachium (Fowler) (Pisces: Carapidae), a species not previously known from Australia. BRUCE: A NEW PONTONIINE SHRm? 239 TABLE 1: Hosts oi Si‘i(iis oi P\r\\(Mishs wn omiir Pomomim. Assoc iahs Paranchistus spp. Hosts Other associates P. armatus (H. Milne- Edwards) Tridacna gigas (L.) Anchistus niiersi (De Man) Conchodytes tridacnae Peters P. ornatus Holthuis Atrina vexillum Born Anchistus custos (Forskal) Conchodytes hiunguiculatus (Paulson) P. nohilii Holthuis Spondylus gaederopus L. — P. spondylis Suzuki Spondylus harbatus Reeve — P. pycnodontae sp.nov. Pycnodonta hyotis L. Platypontonia hyotis Hipeau-Jacquotte Kig. 5: Paranchistus pycnodontae sp. nov., holotype, colour pattern. Systematic Position of P. pycnodontae Paranchistus pycnodontae appears to be most closely related to P. spondylis Suzuki, and may be distinguished from that species by the sub-spatulate fingers of the first pereiopods. In all other features the morphology of the two species appears to be very similar. Other minor differences include: In P. spondylis (a) the palm of the chela of the second pereiopods appears to be stouter; (b) the accessory spines of the ambulatory pereiopods are shorter and stouter; (c) the accessory spines of the dactyls of the fourth and fifth pereiopods are without denticles; (d) the minute processes on the anterior aspect of the dactyls of the walking legs are perpendicular to the surface; (e) the lateral posterior telson spines appear to be marginal; (f) and the intermc^diate posterior telson spines arc of uniform taper. DISCUSSION All species of the genus Paranchistus Holthuis are known to be associates of bivalve molluscs. The genus is not represented outside tropical Indo-West Pacific waters. The host molluscs belong to the families Tridacnidae, Pinnidae, Spondylidae and now also include the Oslrcidae. P. pycnodontae is the only shrimp so far known to associate with Pycnodonta hyotis in Australian waters, but in the western Indian Ocean Hipcau-Jacquotte (1971) has recorded the occurrence of a different pontoniine shrimp, Platypontonia hyotis Hipcau-Jacquotte, which is also known to associate with Pterostrea imhricata (Lam.) in Japanese waters (Suzuki 1971). The only other ostreid bivalve known to have a pontoniine associate is Lopha cristagalli (L.), the host of Platypontonia hrevirostris (Miers), (Bruce 1968). The hosts of the species of Paranchistus are summarized in Table 1, which also indicates other pontoniine associates of the same host. 240 MEMOIRS OF THE QUEENSLAND MUSEUM In addition to the morphological characters that distinguish P. pycnodontae from P. spondylis, it may be noted that the colour patterns also appear to be distinctive. In the figures of the latter given by Suzuki (1971), the specimen shows a comparatively coarse pattern of larger and more sparsely distributed red chromatophores than is found in P. pycnodontae. Most of the pontoniine associates of the bivalve molluscs present a colour pattern of uniformly distributed chromatophores over most of the body surface and appendages. These may be very small and numerous, as in Anchistus custos, Paranchistiis ornatus or Conchodytes tridacnae. In P. spondylis and P. pycnodontae they are larger and more numerous, and in Anchistus demani and A. miersi, they are conspicuously larger and much less numerous. In complete contrast, the colour pattern of Platypontonia hyotis consists mainly of large spots and bands of red. The dactyls of the ambulatory pereiopods are of particular interest as, in the genus Paranchistus. they exhibit a wide range of morphological variations of great value in distinguishing the species. P. arniatus presents a simple condition, with a simple, well developed unguis distally and a broad accessory tooth. There is no trace of ornamentation on the unguis (Bruce 1975). In P. ornatus the unguis is still distinct from the corpus and the outer surface is covered with transverse rows of short spines which become obsolescent towards the tip. The ventral aspect of the unguis is also transversely ridged. The corpus has distal sensory setae present, but the accessory spine is absent, the region of the spine being provided with a low eminence covered with small tubercles. The pit described by Holthuis (1952) appears to be a vesicle below this eminence. In P. pycnodontae, the arrangement is essentially similar except that a well developed accessory spine is present and the unguis is very much larger. A comparable vesicular structure and sensory setae are also present. P. nohilii and P. spondylis have dactyls very similar to P. pycnodontae. Fujino (1975) has provided scanning electron photomicrographs of the dactyl of the walking legs in Anchistus miersi (De Man). Anchistus Borradaile is a genus very closely related to Paranchistus Holthuis, from which it is separated principally by the complete absence of an hepatic spine. In Anchistus a comparable range of variation in dactylar morphology is also present and in species such as A. demani Kemp and A. miersi, with A. gravieri Kemp representing a less well developed stage, and the dactyls very closely resemble those found in some Paranchistus spp. SEM photomicrographs clearly show that the outer surface of the unguis of the dactyls is densely covered with large numbers of short erect spinules. In Anchistus the anterior aspect of the unguis has been reported as scoop-shaped (Kemp 1922; Fujino 1975) but in Paranchistus spp. this surface appears to be convex rather than concave. The precise function of the spinulalions of the unguis is still obscure, but is presumably related to the grip of the shrimp upon its host. The function of the vcntrally situated tooth on the corpus would appear to enable a good grip upon the host's tissues to be obtained during traction by the limb. The distally directed spinules of the unguis may function by preventing the penetration of the dactyl too far into the host's tissues and thereby avoiding excessive damage to the host in a commensal relationship. ACKNOWLEDGMENT I am grateful to Dr Paxton for the identification of the associated fish. LITERATURE CITED Bruci;. a. J., 1968. Notes on some Indo-Pacific Pontoniinae. XII. The re-cxamination of the types of Pontonia brevirostris Miers, 1844, with the designation of a new genus Platypontonia (Decapoda, Natantia). Cru.staceana 15(3); 289-97, figs. 1-3. 1975. Pontonia armata H. Milne-Edwards (Decapoda Natantia, Pontoniinae), — a correction. Crustaceana 29(1); 49-54, figs. 1-3. Fi jino, T., 1975. Fine features of the dactylus of the ambulatory pereiopods in a bivalve associated shrimp, Anchistus miersi (De Man), under the scanning electron microscope (Decapoda, Natantia. Pontoniinae). Crustaceana, 29(3); 252-54, figs. 1, pi. 1. Hiprau-Jacqliottk. R., 1971- Notes de faunistique et de biologic marines de Madagascar. V. Platypontonia hyotis nov. sp., (Decapoda Natantia, Pontoniinae). Crustaceana. 20(2): 125-40, figs. 1-7. Holthlis, L. B., 1952. The Decapoda of the Siboga Expedition. Part XL The Palaemonidae collected by the Siboga and Snellius Expeditions with remarks on other species. IF Subfamily Pontoniinae. Siboga Exped. Mon., 39a***: 1-252, figs. I-IIO, tab. 1. Kfmp. S., 1922. Notes on Crustacea Decapoda in the Indian Museum. XV. Pontoniinae. Rec. Indian Mus. 24: 113-288, figs. I-I05, pis. 3-9. Mac Neill. F. A., 1968. Crustacea, Decapoda and Stomatopoda. Great Barrier Reef Expedition. 1928-29. Sci. Rep. 1(1): 1-98, figs. 1-2, pis. I -2. Suzuki, H., 1971. On some commensal shrimps in the western region of Sagami Bay. Researches Crust. 4: 1-25, figs. 1-12, pis. 1-3. MEMOIRS OF THE QUEENSLAND MUSEUM Plate 39 Paranchistus pycnodontae sp. nov., holotype, in Pycnodonta hyotis L. BRUCE: A NEW PONTONIINE SHRIMP Plate 39 Mem. Qd Mus. 18(2): 243-63. [1978] GASTROINTESTINAL NEMATODES FROM AQUATIC AUSTRALIAN SNAKES H. 1 . JONES Department of Parasitology, University of Queensland ABSTRACT Three new nematodes are described, and another three species are recorded from six species of aquatic snake (Acrochordidae and Colubridae). Four nematodes are from genera which predominantly parasitise vertebrates of other Phyla or Orders, but which inhabit a similar aquatic environment to these snakes. This paper is the first in a proposed series on the gastrointestinal parasites of Australian snakes. Little work has been done on the nematode parasites of the Australian tropica! and sub- tropical aquatic snakes, most of it being confined to identifications and host records (Johnston and Mawson 1941. 1948), with no indication of intensity or prevalence of infection. This paper is concerned with the gastrointestinal nematodes from 26 Australian aquatic Colubrid and Acrochordid snakes preserved in the Queensland Museum, Brisbane. Representatives from all six Australian species in this group were examined, namely Acrochordids javanicus, Acrochordus granulatus, Cerberus rhynchops, Myron ri- chardsoni. Fordonia leucobalia and Enhydris polylepis. The nematodes recovered are as follows: Subclass ADENOPHOREA Order ENOPLIDA Superfamily DIOCTOPHYM ATOIDEA Eustrongylides acrochordi sp. nov. (Fig. I, Table 1) Material Examined HolotypE: Queensland Museum G 1 0275, ^ , collected by H. Jones from stomach of Acrochordus javanicus. QM J2319I, collected by J. Covacevich and C. Tanner near Coen, North Queensland, June 1973. Paratype: QM G 1 0276, ^ , (incomplete), same data as holotype. Diagnosis Double row of six apical papillae, marked terminal striations and row of lateral punctations, nerve ring anterior, oesophagus long and voluminous, anus terminal. Vagina opens into the rectum, forming a cloaca. No tail. Description Worm long, cylindrical and partly coiled. Very marked striations near both ends, diminishing until scarcely visible near centre of worm. Two lateral rows of very small punctate markings, situated in the striations, proceed posteriorly from anterior end, disappearing within a few mm. Two lateral rows of four evenly spaced small papillae at caudal end, replaced anteriorly by punctate markings as at anterior end, gradually disappear- ing. Body of fairly uniform thickness, tapering slightly at posterior end but more markedly at anterior end. Mouth elongated dorsovcnlrally, surrounded by six spined papillae, two laterally and four submedially. Another row of six rounded papillae below these. Another 12 much smaller papillae, two just bclow' each lateral spined papilla, one just anterior to each lateral rounded papilla, and the remaining six at intervals between the two main rows of papillae. A short straight pharynx leads into a wide undulating oesophagus; this increases slightly in width throughout its length, is without a bulb, and measures 0-2 x the total length of the worm. Nerve ring surrounds oesophagus just behind its origin. No excretory pore seen. A wide rectum, flattened 244 MEMOIRS OF THE QUEENSLAND MUSEUM Fi(i 1. hustrongylides acrochordi sp. nov. Holotype ^ G10275, A, anterior end; B, anterior extremity, en face; C, Anterior end and oesophagus; D, posterior end, dorsal; E, posterior end, lateral, showing insertion of vagina into rectum. JONES: NEMATODES FROM AQUATIC SNAKES 245 TABLE 1: Body Measurements (In mm) of Holotype Immature Female Eustrongylides acrochordi sp. nov. Total length 84-50 Width at mid-point 103 Width at 2-0 mm from front 0*56 Head width at 1st row of papillae 015 Head width at 2nd row of papillae 0-22 Width at beginning of oesophagus 0-28 Oesophagus length 15-90 Oesophagus proportion of length 1/5 Oesophagus width near beginning 01 8 Oesophagus width c. 1 /4 along 0-28 Oesophagus width c. 3/4 along 0*42 Pharynx length 0-19 Length of anterior papilla spines 0*01 Nerve ring fr. ant. extremity 0*24 Nerve ring fr. oesoph. beginning 0 03 Caudal papillae distance 0 08 Punclation spacing at anterior end 0-04 Punctation spacing proceeding posteriad 0*13 Anus width 010 Rectum length 0-38 Rectum width 0*13 Post, intestine-rectum 1 00 Width of uterus 4 mm fr. tail 0-15 Width of uterus near rectum 0 06 Lumen of uterus 0*04 Uterine wall thickness 0*01 Distance of post, edge of vagina fr. anus 0*08 dorsoventrally, leads into the terminal anus. This is widened laterally and is surrounded dorsally and venlrally by a rugose ridge. There is no tail. Ovary not seen. The vagina passes posteriorly and just before the rounded posterior end of the worm turns sharply medially and enters the rectum. The uterus was not fully developed and contained no eggs. Discussion Both the holotype and the paratype (which lacked the anterior end) were lying freely in the stomach among a large number of Tanqua ophidis. The worms are characteristic of the genus, but differ from all described species in that the vagina opens into the rectum, forming a cloaca. This feature is sufficient to ascribe them to a new species, even though neither worm is mature. In all other Eustrongylides species the vulva opens terminally, very close to the anus. A cloaca has been described from only one other group of adenophorean nematodes, the free-living Lauratonema species (Gerlach 1953). All previous records of Eustrongylides species have been from large aquatic birds, from the proventriculus or a.ssocialed glands, and it is possible that this snake was acting as a paratenic host. Subclass SECERNENTEA Order OXYURIDA Superfamily OXYUROIDEA Spironoura fordoniae sp. nov. (Fig. 2, Table 2) Material Hxaminfi^ Holotype Queensland Museum G 10277, collected by H. Jones from rectum of Fordonia leucobalia, QM J23200, collected by J. BredI from Edward River, North Queensland, 1973. ALLOTYPES; QM G 10278, 2 ^ (poor condition), same data as holotype. ParatypeS: QM G10279, 3 ^ ^,1 55 , same data as holotype; QM G 1 0280, 8 2 55 , (poor condition), collected by H. Jones from stomach of F. leucobalia. QM J23924, collected by J. Bredl from Edward River, North Queensland, 1973. Diagnosis Head with three lips surrounded by six papillae. Vestibule present. Pharynx short, oesophagus with hour-glass formation at posterior end, stout equal alatc spicules, conspicuous oblique precaudal ventral muscle bands, ten pairs of caudal papillae, one unpaired precloacal papilla, lateral alae from cervical region to near posterior end, vulva just past mid-length. Description Worm tapering towards either end. No constriction behind head. Body finely striated. Lateral alac commence in cervical region and run most of the length of body, diminishing in size and finally disappearing in last quarter of body. Tail slender and finely pointed in both sexes. Mouth with three lips, with two papillae at base of each. Short vestibule leading into muscular pharynx, length about 2-5 x width. Oesophagus muscular, long, gradually increasing in diameter to a prcbulbar swelling (preceded by a constriction) and a large spherical muscular bulb. Intestine straight and wide lumcned. Nerve ring 0-25-0-20 from front of oesophagus. Small but distinct cervical papillae about 0-66 along oesophagus, excretory pore about 0-75 along oesophagus. MALE: Length rather less than females. Caudal end coiled venlrally. Oblique precaudal ventral muscle bands conspicuous; no ventral sucking disc. No caudal alae. Ten pairs of sessile caudal papillae, arranged as three regularly spaced ventrolateral precloacal papillae, three closely set pairs of paracloacal papillae, and four pairs of caudal papillae of which the most anterior two are 246 MEMOIRS OF THE QUEENSLAND MUSEUM Eld 2. Spironoura fordoniae sp. nov. Holotype 3 G10277. A, anterior end; B, anterior extremity, en face; C, anterior end and oesophagus. D, posterior end, ventral; E, posterior end, lateral; F-I, sections at level of upper oesophagus, lower oesophagus, mid-body and near posterior end, respectively. TABLE 2; BOD't Mlasi revients {IN mm) or Spironoura fordoniae. sp nov and Camallanides cerberi sp JONES: NEMATODES FROM AQUATIC SNAKES 247 248 MEMOIRS OF THE QUEENSLAND MUSEUM lateral and the most posterior two ventral. One unpaired precloacal papilla. Spicules equal in length, curved and stout, with wide alae reaching almost to the bluntly pointed tips. Gubcrnaculum chitinized and elongated, with a pointed posterior end. FEMALE: Vulva on a slight protrusion, just past mid-length. Vagina directed anteriorly. No caudal papillae. Eggs large, elongated, with thick shells, unernbryonated. Dicussion These worms differ from all previously described members of the genus in the possession of lateral alae. As absence of alae is given as part of the generic diagnosis (Yorke and Mapicstone 1926; Yamaguli 1961 ), this must now be amended to read; alae present or absent. The genus Spironoura is confined to the digestive tract of fish, amphibia and reptiles. However of the approximately 27 species which have been described from reptiles (Yamaguti 1961) all but three arc from chelonians; only S. rnascula (Rudolphi 1819) and S. nitida (Travassos 1920) have been described from snakes. The only previous Australian record of the genus is S. elseyae Johnston and Mawson 1941 from the turtle Elseya dentata. Order ASCARIDIDA Superfamily ASCARIDOIDEA Family ANISAKIDAE Goezia sp. Mati-riai Examinld Queensland Museum G 10292, 2 collected by H. Jones from stomach of Acrochordus granulatus, QM J28740. collected by J. Covacevich and P. Filewood, Iron Range, North Queensland, June 1976. It was not possible to assign these to a species, but they appeared to be the same as specimens recently recovered from Australian sea-snakes and crocodiles, to be described by Sprent (in press). Apart Irom these, the only previous record of this genus from a reptile has been G. gavialidis, (Maplestone 1930), from an Indian CJavial. All other species have been from fish. Order SPIRURIDA Suborder CAMALLANINA Superfamily CAMMALLANOIDEA Caniallaiiides cerberi sp. nov. (Fig. 3, Table 2) Material Examined Hoi OTVnr Queensland Museum G 1 028 1, (3, collected by H. Jones from upper oesophagus of Cerberus rhynchops, QM J23630, collected by J. BredI, Edward River, North Queensland, June 1973. At l.OTYt’i-;: QM GI0282, j, same data as holotypc. ParatypeS: QM GI0283, 3 (5c?, 11 $5, and 9 incomplete specimens, same data as holotype; QM G 1 0284, I , oesophagus of Enhydris polylepis, QM J20282, collected by S. Sterling near Cairns, North Queensland, 1970. Diagnosis Chitinised buccal valves with 12 14 internal longitudinal ridges. Dorsal and ventral chitinous rods projecting posteriorly from edge of buccal capsule. Four small perioral papillae. Seven or eight pairs of pedunculate precloacal papillae and eight or nine pairs of postcloacal pedunculate papillae in males. Unequal spicules. No guber- naculum. Vulva on a pedunculate prominence; female tail slender and ending in a slight knob. Viviparous. Description Fixed worms pale grey in colour, with a black streak running the entire length due to blood or blood products in intestine. Fairly uniform width, anterior end rounded, tail tapering. All specimens were collapsed so diameter could not be measured accurately. Head capsule chitinized, brown. Cuticle finely striated. Mouth elongated dorso- ventrally, with four inconspicuous sessile papillae, one beside each corner. Buccal capsule consists of two buccal valves, each of which consists of two chitinous masses separated by a broad longitudin- al groove. A smaller chitinous body anterior to each mass, from the median aspect of which is a small projection at the worm's anterior extremity. Twelve to lourteen longitudinal ridges inside each valve, only the central ones of which continue to base of capsule. Chitinous ring at base of buccal capsule. Thin curved chitinous bodies at dorsal and ventral edges of capsule run through ring at base of capsule to anterior end of oesophagus; from exterior aspect of each an irregularly-shaped chitinous bar extends posteriorly. Oesophagus divided into two portions; anterior muscular portion stout, with pronounced swelling towards posterior end. Posterior glandular portion almost cylindrical in shape, widening slightly towards posterior end, lumen narrow and tortuous. Intestine broad lumened, filled with blood or blood products, running directly to anus. Nerve ring surrounds anterior portion of oesophagus, JONES: NEMATODES FROM AQUATIC SNAKES 249 Fig. 3. Callamanides cerberi sp. nov. Hololype S G10281, Allotype $ G10282. A, anterior end, lateral; B, anterior end, dorsal; C, en face; D, male tail, ventral; E, female tail, lateral; F, vulva, lateral; G, embryo in uterus. 250 MEMOIRS OF THE QUEENSLAND MUSEUM excretory pore near posterior end of muscular oesophagus. Very small bristle-like cervical papilla seen in one female only. MALE: Less than half length of female. Well developed caudal alae, terminating just before tip of pointed tail. Alae connected anteriorly (not in all specimens), supported by an inconstant number of papillae; six to nine regularly-spaced pedun- culate papillae precloacally, and eight or nine smaller irregularly spaced papillae between cloaca and tip of tail. Spicules unequal in size and dissimilar in shape. Larger right spicule (projects from cloaca in all specimens) alate, tapering to fine slightly curved point. Smaller left spicule not alate, but also with a finely curved tip. No gubernaculum. FEMALE: Mean length 27 mm. Tail long and tapering, terminating in an almost club-shaped knob, with or without a conical tip. Vulva situated just anterior to midpoint of body on rounded pedunculate appendage directly posteriorly. Vulval aperture on dorsal side of appendage, against body of worm, surrounded by two small lips. Muscular vagina runs anteriorly some distance before opening into opposed uterine tubes. All specimens contained many larvae, many of which contained numerous black granules, indicating that they were obtaining nutriment from the females’ blood intake. Discussion Five species of Camallanides are recognised at present, all from the Indian subcontinent; C prashadi Baylis and Daubney 1922, C. piscatori Khera 1954, C. ptyasi Khera 1954, C. dhamini Deshmukh 1968, and C. hemidenta Majumdar 1965. C. prashadi was recovered from Naja bungarus and Ptyas mucosus, as well as from Bungarus fasciatus by Baylis (1929), and from a frog, Rana tigrina by Karve (1930), C. piscatori from Natrix piscator, C. ptyasi and C. dhamini from Ptyas mucosus and C. hemidentata from a freshwater fish, Channa striatus. In addition, Gupta (1959) recorded a single unidentified female Camallanides from a sea-snake, Hydrophis cyanocinctus. All were found in the intestine of their hosts. The five snake species from which these are recorded are not found in Australia, but their distribution overlaps with both that of Fordonia leucobalia and Cerberus rhynchops in the Indo-Malaysian archipelago. C. prashadi has been identified from C. rhynchops from Thailand (British Museum, unpublished) Camallinides cerberi differs from those species described in being larger, in having four instead of six apical papillae, in the absence of a gubernaculum, and in the rounded anterior end. In addition, it differs from C. prashadi in the shape of the dorsal and ventral chitinous bodies which do not project forwards as in that species, in the thicker post-directed chitinous rods, and in the wider groove separating the chitinous masses of each valve. W'hcther or not C. piscatori and C. ptyasi have dorsal and ventral chitinous bodies seems uncertain (Deshmukh 1968). C. cerberi differs from C. prashadi and C. dhamini in having a relatively shorter and more bulbous vulval appendage; the vulval aperture is on the doral side of this appendage, and not on the ventral side as in C. prashadi. C. ptyasi and C. piscatori. In C. dhamini there are 14-16 longitudinal internal buccal ridges, and less inequality in spicule length. These specimens are assigned to a new species on the basis of these various features. Differences in the number and position of the caudal papillae, however, have not been taken into account; they have been used to some extent in differentiating species in this genus but, at least in C. cerberi, they are variable. C. piscatori and C. dhamini were described from one male, C. ptyasi. C. hemidenta, and the redescription of C. prashadi (by Agrawal 1967) were from two males. In view of this, and the fact that three of these (C. prashadi, C. ptyasi and C. dhamini) have been described from the same host. Ptyas mucosus. it seems desirable that a greater number of specimens be examined before differentiation based on these papillae can be relied upon. Further study may question the validity of these species. Suborder SPIRURINA Superfamily GNATHOSTOMATOIDEA Tanqua ophidis Johnston and Mawson 1948 Material Examined Queensland Museum G 10285, G 1 0286, G 10287, G 1 0288 and G 10293 from Acrochordus javanicus QM J23192, J23718, J23189, J23I91 and J23718 respective- ly (see Table 4); 3 6 S and 1 incomplete specimen in author’s collection, stomach of Enhydris polylepis. J23215. Apart from two in the oesophagus and two in the intestine, all specimens were in the stomach, in numbers ranging from seven to 179. Sexes were approximately equal, with a large number of young worms in the two most heavily infected JONES: NEMATODES FROM AQUATIC SNAKES 251 Fig. 4. Heliconema longissima (Ortlepp 1922), G 10291. A, anterior end, lateral; B, male tail, ventral; C, right spicule; D, left spicule. 252 MEMOIRS OF THE QUEENSLAND MUSEUM snakes; clearly it is a prevalent parasite in this species. It was originally described from a freshwater snake, Amphiesma mairii, and from A. javanicus (see Johnston and Mawson 1948), and has not been reported since then. E. polylepis is therefore a new host record. Superfamily PH YSALOPTEROIDEA Heliconema longissima (Ortlepp 1922) (Fig. 4, Table 3) Material Examined Queensland Museum G10291. 37 stomach, 3 oesophagus, 1 rectum Fordonia leucohalia; J 10262; G10290, 12 stomach and 11 proximal small intestine, F. teucobalia. J877; G 10289, one from stomach, F. leucohalia. .123200. The characters and measurements of the worms agree in almost every particular with the original description by Ortlepp (1922) and the redescrip- tion by Ogden (1969), the only differences being that in these specimens there were never more than nine longitudinal tesselated ridges on the ventral surface of the male caudal region, instead of 12 in Ogden’s description, and in the females the vulva was always just posterior to the midlength of the body. Discussion In the measurements of this species by Ortlepp (1922), Li (1934) from Anguilla pekinensis from China and Ogden (1969), the vulva was always slightly anterior to the midlength of the body. These differences, however, together with the lesser number of tesselated ridges in the male, are not thought sufficient to warrant a new species, and may be host-induced variations. There has been uncertainty about the hosts of the original material examined by Ortlepp (1922), which had been labelled ‘snakes, Australia’. Chabaud and Campana-Rouget (1956) in suggesting that the genus Orlleppina erected for the type species by Schultz (1927) was synonymous with Heliconema, suggested that the original specimens had been wrongly labelled, and that they had probably come, as had all other known members of the genus, from eels, and this view was supported by Ogden (1969). The finding of these specimens in aquatic snakes now suggests that maybe the original labelling was correct. The present specimens are therefore the first certain records of the genus which were not recovered from eels. DISCUSSION AND CONCLUSIONS The nematode species recovered were: Eustrongylides acrochordi sp. nov., Spironoura fordoniae sp. nov., Goezia sp., Camallanides cerberi sp. nov., Tanqua ophidis Johnston and Mawson 1948 and Heliconema longissima (Ortlepp 1922). In 24 of the 26 snakes the stomachs were empty; two Fordonia leucohalia TABLE 3; Body Measurements on mm) of Heliconema longissima (ORTLEPP). Males (4) Females (11) range x ± & Length 17.85 -27.37 28.27 -38.55 32.19 ±3.19 Maximum width 0.192- 0.350 0.402- 0.542 0.499±0.069 Nerve ring* 0.231- 0.315 0.259- 0.350 0.298±0.026 Cervical papilla* 0.231- 0.315 0.245- 0.329 0.290±0.025 Excretory pore* 0.366- 0.413 0.406- 0.546 0.4I4±0.049 Muscular eosophagus 0.371- 0.497 0.455- 0.532 0.425±0.028 length Glandular oesophagus 2.47 - 3.62 2.84 - 4.14 3.33 ±0.29 length Muscular oesophagus 0.063- 0.105 0.070- 0.112 0.091±0.006 width Glandular oesophagus 0.119- 0.189 0.140- 0.210 O.I78±O.OI4 width Tail length 0.231- 0.350 0.091- 0.203 0.145±0.028 Spicule length, left 0.482- 0.616 — — Spicule length, right 0.231- 0.259 — — Vulva position! 50-66 56±2 * from anterior end t % of body length from anterior end Host Qld. Museum Locality and Date Tanqua Eustrongylides Camallanides Heliconema Spironoura Goezia (Habitat and diet) Cat. No. JONES: NEMATODES FROM AQUATIC SNAKES 253 rM I r I a o 00 c w a: ^ cd < Os ON . — --Z z u. o — 2-SgSuS y> 6 • ^ ^ ^ :_:z SSS 5 rN m m — oo Tf — (N O' «N m (N «N CC O (N rj- £Q'Q'£ •Q z z ^ w b_ 5 C C 5 CO 7 4J U > r^s. •o o O T3 ov UJUUUJ- OO ^no ^-so • • yi ZZ c— c — - . sj ^ 4) — < c/5 — o O - •* 4) U U C > .> .> .> oai Cxi m m O' O' aa SSr zz = -o -a "o -o 52 t- »- t- t- Z Uj - UJ ^ UJ - UJ . C 3 = 80| => > a 7 u c^cC -a T3 r- O' O' PO CO *> ^ m o r*-> so SO SO — (N •§“S .5 ■ zz^SS o O fsj PN o m m rs IN ao ^z CQ «3 ou 3 "3 k ' O -3 ' k 3 3 •^* ■ O k C k, k S « o . c k> t/5 *0 3 u. 3 S « E ctJ , . .2 S M S = *3 3 W)S C M 3 3 “3 .3 4> =: o rt t; open sea; mainly J28740 Iron Range, N.Q., June, fish and crabs) 1976 254 MEMOIRS OF THE QUEENSLAND MUSEUM (J877 and J 10262) contained remains of large Crustacea. The differing habitats of the snakes probably reflect the different composition of their food, and hence their largely different parasites. Data in Table 4 suggest that T. ophidis and E. acrochordi probably have fresh-water life cycles, S. fordoniae, Goezia sp. and H. longissima marine life cycles, and C. cerheri an estuarine life cycle. These are the first records of snakes as hosts for parasites in the genera Eustrongylides and Heliconema. Goezia have recently been reported from sea-snakes and crocodiles (Sprent, in press), and the great majority of species in the genus Spironoura are found in fish or chelonians. The usual hosts for Spironoura spp. (fish and turtles), Eustrongylides spp. (aquatic birds) and Heliconema spp. (eels) are ones which live in similar environments to these snakes, and with which they may have common food and hence sources of infection. The presence of these parasites in snakes illustrates their physiological opportunism in adapting to phylogenetically different hosts which inhabit the same environ- ment and may be involved in similar food-chains to their more usual hosts. Both £. acrochordi and S. fordoniae show morphological features not found in other members of these genera (cloaca in one; lateral alae in the other), suggesting that they may be evolving away from the main pattern of forms found in these genera. ACKNOWLEDGMENTS I would like to thank Miss. J. Covacevich for her great help in allowing me access to and facilities for the examination of the snakes in the Queensland Museum, and to Professor J. F. A. Sprent for helpful advice and discussion. LITERATURE CITED Agrawal. V., 1967. Some new Camallanoidea (Spirurida) nematodes from fishes, amphibians and reptiles. Annls. Parasit. hum. comp. 42: 327-42. Baylis. H. a., 1929. On the Hosts of Camallanides prashadi Baylis and Daubney 1922 (Nematoda). Ann. Mag. nat. Hist. (10)4: 50. Baylis. H. and Daubney R., 1922. Report on the parasitic nematodes in the collection of the Zoological Survey of India. Mem. Ind. Mus. 7: 263-347. Chabuad. a. G. and Campana-Rouget. Y., 1956. Le genre Ortleppina Schulz 1927, parasite d‘Apodes, et non de Serpents, est Synonyme du genre Heliconema Travassos 1919. Ann. Parasit. hum. comp. 31: 308-9. Deshmukh. P. G., 1968. Camallanides dhamini n.sp. from Rat Snake Ptyas mucosus Riv. Parassit. 29: 119-22. Gerlach, S. a. Von, 1953. Lauratonema nov. gen. Vertreter einer neuen Familie mariner Nematoden aus Dem Kustengrundwasser. Zool. Am. 151: 43-52. Gupta, S. T., 1959. Nematode Parasites of Vertebrates of East Pakistan. III. Camallanidae from fish, amphibians and reptiles. Canad. J. Zool., 37: 771-9. Johnston. T. H. and Mawson. P. M., 1941, The Gallard collection of Parasitic Nematodes in the Australian Museum. Rec. Aust. Mus. 21: 1 10-15. Johntson. T. H. and Mawson. P.M., 1941-47. Some parasitic nematodes in the collection of the Australian Museum. Rec. Aust. Mus. 21: 9-16. Johnston. T. H. and Mawson. P. M., 1948. Some new records of Nematodes from Australian snakes. Rec. i S. Aust. Mus. 9: 101-6. Karve. j. N., 1930. Some parasitic nematodes of frogs ! and toads. Ann. Trop. Med. Parasit. 24: 481-91. Khera. S., 1954. Nematode Parasites of some Indian Vertebrates. Indian J. Helminth. 6: 27-133. Li. H. C., 1934. Report on a collection of parasitic nematodes mainly from North China pt. 2. Spiruroidea. Trans. Am. Microsc. Soc. 53: 174-95. Majumdar. N., 1965. Camallanides hemidenta sp.n. (Nematoda: Camallanidae) occurring in Channa striatus (Bloch). Zpol. Am. 174-175: 222-5. Maplestone. P. a., 1930. Parasitic nematodes obtained ' from animals dying in the Calcutta Zoological Gardens — Parts 1-3. Rec. Ind. Mus. 32: 385-412. Ogden. C. G., 1969. A revision of the genus Heliconema Travassos 1919, Physalopteridae (Nematoda). J. nat. Hist. 3: 423-31. Ortlepp, R. j., 1922. The Nematode Genus Phvsaloplera Rud. Proc. Zool. Soc. London: j 999-1 107. Rudolphi. C. a., 1819. ‘Entozoorum synopsis cui accedunt mantissa duplex et indices locupletissimi” 81 1 pp. (Berolini). Schulz. R., 1927. Die familie Physalopteridae Leiper 1908. Nematodes und die principen ihre Klassifica- j tion. pp 287-312 In ‘Sammlung Helminth Arbeiten t Prof. K. I. Skrjabin gewidment’. (Moscow). Sprent. J. F. A. (in press) Synopsis of the Ascaridoidea: Terranova, Goezia, and Paraheterotyphlum. J. Helminth. Travassos. L., 1920. Genero Florencioia Trav. 1919. Arch. Esc. Sup. Agric. Med. Vet. 4: 21-4. Yamaguti, S., 1961. ‘Systema Helminthum. HI The Nematodes of Vertebrates.’ (Interscience Publishers Inc.: New York). Yorke, W. and Maplestone, P. A., 1926. The Nematode Parasites of Vertebrates.’ (Hafner Publishing Company: New York). Mem. Qd Mus. 18(2): 255-63, pi. 40. [1978] SOME PARASITIC BARNACLES (RHIZOCEPHALA: SACCULINIDAE) FROM PORTUNID CRABS IN MORETON BAY, QUEENSLAND W. J. Phillips Department of Parasitology, University of Queensland ABSTRACT One new species of Sacculina and two new species of Heterosaccus are described and additional morphological features of Sacculina granifera Boschma 1973 revealed by scanning electron microscopy are recorded. Only three sacculinids have been reported from Australian waters. Boschma (1933) described Sacculina duracina from Parthenope longimanus (Leach) at Port Molle, Queensland, and more recently S. granifera from Portunus pelagicus (Linnaeus) from Moreton Bay, Queensland (Boschma 1973). A further unnamed Sacculina sp. from Thalamita sima H. Milne-Edwards from Sydney was mentioned by Haswell (1888). S. granifera was described only recently although it is a serious parasite of the commercial sand crab (P. pelagicus) and has been known for many years (Thomson 1951). In the course of an investigation into the biology of this association, S. granifera and three other sacc-uiinids found on different host species were examined primarily to determine whether the other crab species served as reservoir hosts of S. granifera. This paper contains descriptions of three previously unnamed sacculin- ids and further observations on the morphology of S. granifera. MATERIALS AND METHODS Externae of the parasites were fixed in 5% formalin, Bouin's Huid or 70% alcohol. For sectioning, a small part of the mantle and parts of the visceral mass containing colleteric glands, cypris cell receptacles and ducts were removed, dehydrated and embedded in wax. Sections were cut at 7 - 10/^ and were stained in borax carmine or chlorazol black E. Retinacula were observed by mounting small pieces of mantle in balsam and examining the inner surface. Pieces of mantle taken from young, clean externae were prepared for scanning electron microscopy by dehydration in graded alcohols and xylol and coating with gold dust. All measurements are in millimetres unless otherwise stated, and follow the convention illustrated in Fig. la where A = anterior to posterior dimension, B = dorsal to ventral dimension (The parasite lies on one side), and C = the left to right dimension i.e. the thickness of the parasite between the crabs abdomen and cephalothorax. Abbreviations used in figures are as follows: col. gl. colleteric gland ex. excrescences ext. cut. external cuticle of mantle g. coat. granular coating of mantle j- junction of male duct and cypris cell receptacle 1. can. lymph canals l.c.r. left cypris cell receptacle l.m.d. left male duct mant. mantle me. mesentery m.o. mantle opening pap. papillae ped. peduncle r.e.r. right cypris cell receptacle r.m.d. right male duct rods supporting rods of retinaculum sept. septae vise. m. visceral mass. 256 MEMOIRS OF THE QUEENSLAND MUSEUM Sacculina amplituba nov. sp. (Fig. la - j) Material Examined Hoitn't Ri Queensland Museum W7145, ex Matuta granulosa, Main channel, Moreton Bay, collector W. Phillips, 6. iv. 1973. Par \t VEi s. W7146, a whole specimen, W7147, serial sections, same data as holotype. Diagnosis Panduriform parasites occurring singly. Cypris cell receptacles and ducts outside visceral mass, completely separated, receptacles large, roughly globular without septae passing abruptly into ducts; left receptacle terminates on the right side of the mesentery; diameters of ducts widen until equal to that of receptacles, septate. Tubes of colleteric glands arranged in approximately seven rows, greatest division shows 150-160 tubes. External cuticle covered with spiny excrescences of 35// length formed of hyaline chiton differing from that of the main layers. Retinacula not present. Parasitic on Matuta granulosa Miers. Description Body form: Mature externae are ovoid to rectangular in shape (Fig. la) and brownish in colour except for a pale patch surrounding the posterior stalk. Fine lines can be seen on the mantle, the opening of which is anterior on top of a short muscular tube. Although some specimens were examined other than those designated types, none w'cre found on crabs bearing more than one externa. Dimensions of holotype: A=15, B = 21,C = 8 and a whole paratype A = 10, B = 15, C = 6. Cypris cell receptacles; The cypris cell receptables are in the anterior region of the stalk completely separated from the viscera! mass (Fig. lb,f-h). They are roughly globular and without septae and open into the ducts by a narrow aperture (Fig. le-g). The ducts widen till they become as wide as the receptacles (Fig. Id) then narrow near the opening into the mantle cavity. The terminal portion is free in the mantle cavity (Fig. Ic). The internal surface of the duct bears septae (Fig. Id). The left cypris cell receptacle terminates on the right side of the mesentery (Fig. Ih). Colleteric glands: The colleteric glands are cushion shaped bodies in the middle portion of the left and right sides of the visceral mass (Fig. lb). In transverse section in the area of greatest division of tubes, there are 1 50- 1 60 lubes in up to seven rows. Fig. Ij shows 155 lubes. Mantle: The external cuticle of the mantle is 75-100// in thickness and bears clusters of spines 35-40// in length. The spines are united at their bases in clumps of two to seven; they are of a hyaline chiton different from the cuticle of the mantle (Fig. li). As with all saccuiinids, granular matter collecting round the excrescences may partially obscure them (Fig. li). No retinacula were observed. Remarks The name Sacculina amplituba nov. sp. is proposed because the male ducts widen to a diameter equal to that of the receptacles. In related species the ducts are narrower than the receptacles. 5'. amplituba is similar to 5. beauforii Boschma 1949 from Scylla serrata (Forsk.), S. leptodiae Guirin-Ganivit from Xantho exeratus (H. Milne-Edwards) and S. vankampeni Boschma from Ozius rugulosus Stimpson, but differs from these in having (i) ducts as wide as the receptacles whose ends lie free in the mantle cavity as opposed to ducts not as wide as the receptacles and without free ends and (ii) the left receptacle terminating on the right rather than the left. Furthermore, the colleteric glands of S. beauforti divide into approximately 50 lubes and those of S. leptodiae into less than 20 (Boschma 1949) and so are quite different from the glands in S. amplituba which divide into more than 100 tubes. S. beauforti has the cuticle of the mantle of 120-130// in thickness (Boschma 1955) considerably thicker than that of iS. amplituba (75-100// thick). Apart from the morphological distinctions, S. amplituba should be considered a new species since each of the most similar species all occur in a different host genus and the work of Fratello (1968) on chromosomes indicates a high degree of host specificity among saccuiinids. Sacculina granifera Boschma, 1973 The most important characteristic of S. granifera is the structure of the excrescences of Fig, 1; Sacculina amplituba: a, whole parasite — left side. A, B and C are the anterior to posterior, dorsal to ventral and left to right dimensions respectively; b, left side with left side of mantle removed, scale = 4 mm; c-h, transverse sections of posterior part of parasite showing male organs, scale = 1 mm; i, transverse section of external cuticle of mantle showing excrescences, scale = 40//; transverse section of colleteric gland in region of greatest division of tubes, scale = 300// . PHILLIPS: PARASITIC BARNACLES FROM CRABS 257 258 MEMOIRS OF THE QUEENSLAND MUSEUM the external cuticle described as irregularly globular (Boschma 1973). Scanning electron micrographs (Plate 40. Pig. l.b) reveal that each of the globular excrescences bears many small spines. F'urther more, Boschma described the collctcric glands of N. granifera as dividing into 40 lubes in one row. Specimens examined by me show the collctcric glands to be divided consistently into up to 100 lubes in the region of greatest division. Hetcrosaccus lunatus nov. sp. (Fig. 2 a-i) MATI RlAl EXAMIM D Hoioiapi Queensland Museum W7148, ex Charyhdis callianassa, Moreton Bay, collector R. Bishop. Feb. 1975. Pnrm'iPIs Queensland Museum W7149 ex Charyb- dis callianassa. Moreton Bay, collector R. Bishop, Nov. 1974, three whole specimens on one host. Queensland Museum W7150. ex C. callianassa. Moreton Bay, collector W. Phillips, 24/v/1973, sections. Diagnosis Kidney shaped parasite, more than one of which may occur on the one host. Cypris cell receptacles within visceral mass and deeply crescentic, terminating near colleleric glands and containing sponge-like meshwork. Receptacles open widely into straight ducts also with sponge-like mcshw'ork; ducts open near stalk; receptacles and ducts surrounded by lymph spaces. Collctcric glands posterior, with 60-70 lubes in five to six rows in region of greatest division. External cuticle with papillae 15/; long and 10// apart, of same material as mantle. Retinacula rounded flaps of tissue occurring singly each with four to six rods. Parasite of Charyhdis callianassa (Herbst). Description Body form: The externa (Pig. 2a) is kidney- shaped having a wide mantle opening with poorly developed musculature. The numbers of cxternac per crab varies from one to three, the exlernae being smaller where there arc more than one on one host. Dimensions of the holotype: A = 10, B = 19, C = 7; and of whole paratypes (i) A = 10, B - 20, C = 9, (ii) A = 8, B = 13, C = 6, (iii) A = 8, B = 11, C 6. Cypris cell receptacles: The male organs are confined to the posterior dorsal portion of the visceral mass w'hcre they are surrounded by lymph canals. The rest of the visceral niass is without pronounced lymph canals. The ducts which open close to the stalk (Fig. 2,h) are of a consistent diameter of about 400// and have a sponge-like structure for most of their length. The lumen of each duct is 40-50// wide. The ducts are not significantly convoluted. They pass gradually into the receptacles which arc only slightly wider than the ducts (Fig. 2 d- g). The receptacles have a deep curvature and terminate near the colleteric glands (Fig. 2g). Colleteric glands: The colleleric glands are cushion-shaped bodies lying in the posterior half of the visceral mass near the middle. In the region of greatest division there are 60-70 tubes in five to six rows (Fig. 2i). There is no chitinous lining but villiform projections arc present on the inside. Mantle: The external cuticle of the mantle is 15-20// thick and is covered with papillae approximately 15// long and 10// apart (Fig. 2c). The papillae are projections of the cuticle of the mantle and may be partially or wholly obscured by a granular coating (Fig. 2b). Scanning electron micrographs (Plate 40, Fig. b) clearly show the shape and distribution of the papillae and the granular coaling. Retinacula (Fig. 2c) are single rounded flaps of tissue about 20// wide each containing four to six rod-like bodies. The retinacula are about 200// apart. Remarks The name Helerosaccus lunatus nov. sp. is proposed because of the deeply crescentic nature of the cypris cell receptacles. Heterosaccus multilacinensis nov. sp. (Fig. 3 a-i) Mati'riai Examimd Hoi.otypi. Queensland Museum W7151, sections, ex Carybdis iruncata, East Moreton Bay, collector W. Phillips 6/iv/l973 pARATYPiS: W7152. cx C truncQta. four specimens on host, same data as holotype. Diagnosis Kidney-shaped parasite more than one of which may occur on the one host. Cypris cell receptacles Fi(. 2: Helerosaccus lunaius: a, whole parasite — left side, scale = 3 mm; b, transverse section of external cuticle of mantle, scale = 10//; c, internal retinacula, scale = 25//; d-h, transverse sections of posterior part of parasite showing male organs, scale =• 1 mm; i, transverse section of colleteric glands in region of greatest division of tubules, scale = 200// . PHILLIPS: PARASITIC BARNACLES FROM CRABS 259 260 MEMOIRS OF THE QUEENSLAND MUSEUM shallow crcsccnlic, not surrounded by lymph spaces; ducts slightly convoluted opening away from stalk; meshwork present in receptacles but not in ducts. Collcteric glands in posterior part of visceral mass, region of greatest division of tubes shows 40-60 lubes in four to five rows. External cuticle with papillae 15//' long, close together at base and separated by 5// at tip. Papillae of same material as cuticle. Retinacula of one to six rounded Haps on a common base with one rod per flap. Parasite of Charybdis truncata (Fabricius). Description Body form: The externae (Fig. 3a, b) are kidney-shaped with a wide mantle opening showing poorly developed musculature. The number of externae per crab may be more than one. Dimensions of paratypes (i) A = 10, B = 17, C = 4, (ii) A = 9, B = 10, C = 7, (iii) A = 10. B = 19, C = 6, (iv) A = 18, B = 14, C = 4. Cypris cell receptacles: The male organs are confined to the posterior dorsal portion of the visceral mass and are not surrounded by lymph spaces although lymph spaces are seen commonly in the visceral mass (Fig. 3 a,d-g). The ducts which open close to the stalk are somewhat convoluted and do not possess a sponge-like structure (Fig. 3 e-g). They have a diameter of 220 240// and the lumen is 50-70// in diameter. The ducts pass gradually into the receptacles which have a shallow curvature and do possess a sponge-like structure. The receptacles arc two to three times as wide as the ducts (Fig. 3 d,e). Collcteric glands; The cushion shaped collcteric glands arc situated in the posterior part of the visceral mass towards the middle. The region of greatest division of tubes shows 40-60 tubes in four or five rows (Fig. 3h). There is no chitinous lining, but the lumen is lined with villiform structures. Mantle: The external cuticle of the mantle is approximately 50// thick with papillae 15// long which are extensions of the cuticle (Fig. 3b). The papillae touch at the bases and taper to a blunt point. They arc about 5// apart at the tips. Retinacula (Fig. 3c) consist of one to six rounded flaps of chiton on a common base, each flap containing two or three rods which may project out of the flaps and appear to extend into the base. The flaps are from 18-20// long and spaced about 200// apart. Remarks The name Heterosaccus multilacinensis nov. sp. is proposed because of the many flaps on each retinaculum. Boschma (1963) described Heterosaccus papillosus from several species of the genus Charybdis including C. callianassa. It is similar to, yet distinct from, the two species described here. //. lunatus differs from //. papillosus in having (i) wider receptacle ducts opening close to the stalk, (ii) deeper curvature of the receptacles, (iii) greater division of the lubes in more posteriorly placed collcteric glands, (iv) longer more closely placed papillae on the mantle, and (v) retinacula consisting of flaps with rods rather than spindles. H. multilacinensis differs from H. papillosus in having (i) ducts opening close to the stalk rather than at a distance from the stalk, (ii) shallower curvature of the receptacles, (iii) greater division of tubules in more posteriorly placed collcteric glands, (iv) papillae closer together and of a different shape, and (v) retinacula consisting of many Haps w ith rods. H. multilacinensis differs from //. lunatus in having (i) narrower more convoluted male ducts without a sponge-like structure, (ii) less division of the tubes of the collcteric glands, (iii) longer, thicker, more closely packed papillae, and (iv) retinacula consisting of several Haps on one stalk. DISCUSSION Undoubtedly the rhizoccphalan fauna of Australia will be found to be more numerous than is presently recorded. The influence of these parasites on the biology of their hosts is considerable and where the hosts are of commercial importance they produce considerable losses to the industry. As yet no sacculinid has been found to parasitize the commercial mud crab Scylla serrata in Australia although S. beauforti and Loxothylacus ihlei Boschma 1949 have been described from this species in Indonesia. Discrimination of species of rhizoccphala is difficult since insufficient is known of the details of their life cycles or the variability of characteristics with the age and maturation of the externa. Scanning electron micrographs of the exterior surfaces of the mantle of young externae should provide accurate and detailed information on one rather important taxonomic feature, viz. the form of the cuticular excrescences. Clear differences can be seen between Sacculina amplituba and Heterosaccus lunatus (Plate 40). PHILLIPS: PARASITIC BARNACLES FROM CRABS 261 Fig. 3: Heterosaccus muliilacinensis: a, whole parasite, left side, with left side of mantle removed, scale - 3 mm; b, papillae of external surface of mantle, scale =10//; c, retinacula, scale = 25//; d-g, transverse section of posterior part of parasite showing male organs, scale = 1 mm; h, transverse section of colleteric glands in region of greatest division of tubes, scale = 300//. 262 MEMOIRS OF THE QUEENSLAND MUSEUM Probably the most promising means of discriminating species would be chromosome techniques as described by Fratello (1968) who was able to discriminate several species previously described as one. ACKNOWLEDGMENTS The sacculinids described in this paper were related to a post-graduate honours project in the Department of Parasitology, University of Queensland. 1 would like to express my thanks to Professor J.F.A, Sprent for permission to use departmental facilities. I would also like to thank Dr L.R.G. Cannon for assistance in preparing the manuscript; the C-S.LR.O.. for use of its boat for collecting specimens. Mr J. Hardy of the Electron Microscope Unit and Mr J. Mines for advice on histological matters. The late Dr H. Boschma of Leiden Museum, Netherlands also provided valuable information concerning this work. LITERATURE CITED Boschma, H., 1933. The Rhi/occphala in the collection of the British Museum. J. Linn. Soc. 38: 473 552. 1 949. Sacculina beauforti and Losolhylacus ihlei, two Rhi/ocephala of the crab Scylla serraia (Forsk.). Bijdr. Dierk. 28: 41-6. 1955. The described species of the family Sacculin- idac. Zool. Verb.. Leiden. 27: 1-76. I 967. A Rhi/ocephalan parasite of the crab Charyhdis callianassa (Hcrbsl.). Prov. K. ned. Akad. Wet. 66C; 132 3. 1973. Sacculina granifera nov. spec., a rhizocephalan parasite of the crab Portunus iPortunusj pelagicus (Linnaeus) from the coast of Queensland. Proc. K. ned. Akad. Wei. 76C: 313-8. Frathi.io. B-, 1968. Cariologia c lassonomia dci Sacculinidi (C'irripedi, Ri/ocefali). Caryologia 21 (4): 359-67. H \s\\ 1 t I , W’.A., 1 888. On Sacculina infesting Australian crabs. Proc. Linn. Soc. N.S.W. 3: 171 1-2. Thomson, J.M., 1951. Catch composition of the sand crab fishery in Morelon Bay. Aust. J. mar. Fre.shwat. Re.^. 2: 237-44. MEMOIRS OF THE QUEENSLAND MUSEUM Pl.ATH 40 A: External surface of mantle of Sacculina granifera 1. x500; 2 x2500 B: External surface of mantle of Heterosaccus lunatus 1. x450; 2 xl350. PHILLIPS; PARASITIC BARNACLES FROM CRABS Plate 40 Mem. Qd Mus. 18(2): 265-71, pis. 41-42 [1978] THE OLD BURIAL VAULT AT NORTH QUAY, BRISBANE J. C. H. Gill Queensland Museum ABSTRACT A brick structure accidentally uncovered in 1972 during construction of the Riverside Expressway, Brisbane, opposite the eastern side of the junction of Herschel Street and North Quay, has been determined to have been a burial vault dating from the penal settlement days of Morcton Bay and Brisbane Town. This probably represents the last convict structure to be located in this area. It has been assessed as fully as possible from the limited record able to be made at the time of discovery, as within an hour of this it had been destroyed totally. In the Queensland Museum are eleven sand stock bricks, registered H9750 to H9760 inclusive, collected by the Museum on 23 August 1972; their in situ provenance is recorded as ‘from suspected grave site on Brisbane River bank 100 yards [90 m] west of Tank Street’. They provide a slender base on which to assert they were part of a structure dating from the convict days of Brisbane Town, yet there is evidence which affords positive indications of the time when the bricks were used, of the structure of which they were part, and of the use to which that structure was put. The narrative which follows will incorporate the historical evidence. Then the structure as such will be dealt with as far as it is possible to make a reconstruction from the relics available. Historical Evidence On 23 August 1972 a bulldozer working on the Brisbane Riverside Expressway construction opposite the junction of North Quay and Herschel Street dislodged some bricks buried in the soil, and disclosed an aperture in the sloping bank behind which lay a buried chamber. Examination revealed a barrel-vaulted, brick structure, over which a coating of lime mortar had been applied internally. Main Roads Department officers supervising the construction halted work and informed the Royal Historical Society of Queensland of the find. The author was contacted and requested Queensland Museum assistance to assess the historical value of the discovery. Very limited ^ime was allowed for investigation before construction work resumed. The urgent situation precluded detailed archaeological investigation. Fortunately a local television station, QTQ 9, filmed external and internal views, and later donated the film to the Queensland Museum. Rough sketches were made of the structure and some bricks were taken for later examination. As nothing further could be done, the Main Roads Department officers were informed and within minutes the bulldozer operator, who had kept his motor running all this time, had destroyed the structure. A press photographer arrived too late to do other than photograph a few bricks lying where the structure had been, resulting in coverage in the ‘Courier- Mair, on 24 August 1972, p. 3. With the wisdom of hindsight it seems to have been a rather inglorious episode. Yet the element of surprise and the pragmatic attitude of the people concerned with the site gave no opportunity for complete survey. Furthermore the nature and purpose of the structure was at first not clear. The situation could not compare, for example, with the discovery, again per medium of a bulldozer, on 30 March 1974, of an early brick vault in what had been the earliest official burial ground in 266 MEMOIRS OF THE QUEENSLAND MUSEUM Fk. 1. Plan based on Assistant Surveyor R. Dixon's 1840 survey, with outlines of modern streets superimposed. Sydney from 1793 to its closure on 27 January 1820. The purpose of the structure was clear — it contained a collapsed wooden coffin within which were bone fragments and Sydney University archaeologists had three days to assemble a team to survey it. Resulting public interest gave the vault a further reprieve and in the end it was dismantled for reconstruction at the Old Sydney Town Project. Finally a thoroughly researched monograph was published (Birming- ham and Liston 1976). The site of Brisbane's earliest burial ground was investigated previously by Gill (1959-60, 1974), who recalled that the structure was only yards away from isolated memorials on North Quay known as ‘the children’s graves’. Relics from Brisbane’s penal settlement era are few. The absence of a burial ground from this era is particularly notable when one considers the plethora of early memorials in the other states of Australia. Controversy has raged as to the site of Brisbane’s earliest burial ground. Yet it has been proved beyond doubt that the penal settlement’s earliest and only burial ground was that shown in the town maps of Assistant Surveyor Robert Dixon (1840)* and Surveyor Henry Wade (1842)t situated on the north bank of the Brisbane River at the northern end of the William Jolly Bridge in the area bounded by the present day Skew Street and Eagle Terrace. One of the causes of controversy was the presence of three (at least) memorials on the river bank at North Quay directly opposite the eastern corner of North Quay and Herschel Street, immediately above a prominent rock face which rises abruptly out of the river. The Royal Historical Society of Queensland has a photo- graph (said to have been taken about 1876) which shows memorials in situ at this point (Plate 41, fig. I). This photograph, acknowledged to Mr H.A. Jones was donated in 1921 and was reproduced in the ‘Brisbane Courier' on 25 October 1913 at p. 12. It w^as captioned: ‘The Soldiers Graves on North Quay. A controversy has for some time been waged in the “Courier” as to the exact location of the soldiers’ graves in the old burial ground on North Quay. Our photo shows their position (indicated by a X).' The memorials have long been gone from North Quay but they are still to be seen in Toowong General Cemetery. On 5 October 1881 the memorials at North Quay and any remains (under licence * Queensland Survey Office Map M.T.3 t Queensland Survey Office Maps 6 and B.il82 GILL: OLD BURIAL VAULT 267 issued by the Colonial Secretary on 23 August 1881) were removed to Toowong.* Only three memorials and remains were removed. Whether there were any more burials on this site is a matter of doubt. The three memorials all commemorate young children — William Henry Roberts, son of Charles Roberts of the Commissariat Department who died of illness on 15 November 1831 aged 3 years and 2 months), Peter, son of Private Peter McCauley of H.M. 17th Regiment of Foot, accidentally drowned on 5 January 1832, aged 5 years and 8 months; and Jane, daughter of John Pittard, former Colour Sergeant of the 57th Regiment, who died on 29 January 1833, aged 12 months and 13 days (Plate 41, fig. 2). The McCauley and Pittard memorials are the usual headstone and footstone types; the Roberts memorial is a catafalque or table monument type with the inscription on the flat top. The memorials show evidences of the ravages of time. In 1962 the Brisbane City Council took steps to refurbish them. The decay of the first two memorials has been arrested, but the inscription on the Roberts memorial is now lost and is, in fact, covered by a coat of stucco. Fortunately it was recorded in its entirety in 1913 with some errors which, except for the age (5 instead of 3 years) were corrected by Gill (1959-60). Assistant Surveyor Dixon in his 1840 map, in addition to the official burial ground, showed two small squares, habitually used by surveyors to indicate isolated graves, at a point corresponding to the location of The children’s graves’ (Gill 1959-60). Steele (1975) indicates that the Reverend John Vincent, in his initial report to Archdeacon Scott, complained that the settlement’s ‘burial ground is at a very remote distance, is too small and so very difficult to be dug . . . that two men with the utmost exertion are scarcely able to prepare a grave in six or seven hours’. If the children were buried where the memorials were located, it would have required the cutting of kists in solid rock, and the purpose of the structure unearthed on 23 August 1972 now becomes clear. It was a burial vault sited on or slotted into the rock in which bodies were placed whilst sorrowing relatives were allowed to place memorials around it, a much easier process than cutting a full-size grave into the rock each time. Evidence of the existence of the vault was available, but the disappearance of the structure beneath the earthen embankment of the widened North Quay in the latter part of the nineteenth century and the presence up to 1881 of the table monument to the Roberts’ child led to the belief that references to the vault were references to the table monument. The removal of the memorials from North Quay on 5 October 1 88 1 excited no comment from the press of the day. In fact it was not until Friday, 21 October 1881 that the ‘Brisbane Courier’ on page 2 remarked: ‘The old landmarks of Brisbane are rapidly disappearing and although by many the removal of these mementoes of early days may be regarded with some lingering feelings of regret the improvement which will result in the general appearance of the city must be a matter of satisfaction*. The disappearance of the old convict barracks and old buildings on the corner of Queen and Albert Streets is favourably commented upon, whilst the fact that the dreary looking goal at Petrie Terrace will soon become a thing of the past also is editorially approved. Then: ‘Another “landmark" that has disappeared from within the boundaries of the city are the old graves on the North Quay. In these graves were the remains of three bodies that have been under the ground for some fifty years. By the order of the Colonial Secretary, the remains with the tombstones that were over them, have been removed to the cemetery at Toowong, that being considered a more fitting place for them, where it will be less trouble to keep the graves in order, and where they will be subjected to le.ss desecration*. It was not on-the-spot reporting by any means and it is not surprising that the vault escaped notice by the press. There the matter rested for 32 years whilst improvement works along the North Quay greatly altered the old landscape. Professor F. W. S. Cumbrae Stewart through the columns of the ‘Brisbane Courier’ of 30 September 1913 (p. 4) drew attention to the childrens memorials in Toowong Cemetery, expressed the belief that they had been removed thence from North Quay in 1876, and sought information as to where the gravestones originally stood on North Quay. * John Oxley Library MS.O.M. 65/18, Queensland State Archives [QSA.] COL/A319, 3694; QSA. COL/G15, p. 585, 81/1233. , „ . u u t John Oxley Library ibid. Mr Melville the sexton at Toowong Cemetery in 1940 says it appears to be 3 but could be 5 years, on account of weathering of the stone. In 1881 (59 years earlier) the age is given as 3 years and 2 months. The stone was then 50 years old and the inscription much more legibde. 268 MEMOIRS OFTHE QUEENSLAND MUSEUM Between 30 September and 5 November 1913 the ‘Brisbane Courier' published fourteen letters on the subject of the ‘Old Burial Ground' (Gill 1959-60). On 7 October 1913 the ‘Brisbane Courier’ (p. 4) published a letter from a Mr Thos. Illidge who stated he was on the spot when the remains were dug up by Mr D. Hannah of Paddington. Published with the letter was an interview by a Courier reporter with Mr David Hannah of Rosalie who said he had assisted in the removal of headstones on the river bank from North Quay to Toowong. It is significant that Mr Hannah did not refer to the removal of remains. On 16 October 1913 (at p. 7) ‘Sixty Years a Queenslander' wrote about ‘the monumental Slone' which in earlier years marked the spot known to old residents as the ‘soldiers' graves'. He referred to the difference of opinion which seemed to exist as to the exact site. This he regarded as quite natural for during the last few years (i.c. prior to 1913) ‘surface appearances have changed or become obliterated’. He also says: ‘In the 'fifties and 'sixties of last century much of the river frontage along the North Quay was covered with a tangled growth of lantana. which could only be penetrated by a bird-nesting boy or a billygoat. As the former, 1 have often “worked my passage" through this scrub and my memory fixes the site of the graves as about midway between Tank and Herschel Streets and just above a prominent rock which here rises abruptly over the river.’ (Gill 1959-60). He was almost on the correct site and indicates that with the profusion of under-growth making the site inaccessible the general public had come to refer to them as the ‘Soldiers' Graves’. On 20 October 1913 (p. 6) the ‘Brisbane Courier’ published a letter from a Mrs Lucy Sonnonschein (nee Wynn) of Warwick. She said: ‘The old burial grounds were situated opposite the present Roma Streets goods shed gates. After the burial ground was removed, the land was built upon and occupied by Davies (Davis) the Jew. Joseph Jayes, Bill and Joe Jewell, carpenters, Joshua Peter Bell, and Bulger, a wine merchant. New shops have replaced these dwellings. This burial ground was known as the “Soldiers’ Graves". On the bank of the river in North Quay there was a large vault, opposite Paddy Pacey's milking yard, which marked the spot where some officer was buried.' (Gill 1959-60). The existence of the burial vault, rediscovered in 1972, being unknown, the author as already slated believed this to be a reference to the table monument to the Roberts' child. Yet on 5 November 1913 the ‘Brisbane Courier’ published (in the Courier Home Circle Sup- plement at p. 6) the final two letters in the burial ground debate. Mr A. E. Campbell of Rockhamp- ton refers to the sketch in the ‘Courier' of 25 October of the “soldiers' grave'*. ‘Yes. that is about the spot, only high up, about ten yards from the fence, [the X was placed too low on the photograph], and there is a high stone face projecting towards the river. It was the grave of an officer who went out riding and his horse carried him against a tree and broke his neck. I think his name on the lop slab was “Lawson", The tomb must have been shifted in the early ’sixties. The bottom end near the river was broken, and it acted as a cave. Many a one slept there for the night, for it was a cosy camp for a cold night.’ (Gill 1959-60). Here again the author believed the reference w-as to the table monument; but it would have made cramped quarters and access would have been grossly difficult. The mean dimensions of the monument are only 1-905 m by 0-990 m by 0-852 m high. The brick vault was in a different category — in the 'fifties and 'sixties it was buried beneath lantana; any remains in it would have been moved in 1881; (in any event the remains of young children would not have the durability of those of adults); the access, if the end near the river had broken away, would have been easier, and air circulation and animal scavengers would have removed any associations with its grisly past by the time ‘down and outs' began to camp in it. It is considered that these reports provide sufficient evidence of the existence of a vault. In addition to its existence we also have evidence ol its use. .A macabre story recounted by J. J. Knight (1895) furnishes this. Talking of the offences for which a convict could be sent to the triangle Knight says: ‘A very common offence was the purloining of a few cobs of corn and potatoes, the chief ingredients of the convicts' much prized “fiddle-cake", the love for which caused many sore backs at the triangle, and weary legs on the treads. These two things having been obtained, the corn was ground on an improvised grater made usually of a piece of tin or zinc, in which holes had been punched.' Alter describing the grating and recipe for the delicacy, Knight goes on ‘But it was often more difficult to obtain the material for the grater than the corn and potatoes, and in one case at least a convict finding this so resorted to a very questionable means of gratifying his desires. The overseer at Eagle Farm was unfortunate enough to lose one of his children, and the body having been placed in a tin box, was laid in one of the vaults on the river bank near Herschel Street. This fact was. of course, known to the prisoners, and a day or so after the funeral one who had so urgently desired the possession of a “grater" GILL: OLD BURIAL VAULT 269 effected an entrance to the vault and taking out the dead body of the child found in the box the material for the manufacture of this necessary implement of the “fiddle-cake" maker. Fortunately for the sacrilegist, he was not found out, and was enabled to make many specimens of that delicacy, which was described by one who had often partaken of it as being “better than any pie going".’ Apart from anything else this indicates that a measure of improvisation was required still in the undertaking line. If there were no coffins readily available for children of officials one wonders what sort of hurriedly thrown together boxes were used for convict burials. Knight talks of one of the vaults on the river bank near Herschel Street. The available evidence indicates there was only one such vault. It appears to have carried no marks of identification (the adjacent memorials made this unnecessary). Mr Campbell does mention the name Lawson, but he appears to be confusing the inscribed Roberts' monument with the vault. In Plate 41, fig. 1 the Roberts' monument is plainly discernible but the vault is not, although Mr Campbell has no doubt about the location of the latter. It was because the vault carried no identifying marks or inscription that all the stories about soldiers’ graves and officers' burial places came into being. The correct function of the vault probably occurred to no one; the memorials were there and it would not be known generally that they were sited on solid rock. Once North Quay was widened in the late nineteenth century the footpath on the river side of the roadway was cut off from the steep river bank by a wooden picket fence of sturdy construction. Apart from wooden stairway accesses to the Commercial, Brisbane and Brisbane Grammar School Rowing Club Boatsheds there was no access and for upwards of half a century until World War II the North Quay river bank from the O'Connor Boathouse (Commercial Rowing Club) upstream to the Grey Street (William Jolly) Bridge became a terra incognita. Even the erection of air raid shelters in 1942 resulting in the partial disappearance of the fence followed by its nearly complete demolition in the post-war period brought about no interference with the river bank itself. It was not until the construction commenced of the Riverside Expressway section of the South East Freeway that the riverbank began to be altered substantially. In August, 1972 the work began on the expressway exit to Herschel Street. This meant carving away a section of the riverbank; before the bulldozer got down to the rock, it blundered into the old vault with the results already indicated. The vault could well have been coeval with the Windmill and the Commissariat store*, and may well be the last of the Brisbane Town convict structures which remained to be rediscovered. Description of the Vault The vault itself was constructed of sand-stock bricks (as they are commonly known) with mean approximate dimensions of 205-9 mm x 63-5 mm X 107-9 mm. The dimensions of the vault as estimated by Mr A. Sweetser, Queensland Museum, in a drawing he made (Figure 2) are given as 1-53 m x 1-83 m with a barrel vaulted roof rising above 1-53 m high side and end walls. The entrance at the northern end had been bricked up after any remains had been cleared from the vault. Two wooden slabs lay on the floor covered by rubble and a partly burned piece of hardwood plank lay near the southern end of the vault. There was some evidence of dampness in the earth floor but no signs of the vault ever having been inundated with water; the run-off from North Quay to the Brisbane River was steep enough to prevent water from gathering and lying in the vault. The only wall of the structure to show up in full width in the movie film was the northern wall, and estimating from the known lengths of the bricks collected from the site the mean width of the wall could be 1-72 m, 019 m more than Mr Sweetser's estimate (Figure 2). The side wails were grooved for their entire lengths at two levels, which indicated an intention to have the coffins resting on planks (fitted into the grooves) above ground level and provisions for an upper layer of coffins if needed. An analysis of the lime mortar used to bind the bricks and plaster the interior of the vault has been made by the Queensland Government Chemical Laboratory. The analytical report (Appendix I) established the source of the burnt lime used in the mortar as Ipswich; a clear link with the penal settlement days. As regards the bricks, what were known as the old and new brick kilns respectively both were located within 230 m of the vault as shown on Figure i (Steele 1975). So the raw materials for the brickwork were close at hand. Plate 42. fig. 1 shows the intrusion of some tree roots, but the lack of moisture had inhibited the growth of these. Owing to the gross interference with ground levels, even before the discovery of the vault as ♦Both completed in 1829. 270 MEMOIRS OF THE QUEENSLAND MUSEUM well as subsequently, no soil profile could be taken. However it was obvious to the eyewitness that the vault had been covered by fill from elsewhere than the river bank itself and that the river bank proper began at the base of the structure. Apart from the planks within the vault no timber beams or lintels were noted in the walls of the structure; it appears to have been small enough not to require limber to span any openings or archwork which the bricks themselves could not support. No other features remain to be remarked upon. Thus a few feet of film and eleven bricks at the Queensland Museum, with this account, will be the only record of a structure dating from the early penal settlement days of Brisbane Town. LITERATURE CITED Birminomam. J., and Liston, C., 1976. Old Sydney burial ground 1974 — emergency excavation. Studies in Historical Archaeology No. 5, vii + 40 pp. Gll.L, J. C. H., 1959 60. Some investigations into the site of Brisbane’s earliest burial ground. J. R. Hist. Soc. Qd. 6 (2): 388-417. 1974. Dc Mortius. /^7Wiiso\. P. D. (Edit.) Moreton Bay Sesqui - Centenary 1824-1974 (Library Board of Queensland: Brisbane). Kmout. j. j., 1895. In the Early Days, xi + 390 pp. (Sapsford and Co.: Brisbane). STHta.i:. J. G., 1975. ‘Brisbane Town in Convict Days 1824-1842.' xxii + 403 pp. (University of Queensland Press: Brisbane). FiG. 2. Diagram of the vault, after a sketch by Mr A. Sweetser held in the Queensland Museum. GILL: OLD BURIAL VAULT 271 APPENDIX 1 Government Chemical Laboratory William Street, Brisbane, Q. 4000 16th November 1976 Dr. D. J. Robinson Qld. Museum Gregory Terrace & Bowen Bridge Road, BRISBANE. Ref: DR;DF HlO/5 Dear Sir, On Tuesday 8lh November, soft portions of the white material in mortar samples H9255 and 9578 provided by the Museum were scraped out and this material on analysis gave carbon dioxide, 9% acid soluble silica and 6% magnesium as magnesium oxide. Aslo present is calcium oxide, iron oxide and alumina. The Geological Survey Office was contacted and confirmed that the results of an early analyses of the Limestone Hill Limestone was Loss of Ignition 43.8% Total Silica 5.7% Iron oxide 0.9% Alumina 1.1% Calcium oxide 27.5% Magnesium oxide 20.6% This indicated that this mineral is an impure mixture of calcium and magnesium carbonates. Other areas of limestone in the Ipswich area have similar compositions. The presence of acid soluble silica and the magnesium oxide indicates that the burnt lime used in the mortar sample probably came from the Ipswich district which seems to be the only source of limestone in the local area other than sea shells. The Ipswich deposits have variable compositions but all seem to have more than 5% magnesium carbonate. Microscopic and hand lens examination showed no shell particles and it is not expected that shells burnt to calcium oxide would contain acid soluble silica and magnesia. Warwick and Gympie limestones are much lower in magnesia. D. Mathers Director MEMOIRS OF THE QUEENSLAND MUSEUM Plate 41 Fig. 1 : The memorials in situ ca. 1 876 - photograph in Royal Historial Society of Queensland collections. Fig, 2; Children’s memorials at Toowong Cemetery. GILL; OLD BURIAL VAULT Plate 41 1 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 42 Fig. 1: Interior of vault-photograph from QTQ 9 16 mm film. Fig. 2: Riverside Expressway showing remains of bold, rocky face above which the vault was located. GILL: OLD BURIAL VAULT Plate 42 Mem. QdMus. 18(2): 273-319, pis. 43-103 [1978] THE CORAL GENUS ACROPORA (SCLERACTINIA: ASTROCOENIINA; ACROPORIDAE) IN THE CENTRAL AND SOUTHERN GREAT BARRIER REEF PROVINCE Carden C. Wallace Queensland Museum ABSTRACT The genus Acropora (excluding subgenus Isopora Brook) is revised from zonation studies on two reefs, and incidental collecting from many other localities in the Great Barrier Reef Province. As a result of field studies, many species are placed in synonymy, and a resulting forty species are redescribed. The revision includes description of field characteristics of the species, many of which were previously known only from small skeletal series. Problems of identification and delimitation of the various species are discussed. In the Great Barrier Reef Province, as in most Indo-Pacific reef areas, the regions of densest coral growth are regions dominated by the branching coral genus Acropora. Over 300 species have been described for this genus. Some 80 have been recorded from the Great Barrier Reef, and many more would have been added to this list had the reef slopes and deeper waters been more accessible to early collectors. The original 300 rtiay be reduced to around 70 by synonymy, and the 40 species here treated are considered to represent at least 90% of the branching Acropora in the Great Barrier Reef Province. The following is a working paper on the taxonomy of this neglected genus. It is based on regional studies in the Great Barrier Reef region, and in particular on line transect studies carried out at Big Broadhurst Reef (Wallace 1975, Wallace and Dale 1977) and Bushy-Redbill Reef (Wallace and Lovell 1977, Wallace unpublished). It is an attempt to open the study of the genus to workers in all aspects of coral study and to this end it concentrates on field description of the species. it is clear to the author after some experience with the genus that few species can be fully understood by simple morphological studies of colonies, even with the addition of ecological information. Even the population cannot be considered a sufficient unit for study as there are often other key species in the surrounding assemblage which may look like, physically interact with, or affect the members of the population in some way. This paper seeks to indicate such problems. Taxonomic History The name Acropora Oken, 1815 was officially validated in 1963 (Boschma 1961, China 1963). It came into general use with Verriil (1902), although not without some argument and nostalgia for the previously widely used Madrepora Lamarck: ‘it is with a feeling of regret that we are forced to abandon the use of a generic name which remained unquestioned by four generations of authors during the golden days of systematic zoology’ (Mayor 1924, p.vii). Many authors have described species in this genus. Few have attempted to synonymise or group species, fewer still to define and standardise terms. The most notable single contribution to the taxonomy of the genus was that of Dana (1846) who gave very careful attention to his specimens collected during the United States Exploring Expedition. His arrangement of species according to colony shape and radial corallite shape without defined subgeneric status remains the simplest and most acceptable treatment of the genus. Brook (1893) produced the only true mono- graph of the genus, summarizing all previous work and re-describing all described species. He ordered the species into subgeneric groups, attempting to 274 MEMOIRS OF THE QUEENSLAND MUSEUM take into account all skeletal characters, but his subgenera have not found general acceptance: indeed, current synonyms often span the groupings. Brook unfortunately did not examine Dana’s specimens, although he did see most of the European located types, and most Dana species can be shown to have a Brook synonym. Verrill (1902) also split many Dana species needlessly. Nemenzo (1967) revived the use of group names, although not affording subgeneric status to his groups. Of the twentieth century authors, Vaughan (1918), Hoffmeister (1925), Crossland (1928. 1952) and Wells (1954) have treated the genus in detail, with ecological data, although not concentrating exclusively on Acropora. Detailed underwater studies of Acropora were first made by J. Verwey in the Bay of Batavia in the 1930’s, and the taxonomic results of his study are still unpublished (see Umbgrove 1939, p.56, 1940, p.303; Wells 1954, p.4I4). In discussions with Dr Verwey I have found that we are usually in agreement in our general assessment of species limits (though obviously not always on names). His opinions pre-date those of this paper by some forty years, and it is hoped that his work, which is to be a complete monographic treatment of the genus, will soon be available. Characters and Terminology (For coral terminology not specifically referrable to Acropora see Moore, Hill, and Wells 1956). Acropora lacks most of the skeletal characteris- tics used in the species identification and study of variability of other corals, e.g. columella, dissepiments, septal structures. The main character defining the genus is the method of branching: an axial polyp builds a corallite of increasing length, and buds off radial polyps from ' its growing tip. Any radial polyp has the potential : to take up an axial role. A variety of branching patterns occur, and these (with commonly applied terminology) are summarized in Figure 1. The polyps have twelve tentacles, one of which (overlying a directive septum) is longer than the others. Two cycles of simple trabecular septa can be expressed in the corallites, although further cycles may be apparent in the walls. The walls, regarded as synapticulothecac (Wells 1956) have recently been shown to have septal contribution (Ricart y Menendez and Freidman 1977). See plate 43 and Figure 1 for a summary of these features). The axial corallites are usually described by measurements, (in this paper expressed as inner and outer diameter) and by septal development (here expressed as maximum observed develop- ment, to an approximate fraction of the radius, e.g. to 1/3R). The radial corallites are described according to their shapes, summarized in Figure 2 . ‘Coenosteum’ as described for Acropora commonly refers to both external corallite and intercoraliite features. The walls of corallites are often clearly costate, in which case the intercoraliite material is usually a spongy reticulum, with or without spines. Sometimes there is little or no difference between the appearance of the two areas. This is usually when the coenosteum is a dense arrangement of elaborated spines. With electron microscopy i different categories of spines can be seen and the ' coenosteum can be more accurately described j (Wallace and Grimmer, in preparation). j i Variability All Scleractina are capable of broad mor- phological variety, but in Acropora this can be caespitose caespito-corymbose table plate Fig. 1: Common colony shapes in Great Barrier Reef Acropora and their nomenclature. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 275 particularly subtle. A number of features contribute: the axial method of growth allows a continuum of shape possibilities, which may be exploited differently by the species, the popula- tion, the colony, or even parts of the colony. The simplified skeletal structure allows the application of only a limited descriptive terminology, and description of small variations is always difficult. Detailed measurements for analysis of variation often show similar variability within the colony to that in the entire population (Wallace, unpub- lished, for A. millepora and A. aspera; similarly high variability in Porites is shown by Brakel 1977). In fact the absolute variation in size of most skeletal features within the entire genus is relatively small for a coral group. In this much speciated genus there is a tendency for numbers of species to co-occur. Often differences between species can be seen in population strategies and slight differences in habitat preference. Despite the above, some of the taxonomic confusion so repeatedly quoted in the literature disappears when the taxonomist enters the water. This is particularly so with reef slope species where growth form and other features can often be seen to alter gradually with depth. It can be staled unequivocally that the most complex species laxonomically are those occurring on the shallow reef flat, and none of these can be considered to be fully described in the present paper. Some generalizations can be made, and should be born in mind when interpreting the taxonomic section. (1) Some reef regions support predominantly characteristic colony shapes, which can be attributed to physical parameters (in particular low-profile corymbose shapes on the outer reef flat where exposure is great (low water depth); small flat plates on the deep reef slope where light availability is limited). In these areas the species composition can be expected to include (a) species capable of only that shape; (b) species having the shape within a range of phenotypic variability; (c) morphs of polymorphic species. (2) All species which extend down the reef slope exhibit a gradual flattening out of shape with depth, and the ratio of radial to axial corallite numbers decreases. These flat colonies can be recognised for what they are if the observer investigates shallower sections of the populations. (In skeletal collections they are more difficult to sort). A few species are characteristically flat but with high radial to axial corallite ratio, and do not extend into deep water. At least one species {A. granulosa) is apparently adapted to low-light situations, and has a characteristic distribution which includes shallow but shaded situations. tubular, round opening tubular, oblique opening tubular tubular, tubular, nariform appressed dimidiate opening opening (tubo-nariformJ tubo-nariform dimidiate with rostrate development (front) lipped (labellate) flaring lip cochleariform sub-immersed immersed Fig. 2; Radial corallite shapes in Great Barrier Reef Acropora and their nomenclature. 276 MEMOIRS OF THE QUEENSLAND MUSEUM (3) Some basically arborescent species which occur on the reef flat may exhibit diverse growth forms adapted to different habitats. (4) The greatest variety of colony shape seems to occur in areas with good water cover and reasonable water circulation, such as deep ‘middle reef flat' areas and leeward patch reefs. Many species can be thought of as achieving their most ‘characteristic’ shape in these areas. Here also many colonies achieve colossal size. Many species have characteristic colouration, at least locally, which can be used as a guide to identification but it should never be supposed that this is the only possible colour for the species. Methods Field Studies: The observations in this paper are the result of a variety of studies on reefs in the Centra! and Southern Great Barrier Reef Province (sec Maxwell 1968). They concentrate on reef slope and other below-L.W.M. areas, as the corals here have previously been least observed. On Big Broadhurst Reef (18°55’S; 147o44^E) observations and collections were made serially across the outer reef flat and down the reef slope (see Wallace 1975; Wallace and Dale 1977). Similar techniques were used on the reef Oat at Redbill Island {20^57’S; 150"05’E) (Wallace, unpublished). Collections with habitat data were made at Bushy-Redbill Reef (see Wallace and Lovell 1977), Darlcy Reef (19'^24’S; 148°05’E), Bowden Reef (19^02'S; 147^06’E), Prawn Reef (19“02’S; 148°05’E), Viper Reef (18°53’S; 148°10’E), Tryon Reef (23°15’S; 151^47’E), Lady Musgrave Reef (23°55’S;' 152°24’E), and Heron Island reef (23°27’S; 151°55’E). Detailed samples of populations of A, miilepora and A. aspera for a morphometric study were made at Heron Island. A reference collection was made in the Fiji Islands, type locality of many of the species. Occasional reference is also made to observations by the author at Enewetak, Marshall Islands and Tongatabu, Tongan Islands. A recent opportunity to study specimens at James Cook University (JCU) collected at Lizard Island (14^40’S; 145''28’E) and the Palm Island group (18‘"40'S; 146°33’E) allowed further notes to be added for some species. Other workers contributed specimens from various parts of the Great Barrier Reef Province and the Murray Islands. Information on this material is given to the text, and locations of reefs can be seen in Maxwell (1968, figs. 17A-D). Other material examined in the Queensland Museum included the collections of the Great Barrier Reef Committee (G.B.R.C.) and specimens mentioned by Stephenson and Wells (1956). Type Material: Types were studied in the Smithsonian Institution (USNM), Yale Peabody Museum (YPM), British Museum of Natural History (BM), Mu.seurn National d’Histoire Naturellc, Paris (PM), University of the Philippines (UP). Fragments of type material were received on loan from the first three institutions, and photographs of type specimens were received from the Museum fiir Naturkunde, Berlin (MNB), the USNM and the BM. Collections were examined at the Rijksmuseum van Naturlijke Historie, Leiden (Verwey collec- tions), Zoological Museum Amsterdam, and the University of the South Pacific, Fiji. Laboratory Studies: Most specimens were stored as cleaned skeletons (preferably macerated and cleaned in water). Small fragments of some specimens have been preserved entire in buffered formalin. Observations on skeletons were made with a Wild M5 binocular microscope with linear measuring eyepiece. W'here possible, particularly with type material, five observations per character were made for each specimen, but expediency j prevented this procedure being used for all ( specimens. Additionally, representative specimens of each species were examined with scanning electron microscopy, and a standardized procedure for photographing axial and radial corailites and inter-corallite spines was followed. This information is sometimes used to clarify descriptions, but it is mainly to be used for a study further defining species groups and pursuing a phylogenetic grouping of them (Wallace and Grimmer, in preparation). Use of the Text Any taxonomic text has two main categories of readers: those interested in the taxonomic interpretation as such, and those requiring an identification tool. For this genus, I suspect the second category will be in the majority. I recommend that no attempt be made to identify single specimens without field observations, and conversely that field work (even when done without taxonomic emphasis) include casual assessment of affinities of colonies. Any field notes which allow specimens to be later grouped as series are useful. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 277 For observing skeletal specimens with an incident light microscope a finger held between the light source and the specimen (to diffuse the light) enables corallite shape and septal features to be viewed clearly. The photographic plates are considered to form a major part of the taxonomic description, because they (1) display field appearance of colony and habitat, (2) indicate size and shape of corallites, and where possible coenosteai texture, (3) indicate variety in all features. A fourth and most important role intended for the plates is that of a visual key. The identification process will be simplified if the user scans the plates before attempting to identify material. Reference to plate numbers is made once only for each description. Where synonymy lists of other authors have been accepted without investigation these are quoted as (synonymy) after the author reference. Check-lists and catalogues (e.g. Ralhbun 1887) are not used for synonymy lists. In this context Faustino (1927), being mainly a reiteration of previous descriptions, is regarded as a check list unless it makes an addition to information on a species. The arrangement of species is partly in accordance with previous arrangements, and reflects my own views of affinities. Strongly supported affinities are grouped as species-groups. As both detailed morphological and phylogenetic affinities of species will be pursued in a later paper this arrangement must be considered a working guide only. One species group, that including A. palifera (Lamarck) (i.e. the subgenus Isopora of Brook, 1893), has been omitted as this is under scrutiny in population studies by Potts (1976, 1977) and will be given separate taxonomic treatment (Potts and Wallace, in preparation). Specimens quoted in the text are registered specimens, and in most cases further unregistered material and spirit specimens are located in the Queensland Museum (QM). Smalt reference collections will be lodged in the USNM, BM, and MNB. Depth ranges recorded for species apply to the areas studied, and these may differ in other areas. Finally, some of the taxonomic format used by other authors has been omitted. For example, many authors describe undersurface details of colonies. As these depend on general colony shape and are reductions of upper surface features their usefulness is questionable. Radial corallite measurements are not given, as a statistically acceptable assessment of these which is compar- able across different shapes has not been arrived at (the illustrations and previous authors’ descriptions give an indication of size range). SYSTEMATICS Family ACROPORIDAE Verrill, 1901 Acroporidae Verrill, 1901, p.I63; Wells, 1936, p.99; Wells, 1956. P.F372. Massive or ramose colonies by extralentacular budding; hermatypic. Corallites small, synap- ticulolhecale or septothecate, pseudocostate or costate, slightly differentiated from coenostcum. Septa non-exsert, in 2 cycles, formed by simple spiniform trabeculae projecting inw'ard and upward from vertical mural trabeculae, commonly fusing to form laminae. Columella absent or trabecular and weak. Dissepiments thin and tabular when developed. Coenostcum extensive, light reticulate, flaky, generally spinose or striate on surface. (Modified from Wells 1956, P.F372). Genus Acropora Oken, 1815 Acropora Oken, 1815, p.66; Verrill, 1901, p.l64; Verrill 1902, p.208 (synonymy); Vaughan, 1918, p.l59; Wells, 1956, p.F373 (synonymy); Nemenzo, 1967, p.47. Typi Spi-CII S: Millepora muricata Linnaeus, 1758. Acroporidae which are ramose, rarely massive or encrusting; branching with an axial or leading corallite larger than the more numerous radial corallites budded from it; united by light, reticulate, spinose or pscudocostate or costale coenostcum. Columella and dissepiments absent. (Modified from Wells 1956, p.F374). The ‘Acropora robusta' GROUP The four species Acropora palmerae. A. robusta. A. rotumana, and A. abrotanoides are very similar in corallite morphology but not colony shape. A. palmerae, described as a completely encrusting species (Wells 1954) can bear short vertical branches (Wallace, in prep.). A. rotumana is stalked, with thick alciform branching units or even forming a sturdy arborescent table. A. robusta can vary phenoty- pically from encrusting to open branching. A, abrotanoides, still poorly categorized, occurs as clumps of mainly vertical branches. 278 MEMOIRS OF THE QUEENSLAND MUSEUM Ignoring this last species, the other species have strong habitat preferences: A. palmerae for rocky (typically algal ridge) locations; A. rotumana for deeper-water reef flat areas, A. robusta for the shallow outer reef. On Enewetak atoll in the Marshall islands A. palmerae and A, rotumana arc very abundant and A. robusta rare {Wallace, in prep.). On the Central Great Barrier Reef where there is no distinct algal ridge, A. palmerae is absent, any encrusting colonies being recognis- able as members of an A. robusta population: A. rotumana appears as a rare species. Within each species there is also a phenotypic component to radial corallitc variation: the radials are best developed on the most freely branching parts of the colony. These features, and the added fact that the clumsy colonics never lend themselves to the collection of a truly representative sample, have led to the naming of many species from single or few fragments. As well as the species placed in synonymy here, a number of more obscure species-names possibly belong with these species, e.g. A. cyclopea (Dana), A. cuspidata (Dana), A. conigera (Dana), A. smithi (Brook). With the current lack of geographic studies on Acropora it is difficult and unwise to lump these. The separation of the three species treated here is maintained on the basis of their apparent distinctness in Great Barrier Reef populations, and pending further regional studies. Acropora robusta (Dana, 1846) (Plates 44, 45) Madrepora robusta Dana. 1846, p.475, pi. 39, fig.3, 3a, pi. 31, figs. 3a-c: Brook, 1893, p.42 (synonymy). Madrepora pacifica Brook, 1891, p.465; 1893, p.39, pi. 30, fig.B. Acropora pacifica: Crossland, 1952. p.202, pl.31, fig. 2, pi. 32, fig. 2; Nemenzo, 1967, p.54, pM8, fig.4. Madrepora decipiens Brook, 1892, p.456; 1893, p.51, pi. 14, figs. B to D. Acropora decipiens: Vaughan, 1918, p.l65, pi. 67, figs. 2, 2a, 2b; Nemenzo, 1967, p.60. Material Examined USNM: Fiji Islands, U.S. Expl. Exped., A. robusta holotype 297. BM: Samoa, Rev. Whitmore, A. pacifica holotype 1875.10.2.13.’ Rocky Is., G.B.R. Saville Kent A. decipiens syntypes 1892.6.8.82; 1892.6.8.85. QM: Big Broadhurst Reef: SW. side, reef slope: 4m, 27.111.1973, C.W., G10190; 2-2m, Il.x.1973, C.W., G10250; l-5m, 18.x. 1973, C.W., G10191; Im, 26.111.1973, C.W,, G10194; SW. side. Outer reef flat: Il.x.1973, C.W.. G10246. 18.x. 1973, C.W., G10248; 22.X.1973. C.W., G10247. Bushy-RedbilLReef; SE. side, reef crest, 27.xii.1972, C.W., G10193; S side, reef crest; 27.xii.1972, E. Lovell. G10I92; 25.xii.l972, E. Lovell, GI0257; W. side, reef crest, 22.xii.1972, C.W'., G 10258; W. side, reef slope, I9.xii.l972, E. Lovell, G10259; NW. side, reef crest, 30.xii.l972, C.W., G10I93; NNE side, reef slope, 30.xii.l972, C.W., G10193; NNE side, reef slope, 30.xii.l972, C.W., G10197. Bowden Reef:' SW. side, reef crest, 15.vii.l972, C.W., G 101 98, G 10199; slope of opening in SW. side, 26.vii.1972, C.W., GI0256. Bramble Cay: Dec. 1924, C. Hedley, G 10201; Oct. 1924, C. Hedley, G 1 0262. Coates Reef. 13.viii.l924. ‘Geranium’, G10261. Feather Reef, ILviii.1924, ‘Geranium’, G10260. Flinders Reef, Moreton Bay. W. side, 6.ix.l973, C.W., G 10200. Masthead Reef, NNE. side, reef crest, August 1974, J. Buhman, Gi0254, G10255. Field Diagnosis Colonies have stout branches which can be vertical cones, distorted humps, or freely branching horizontal units on different parts of the same colony. Sections of the colony can be completely encrusting. Those parts of the colony on solid substrate have the solid vertical cones, but these extend horizontally and branch freely where small holes in the reef, or the reef edge itself, allow. The low, distorted rounded humps are formed in very shallow (e.g. reef crest) areas. Prominent tubular radial corallites mixed with shorter ones are obvious at the tips of the cones and branches. Common colours are bright green with deep pink branch-tips and pink-brown, yellow-brown or cream. Laboratory Diagnosis Laboratory difficulties occur because usually only a piece of the heavy colony is collected. Two very different fragments are illustrated in Plate 45 figs, a and c. Crossland’s plate 32, fig.2, shows another shape the branches can take. Branching pattern: This is described above. The free branches are usually 20 to 35 mm diameter, but branchlets as narrow as 10 mm can occur; maximum length measured is 250 mm. The cones may be as thick as 40 mm at the base. The humps are either solid or encrusting, taking the shape of the substrate. Axial corallites: Outer diameter 2-5 to 3 0 mm; inner diameter L2 to 1-5 mm. Septation: primaries to 3/4R, secondaries to I /4R, usually incomplete. The axial corallites are not strongly different from large radial corallites, and on the rounded humps they may be absent altogether. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 279 Radial corallites: Tall and short radial corallites mixed. The differences between the two are emphasised at the growing tips where the tall radials may be 5 mm long. On the low humps the differences may disappear entirely. The tall radials are tubular, with rounded, oblique or dimidiate openings. Their angle of extension from the branch decreases from 90*^ at the base to about 45° at the tips. The short radials usually have only a small section of outer wall present. Septation is usually well-developed in the tall radials (primaries to 1 /2R, secondaries to 1 /3R), and less developed in the short radials (sometimes only the directives visible). Coenosteum: Costate on corallites, reticulate with simple spines in between. Within Reef Distribution The species occurs only in the shallow outer flat-reef crest area, and flourishes at the windward (SE.) end of the reefs. Identification Difficulties and History In the field this species is conspicuous and distinctive. In the laboratory the colony is poorly represented by fragments and there is the temptation to find a ‘best fit’ with one of the many described robust species, few of which have been described from the field. Only the most certain synonyms have been included; some of the robust species mentioned in the introduction may also be this species. The more slender free-growing branches resemble A. intermedia, and specimens of the two species can be confused in the laboratory (but never in the field). A. palmerae Wells may be synonymous. An encrusting colony of A. robusta with few vertical branchlets is illustrated in colour in Roughly 1936 (plate 6 upper). Geographic Distribution Indo-Pacific: China Sea, Philippines, Great Barrier Reef, Fiji Islands, Samoa, Marshall Islands (C.W.). Acropora rotumana (Gardiner, 1898) (Plate 46) Madrepora rotumana Gardiner, 1898, p.258, pi. 23, fig.2. Acropora rotumana: Hoffmeister, 1925, p.69; Wells, 1954, p.419, p.ll2, rig.1-3, pl.ll3, figs.4, 5. Acropora tutuilensis (part) Hoffmeister, 1925, p.7L Material Examined USNM: Pago Pago Harbor, Tutuila, Samoa A. tutuilensis no. 4 (mentioned Hoffmeister, 1925), Rongerik Atoll 44477; Rongerlap Atoll 44478; Bikini Atoll. 44469, 44470, 44472, 44473, 44474, Enewetak Atoll 44475: A. rotumana (mentioned Wells, 1954). QM: Big Broadhurst Reef: SW. side, outer reef flat: 14.X.1973, C.W., GI0233; 22.X.1973, C.W., G10234, G10235; E. side, reef slope, Im, 21.X.1973, C.W., G10236. Bowden Reef: SW. side, reef crest, 15.vii.l972, C.W., G10263. Field Diagnosis Mature colonies are about 1-5 m in height and width. They have a sturdy tree-like appearance due to the small number of very stout branches which have some proximal vertical growth component, then become horizontal for most of their extent, and proliferate into small branchlets at the tips. This is not the full range of shapes possible in the species. At Enewetak Atoll, for example, it can form a stalked table (Wallace, in preparation). Colours: deep pinkish-brown or apple green. (See Gardiner 1898 for detailed field diagnosis). Laboratory Diagnosis The fragments collected from this species are usually branch-tips with a greater or lesser amount of branch remaining. They often have a ‘distorted' appearance, due to the irregular secondary branching pattern. Branching pattern: described above. Axial corallites: Outer diameter 2-0 to 2-5 mm (in the G.B.R. specimens); inner diameter 0-8 to 1-2 mm. Septation: primaries present to 1/2R, few secondaries, up to 1 /4R. Radial corallites: Tall and short radial corallites mixed. Tall radials extend from branch at 45° to 90° and are tubular with round, oblique, dimidiate or nariform openings. Short radials usually have less than half the wall developed, and range down to immersed. Primary septa not well developed, up to 1/3R; secondaries absent or a few just visible. Coenosteum; Costate or pseudocostate on radials, reticulate with simple spines in between. Within Reef Distribution Occasional colonies found just below the reef top on the upper reef terrace or in surge channel openings on the outer reef. Elsewhere (e.g. on Enewetak Atoll) it occurs in deep-water reef flat situations. Identification Difficulties and History This is one of the few Acropora species to be originally described from the field as well as from 280 MEMOIRS OF THE QUEENSLAND MUSEUM skeletal material. Both Hoffmeister and Wells regard it as a good species because of its distinctive colony shape. Gardiner states it to be ‘by far the most abundant coral on the reefs of Rotuma’. It cannot be distinguished from A. robusta or A. abrotoftoides on radial corallite shape. Geographic Distribution Pacific Ocean: Great Barrier Reef, Rotuma, Samoa, Marshall Islands. Acropora abrotanoides (Lamarck, 1816) (Plate 47) Madrepora abrotanoides Lamarck, 1816, p.280; Brook, 1893, p.56 (synonymy). Acropora abrotanoides: Crossland, 1952, p.204; Wells, 1954, p.418, pl.l23, figs. 1, 2. non Acropora abrotanoides: Vaughan, 1918, p.I66, pi. 68, fig. 2. Madrepora deformis [non Michelinl; Dana, 1846, p.484, pL43, fig. 1. Madrepora danai Milne Edwards and Haime, 1860, p.560. Madrepora danae: Brook, 1893, p.57 (synonymy). Acropora danai: Wells, 1954, p.418, pi. Ill, figs. 4-6. Materiai. Examined USNM: Tahiti, U.S. Exp). Exped., A. danai holotype 303 (M. deformis of Dana); Marshall Islands, J. W. Wells. A. danai 45175 (mentioned Wells 1954). YPM; Tahiti, fragment of type (labelled A. deformis) 4162 (mentioned Verrill, 1864) QM: Big Broadhurst Reef, patch reef in lagoon, 28.iii.1973, C.W., G10238. Bushy-Rcdbill Reef, NW. side, reef crest, 22.xii.1972, C.W., G 10239. Bowden Reef, slope of opening in SW. side, 26.vii.1972, C. W., G10240-2. Viper Reef, patch reefs in lagoon, L5m., 16.vii.l972, C. W., G10243. Great Astrolabe Reefs, Fiji Islands, 1974, C. W., G 10244. Field Diagnosis Colonies occur as tufts of sturdy vertical to oblique branches with appearance roughened by the presence of numerous incipient branchlets. Colour: yellow-brown. LABORATORY DIAGNOSIS Branching pattern (taken from five specimens): From a basal area (usually dead) a small number of sturdy branching units is given off. Each branching unit has a small number of main branches which are vertical or almost so, and the branches may branch again. In addition, each branch is roughened by the presence of incipient branches. The longest branch is 150 mm, and the widest 40 mm. Axial corallites (on main branches): Outer diameter 2-5 to 3-8 mm; inner diameter 0-8 to 1-2 mm. Scptation: primaries present to 1/3R, secondaries sometimes complete to I/4R. Radial corallites: Tall and short radials mixed. Tall radials tubular with round, oblique or dimidiate openings. Those with round openings usually can be recognised as incipient axials, themselves budding radials from their circumference. Coenosteum: Costate on radials, reticulate with simple spines in between. Within Reef Distribution This species seems to be restricted to upper reef areas, mainly the outer reef flat and crest, and the tops of patch reefs in deep lagoons. Identification Difficulties and History Brook describes his specimens in detail, and apparently examined Lamarck’s type, which I have been unable to locate. AH other authors had meagre or immature specimens (e.g. Vaughan’s Three small immature specimens'). There are no morphological characters supporting the separa- tion of A. danai. A. listen (Brook) may also be a synonym, but its validity could be tested by a study on the Tongan reefs. This species still remains categorized only as a group of specimens, with no field identity. The fact that it eludes categorization in the field suggests that it is not a ‘good’ species. The abundance of incipient branches is a poor species character, but this is at present the main feature defining this species. Geographic Distribution Indo-Pacific: Singapore, Great Barrier Reef, Marshall Islands, Tahiti. Acropora intermedia (Brook, 1891) (Plates 48, 49) Madrepora intermedia Brook, 1891, p.463; 1893, p.3L pl.l, fig. C. Acropora intermedia: Crossland, 1952, p.200, pi. 32, fig. 1; Stephenson and Wells, 1956, p.l6; Pillai and Schcer, 1976, p.24, pi. 2, fig. 1. Material Examined BM: Maidive Islands, A. intermedia syntype 1886.11.22.6. WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF 281 QM: Big Broadhurst Reef, SW. side, reef slope: 2m, 11.X.1973. C. W., GIMOO; 2m, 26.iii.1973, C. W., G1 1295; 3m, 28.iii.1973, C.W., GI1310; 5m, 25.ii.1973, C.W., GI1302; 5.3m, 13.x. 1973, C.W., Gn576; 6m, 25.iii.1973, C. W., G1I301; 6 1m, 13.X.1973, C. W., Gn582; 6-3m, 13.X.1973. C. W., G1I579; 7-6m, 13.x. 1973, C. W., G1I575; 7-7m, I4.X.1973, C. W., Gl 1578, Gil 580; 8m, 26.iii.1973, C. W., Gl 1300; 8-3m, I5.X.1973, C. W., 011581; 8-6m, 15.X.1973, C. W., 011577; SW. side, outer reef flat: 1 Lx. 1973, C. W., G11308, G11583; 22.X.1973, C. W., G1 1327, G11584; SW. side, floor of surge channel opening: 20.x. 1973, C. W., G11307, GU309, GI 1585. Bowden Reef, SW. side, upper reef slope, 26.vii.1972, C. W., G11316. Bushy-Redbill Reef, S. side, reef slope, 3m, 27.xii.1972, C. W., G11311; adjacent Redbill Is., reef crest, 18.xii.l972, C. W., G11305; NW. side, first reef crest, 22.xii.1972, C. W., G11312, G11313; NNE. side, reef slope, Jan. 1973, C.W.. G11391. Heron Is., W. side, reef flat, 6.vii.l973, Y. Loya, G11297. Field Diagnosis Colonies vary from small clumps to small thickets of around 3m diameter and are sturdy-branched arborescent. The radial corallites are a mixture of tall and short tubular and are obvious under water. Colours are cream to pale brown, pale green or greenish brown or bright blue. Laboratory Diagnosis Branching pattern: Branches are given off at wide angles (45° to 90°) and branch width varies from 15 to 25 mm. Branches usually taper strongly. Axial corallites: From barely exert to 2 mm exert. Outer diameter 3 0 to 4-0 mm (in QM specimens); inner diameter 11 to 1-5 mm. Septation: primaries well developed, often reaching 3/4R, secondaries usually present, up to 1/4R. Radial corallites: Tall and short radial corallites are mixed. The tall radial corallites are tubular, up to 5 mm long, with round, oval or dimidiate openings and usually extend from the branches at 90° to 45°. Lower wall may be slightly thickened. Primary septa are developed up to 2/3R, and secondaries present, up to 1/4R. The smaller radials are tubular appressed or emergent, or sub-immersed, and oriented with their openings facing in random directions to the branch. Coenosteum: Strongly costate on radial corallites, open reticulate with simple spines between radials. Within Reef Distribution Middle and outer reef flat, upper reef slope to about 12 m, surge channel floors and sandy bottoms around patch reefs. Identification Difficulties and History This species appears to be morphologically similar in radial corallite and coenosleal features to the A. robusta group. Many specimens are difficult to place exactly in either this species, A. formosa or A. grandis, and clearly all three species require further field study. Geographic Distribution Indo-Pacific: Maldives, Great Barrier Reef. Acropora grandis (Brook, 1892) (Plate 50 A,B) Madrepora grandis Brook, 1892, p.457; 1893, p.42, pi. I, figs. A, B. Acropora grandis: Crossland, 1952, p.202, pl.31, figs. 1, 3, 4, Material Examined BM: Palm Island, Saville-Kent, A. grandis syntype 1892.6.8.60; Herring Island, Bowen, Saville-Kent. A. grandis 1892.6. 8. 314; Rocky Island, Saville-Kent, A. grandis var. 1892.6.8.261. QM; Big Broadhurst Reef, SW. side, reef slope: 6m, 25.iii.1973, C.W., 011320; 6-Im, I3.X.I973, C.W., GII366; 7m, 27.iii.1973, C.W., G11299; 7-lm, 13.x. 1973, C.W., GI1367; 7-7m, 14.x. 1973, 011368. Bushy-Redbill Reef, NW, side, first reef crest, 22.xii.1972, C.W., 011306, G1I315, G11317. Darley Reef, patch reef in lagoon, 4 m, 24.iii.1973, C.W., 011298. Field Diagnosis Openly arborescent colonies have scattered radial corallites of mixed size with large obvious openings, and may be brown, bright blue, bright purple or even greenish-brown with blue tips. Laboratory Diagnosis Branching pattern: Branches commonly spread at from 60° to 40° . Branches are from 10 to 30 mm thick, and may be as long as 30 cm without branching. They usually taper gradually. Axial corallites: Up to 3 mm exert; outer diameter 2-5 to 3-5 mm; inner diameter 1-0 to 1-7 mm. Septation: both cycles may be present, the primary septa up to 1/3R, the secondary up to 1/4R. Radial corallites: Two sizes of radial corallite occur. Towards the branch tips the large radials reach 3 mm length, but elsewhere they are much 282 MEMOIRS OFTHE QUEENSLAND MUSEUM shorter than this. The openings are round to oval, and are directed straight out from the branch, or nearly so. The septa are poorly developed: both cycles may be completely absent, or primary sepia may be present up to 1/4R, plus a few secondaries. Coenostcum: The surface has a light, crumbly appearance, the coenostcum being costate or reticulate on the radial corallites and open reticulate between. Identification Difficulties and History This species has only been recorded from the Great Barrier Reef, and in my experience it is not common on the outer reefs, but is more characteristic of the fringing reefs of the continental islands. Crossland treats the species in detail. Although this species is morphologically different from A. formosa and A. intermedia in many features, there remains a suspicion that it may be related to one or other of these, and all three species require further study in the field. Geographic Distribution Recorded from the Great Barrier Reef only. Acropora formosa (Dana, 1846) (Plates 51, 52) Madrepora formosa Dana, 1846, p.473, pl.3I, fig. 2a, 2b, pi. 38, rig.4. Acropora formosa: Hoffmeister. 1925, p.55. pi. 8, figs. I -3 (Synonymy); Wells, 1950, p.35; 1954. p.4I5, pi. 102, figs.1-9. pi. 103. figs.I-5, pi. 104, fig.4 (synonymy); Stephenson and Wells, 1956, p.14 (synonymy); Ncmenzo, 1967. p.6l, pl.21, fig.3; Pillai and Schecr, 1974, p.453; Pillai and Scheer, 1976, p.23. Madrepora hrachiata Dana. 1846, p.474, pi. 38, fig.3, 3a, 3b. Madrepora gracilis Dana, 1846, p.482, pi. 41. fig 3 3a, 3b. Madrepora nohilis Dana, 1846. p.48l, pi. 40, fig.3, 3a. Acropora nohilis: Hoffmeister, 1925, p.59, pl.l I, fig.l (synonymy); Wells, 1954, p.416. pi. 104, figs.l, 2; Nemenzo, 1967, p.62, pl.21, fig. 5. non Acropora nohilis: Pillai and Scheer. 1974, p.453. figJc. Acropora laevis (part) Crossland, 1952. p.230, pi. 45, figs.l, 2. M \ 1 1 Ki \i E\ \\ii\i I) USNM: Fiji Islands. U.S. Expl. Exped., A. formosa syntypes 888, 282; Fiji Islands, U.S. Expl. Exped., A. gracilis holotype 333; Sooloo Sea, U.S. Expl. Exped., A. hrachiata holotype 295; Singapore, U.S. Expl. Exped., A. nohilis holotype 427. QM: Big Broadhurst Reef: SW. side, reef slope: 1-6 m, Oct. 1973. C.W.. GI1388: 2 m, 20.x. 1973, C.W.. G1I339; 2 m, II.x.1973, C.W., GI1343; 2 m! 26.iii.l973, C.W., GI1398, G11405; 2-3 m, 20 x 1973 C.W., GI1349, GII353, G11360; 2-6 m. 22.x.l973i C.W., G1I346; 2-8 m, I2.X.1973, C.W., G11356; 3-8 m. 12.x. 1973, CAV.. GII335, GII347: GII362;’ 5- 3 m, 13.X.1973, C.W.. GI1365; 5-7 m. 23.x. 1973, C.W.. GII348, GII364; 6 m, 25.iii.I973, C.W„ GII320; 6 m, 13.X.I973, C.W., G11336; 6- 1 m, 13. X. 1973, C.W.. GI1363, G1I373, 6-3 m,’ 20.x. 1973. C.W., GI1334: 6-3 m. 23.x. 1973, C.W., GI1361; 6-3 m, 13.x. 1973, C.W., G11369, G11371; 7- 6 m, I3.X.1973, C.W., GII367; 7-7 m, 14x1973 C.W.. GI1333, G11341, GI]368, G1I372: 7-8 m’ I4.X.1973, C.W., GII330. GI1337, Gl]358, Gn37o! 8- 1 m, 22.x. 1973, C.W., GII340; 8-7 m. I5.X.1973, C.W., G11352; 12-4 m, 23.X.1973. C.W., GII363; 12-5 rn, 17. x. 1973, C.W., Gl 1350, G1 1355; 28.iii.l973i C.W,, G 1 1396; SW side, outer reef flat: 20.x. 1973, C.W., GI1331, G1I354; ll.x.1973, C.W., G1I359; 1 8.x. 1973, C.W., Gl 1344; SW. side, reef crest: 1-3 m, 22. x. 1973, C.W., GII345; 1-3 m, 23.x. 1973, C.W.] G11351; SW. .side, surge channel, 7-5 m 20x 1973 C.W., GI1342. Bowden Reef: SW. side, upper reef slope: 26.vii.1972, C.W., GII318. GI13I9. Gl]322, GII323, GI1325; floor of opening in SW. side, 5 m, 26.vii 1972 C w' GI1380, G1138I, 011375. Bushy-Redbill Reef: NW. side, first reef crest: 23. xii.t972, E. Lovell, Gl 1374, G11376, 01 1377; 22.xii.l972. C.W., GII306, G1I378, G1I379, G1I384 G11385. Darley Reef, patch reef in lagoon: 1 m, 24.iii.1973, C.W., GII401; 4 m, 22.iii.1973, C.W., GI1404. Viper Reef, .luly 1972, C.W., GI1326, G11394. Field Diagnosis This is the common, thicket-forming arbores- cent species with numerous small tubular radial corallites. commonly coloured cream to brown, less often bright blue or blue-tipped. Laboratory Diagnosis Branching pattern: Branching is always openly arborescent. Branch diameters vary from slender (10 mm) to thick (35 mm). Axial corallites: From barely exert to 1-5 mm exert. Outer diameter 2-0 to 2-8 mm; inner diameter 0-8 to 1-2 mm. Septation: Primary septa well developed, up to 1/2R; secondary septa usually present, but poorly developed, up to 1/4R. Radial corallites: Tubular, varying from sub-immersed to 5 mm, prominent, with openings from round to sharply oblique, the corallites oriented at anything from 90° to the branch to fully appressed. Septal development varies greatly, but most commonly secondary septa are absent or only poorly represented. WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF 283 Coenostcum: Coslate or neatly arranged simple spines on radial corallites, reticulate with simple spines between. Within Reef Distribution Reef crest and slope to position of surge channel floor (10-12 m), lagoonal floors and lagoonal patch reefs (may even form entire basis of lagoonal patch reefs), middle reef flat, occasional small colonies on outer reef flat, fringing reefs. Identification Difficulties And History The interpretation that I have placed on this species follows that of Wells (1954) and Stephenson and Wells (1956) preceded by Hoffmeislcr ( 1 925). The reader is referred to their discussions. The species, as interpreted, may well be a mixture and there is certainly no satisfactory interpretation of variability (for example slender and sturdy branched thickets may occur adjacent to each other). There is some suggestion (J. Collins, pers. comm.) that the sturdy branched colonies may be old colonies. The colonies offer no readily perceplable clues in the field. If there are several species involved, the mosaic of their variabilities may be adding to the confusion. This species requires individual attention, and the only assistance that can be offered at present is in distinguishing from the field the other ‘easily recognizable’ arborescent species; A. haimei, A. microphthalma, A. intermedia, A. aspera, A. pulchra and A, splendida. A. acuminata Verrill does not seem to occur in the areas studied. Geographic Distribution Indo-Pacific: Ceylon, Cocos-Keeling, Amboina, New Ireland, Great Barrier Reef and Torres Straits, Fiji Islands, Samoa, Marshall Islands, Tahiti. (Further Indian Ocean localities in Pillai and Scheer, 1974). Acropora splendida Nemenzo, 1967 (Plates 53, 54) Acropora splendida Nemenzo, 1967, p.51, pi. 17, fig-2. Matkriai Examiniuj UP: Hundred Islands, Pangasiman, 1959, A. de la Cruz, holotype C931. USP; Fiji Reefs 3916. QM: Big Broadhursl Reef, SW. slope: 23-5 m, 17.x. 1973, C.W., G8715; 15 m, 16.x. 1973, C.W., G8716; 9-7 m, 15.x. 1973, C.W., G8714; 8 m, 28.iii.1973, C.W., G8704-6; 7-H m, 15.x. 1973, C.W., G87I3; 7-5 m, 15.x. 1973, C.W., G87I2; 7 m, 26.iii.l973. C.W., G870], G8702; 6-9 m, 14.x. 1973, C.W., G8710, 08711; 6-6 m, 14.x. 1973, C.W., G8708, G8709; 6 m, 25.iii.1973, C.W. G8699; 5 m. 26.iii.l973. C.W., G8700; 4-9 m, 13.X.I973, C.W., G8707; 3 m, 27.iii.1973. C.W., G8703. Bowden Reef: Lagoon slope, Nov. 1972, R. Pearson, G8696; floor of opening in SW. end of reef, 26.vii.1972, C.W., G8697. Darley Reef, patch reefs in lagoon, 4 m, C.W., G8698. Field Diagnosis Colonics arc large (commonly around Im diameter) bowl to bracket shaped, the shape given by long openly arborescent branches which curve and divide to varying extent, depending on their position within the colony. The bowl shape occurs in specimens on flat surfaces and attachment becomes lateral with steepness of the attaching surface. With increasing depth the colonies become flatter and smaller. A specimen collected at 23-5 m (G8715) is completely horizontal. Common colours are dark olive green with paler green tips to the branches, dark blue with paler blue tips, brown with pale blue lips, brown or cream. Laboratory Diagnosis Branching pattern; As the colonies arc large, fragments are usually collected. These are often easily confused with other large arborescent species. Specimens from the centre of the bracket usually have long curving branches with little secondary branching; those from the edge zone arc more proliferous, with shorter branches and some fusions, and may be flat underneath. Axial corallites: Outer diameter 2*0 to 3*5 mm; inner diameter 0-8 to 1-5 mm. Septation: primaries to 2/3R, all or some secondaries present to 1 /2R. Radial corallites: Tubular, projecting from branches at 90'" (less towards tips) with openings round, oval, nariform, or dimidiate. Radials are usually evenly sized and distributed but smaller radials may be present amongst these, and their openings may be variously directed. Septal development varies, but primaries arc usually present to 1/3R, and some secondaries visible. Coenostcum: Usually neatly costate on radial corallites, open reticulate with simple or laterally flattened spines in between. Within Reef Distribution Reef slope from a few metres below crest to limits of Acropora distribution (species no. 3 in Wallace 1975); deep water lagoonal patch reefs; fringing reefs. 284 MEMOIRS OF THE QUEENSLAND MUSEUM Identification Difficulties and History Nemcnzo’s species was based on a single fragment, and he was not aware of the shape of the colony. The species seems to have been neglected in all other literature, although it is possible that fragments may have been identified with other arborescent species. This is the only Acropora with an ‘arborescent bracket’ shape and it is very distinctive in the field. Laboratory specimens without field notes become unnecessari- ly confused with other arborescent ‘problems’. Geographic Distribution Indo-Pacific; Philippines, Great Barrier Reef, Fiji Islands, (common in Fijian fringing reefs, from my personal observations), Palau (M. Pichon, pers. comm.). Acropora horrida (Dana, 1846) (Plates 55, 56) Madrepora horrida Dana, 1846, p.472, pi. 39, fig.2, 2a; Verrili, 1864, p.4I; Brook, 1893, p.l88 (synonymy). Acropora horrida: Wells, 1954, p.417, pi. 107, fig.l. Madrepora angulaia Quelch, 1886, p.l60; Brook 1893,p.l95. Madrepora inermis Brook, 1891, p.462; 1893, p.194, pi. 29, figs. A, B. Acropora inermis: Wells, 1954, p.43T MaTI'RIAI EXAMlNtD USNM; Fiji Islands, U.S. Expl. Exped., A. horrida holotype 291. YPM; Fiji Islands, U.S. Expl. Exped., A. horrida fragment of type 2013. BM; Zamboangana. ‘Challenger' Exped., A. angulata holotype 1886.12.9.233; South Seas, purchased, A. rVif'/'w/.y syntypes 1841.12.11.6, 1841.12.11.7. QM: Big Broadhurst Reef: SW. side, reef slope: 7-3 m, I5.X.I973, C.W., G8755; 7-6 m, 23.X.I973, C.W., G8757, G8758; 5 m, 27.iii.1973, C.W., G8759: 8 m, 28.iii.1973, C.W., G8761; 4 m, 28.iii.l973, C.W., G8763; SW. .side, surge channel, 8-5 m, 20.x. 1973, C.W.. G8756. Bowden Reef: Slope of opening in SW_ side: 26.vii.1972, C.W., G8764-7; 1-5 m, 26.vii.1972, C.W., G8768; 1-2 m, 26.vii.I972, C.W., G8769; 25.vii.1972, C.W., G8771; floor of opening in SW. side, 26.vii.I972, C.W., G8770. Bushy-Redbill Reef: NW. side, floor outside slope: 12 m, 30.xii.I972, C.W., G9076; 8 m, 3I.xii.l972, C.W., G9077; NW. side, middle reef flat, 22.xii.I972, C.W., G9078; 20.xii.l972, C.W., G9079; NW. side, reef patches, 12 m, 14. vi. 1975, C.W., G9080; W. side, reef slope: 8 m, 3.vi.l975, C.W., G9081; Dec. 1972, C.W., G8774; adjacent Redbill Is., reef slope, 5 m, 2.vi.l975 C.W., G9084. Darley Reef, patch reef in lagoon: 22.iii.l973, C.W., G8773; 6 m, 22.iii.1973, C.W., G8762, 23.iii.1973, C.W., G8760. Heron Island, W. side, reef slope, July 1973, CW G8773. Fiji Islands, Great Astrolabe Reefs (Kadavu), W.side of Yaukuvi Levu fringing reef, l.ii.l974, C.W., G9085. Field Diagnosis Occurs as sprawling arborescent to shrubby patches. Branches are slender and have a ragged appearance due to the scattered distribution of the radials and their poorly-formed walls. The polyps of this coral are usually extended, an unusual feature in Acropora. The colour is usually light powder-blue to light grey. Laboratory Diagnosis Branching pattern: Branching is primarily open-arborescent. Specimens from loose substra- tum areas tend towards horizontal primary growth; those From firmer substrates are more upright. Main branches are 7 to I 5 mm in width, and taper gradually towards the tips. Secondary branchlets are irregularly sized and placed. They can be short, scattered twigs which alter the shape of the colony little, or bundles of branchlets which give a shrub-like appearance, or bottlebrush formations of subequal length arranged all over the branches. Axial corallites; Outer diameter 1-6 to 2-3 mm; inner diameter 0-6 to 1-2 mm. Septation: Primary septa present, up to 2/3R, secondary cycle poorly developed, usually a few septa just visible. Radial corallites: Scattered, sub-immersed to emergent tubular, sometimes appressed, with round openings. Walls are thin and very fragile around the opening. Primary septa developed, up to 1/3R, secondary septa usually absent, or one or two just visible. Cocnosteum: Open reticulate with simple to forked spines, both on corallites and between. Within Reef Distribution This species occurs where there is some constant water cover, down to at least 15 m, and apparently favours high sediment areas, i.e. deeper reef flat areas, reef slopes, rubble and sandy bottom areas around patch reefs and on surge channel fioors. It occurs with A. vaughani in the deeper areas. Identification Difficulties and History The species is distinctive in the field because of the extended polyps, ragged appearance and WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 285 unusual colouring. The extended polyps also give it a silky sheen. Laboratory differences from A. vaughani are explained by Wells 1954 (p.417). Brook’s A. inermis specimens arc apparently slender branches of this species. Quelch’s A. angulata is named from a single branch lip. Geographic Distribution Indo-Pacific: Philippines, Great Barrier Reef, Fiji Islands, Marshall Islands. Acropora vaughani (Wells 1954) (Plate 57) Acropora vaughani Wells, 1954, p.416, pi. 105, fig. 1, pi. 106, figs. 1-8, pi. 107, figs. 2-6. Material Examined USNM; Bikini lagoon, 18 fms, holotype 44452; Seaward slope, Bikini Atoll, 21-25 fms, paratype 44457. QM: Big Broadhurst Reef, SW, side, reef slope: 9-8m, 16.X.1973, C.W., G10264; 8Tm, 14.X.1973, C.W., G10265; 7m, 13.x. 1973, C.W., G10266; 6-lm, 13.X.1973, C.W., G10267, G10268, 3m, 27.iii.1973, C.W., G10269; 7m, 28.iii.l973, C.W., G10270; 8m, 28.iii.I973, C.W., G10271. Bushy-Redbill Reef, NNE. side, off-reef floor, 12m, 30.xii.l972, C.W., G10272. Field Diagnosis Occurs as scattered ‘shrubby arborescent’ patches, particularly on areas of loose bottom material (sand or rubble). Scattered short final branchlets are given off from the main branches. Colour is usually cream or pale brown. Laboratory Diagnosis Branching pattern: Branching is basically arborescent, but the length and positioning of secondary branches varies greatly. Branch widths are from 10 to 20 mm. Secondary branches (or ‘branchlets’) can be regularly arranged along the main branches, or scattered. In general, the slender main branches tend to be more proliferous than the sturdy ones. Axial corallites: Outer diameter: 1-6 to 2-5 mm; inner diameter: 0-6 to 0-9 rnm Septation: both cycles usually complete, the primaries up to 2/3R, secondaries up to 1 /4R. Radial corallites: Tubular with round or occasionally oval to nariform openings. They are unequal in length and orientation (usually oriented within a range 45° to 90°, except towards basal areas, where they may be appressed). Coenosteurn: Spines are distributed evenly over corallites and inter-corallite areas. In heavily calcified specimens these have elaborated tips; in less calcified specimens the tips are simple to laterally flattened. (For more detailed laboratory diagnosis see Wells, 1954). Within Reef Distribution From about 3m to 15m on reef slope, deep-water lagoons, and patch reef areas, on rubble or sandy floors, fringing reefs. Usually it occurs with A. horrida (Dana), from which it can easily be distinguished, on the Great Barrier Reef, because A. vaughani is more sturdy, does not usually extend the polyps in daylight, and is commonly cream or pale brown, while A. horrida is pale blue or grey. The upper range is more limited than for A. horrida, which can occur in some reef flat situations. Identification Difficulties and History The appearance of the branchlets can be very varied (see Wells), but with good field notes the species can be identified without confusion with other arborescent species. Geographic Distribution Pacific; Marshall Islands, Great Barrier Reef. Acropora pulchra (Brook, 1891) (Plates 58, 59, 60C) Madrepora pulchra Brook, 1891, p.468; 1893, p.44, pl.28, fig. A. Madrepora pulchra var. stricta Brook, 1893, p.44, pl.28, fig. B. Madrepora pulchra var. alveolata Brook, 1893, p.45, pl.28, fig. C. Acropora pulchra: Vaughan, 1918, p.l62, pi. 66, figs. 3, 3a; Crossland, 1952, p.203; Stephenson and Wells, 1956, p.l7. Acropora pulchra var. stricta: Crossland, 1952, p.204, pi. 34, fig. 2. Acropora pulchra var. alveolata: Vaughan, 1918, p.I62, pi. 66, figs. 1, 2; Crossiand, 1952, p.203. Material Examined BM: Keeling Is., A. pulchra holotype 1884.2.16.1. QM: Bushy-Rcdbill Reef, adjacent Redbill Is., reef flat: 7.vi.l975, C.W., Glil02, 011105; ll.vi.l975, C.W., GI1097. 011103, 011104; I2.vi.l975, C.W., G1I093, 011095, 011096, GlllOO, GlllOl. Heron Island, W. side, reef flat: 2.vii.l973, Y Loya, 01 II 17; 3.vii.l973, C.W\. 011120; July 1973, C.W., 011115; 011119, lLxii.1973, C.W., GllllI, 011116, 011121, 011122; 15.xii.l973, C.W., 011118. 286 MEMOIRS OF THE QUEENSLAND MUSEUM Fiji Islands. Great Astrolabe Reefs. Feb. 1974, C.W., Gil 123-5. Low Isles, August 1954, W. Stephenson (Stephenson and Wells 1956): G2617-20; G2699-G2712, G2715, G2716, G2721-5. North West Island, C. Hcdley, 3-4 June 1924, GB.R.C. 170. Field Diagnosis Colonies of this species are arborescently branching, and can vary in overall shape from open thickets to compact corymbose form. Small radial corallites with oblique apertures give the branch surface a smoother appearance than that of other arborescent or corymbose species. Colour is pale to dark brown, often with pale blue tips. Laboratory Diagnosis Branching pattern: The angle of emergence, length, and width of branches varies widely, as the plates show. Branch widths are from 7 to 15 mm, and the overall general colony form can be anything from an arborescent thicket to a neatly caespito-corymbose clump. Axial corallites; 1 to 2 mm exert; external diameter 2-0 to 3-5 mm, internal diameter 0*6 to 1-2 mm. Septation: primary septa well developed, up to 2/3R, secondary septa absent or poorly represented, occasionally all present, up to 1/4R. Radial corallites: Mixed sizes, all very small, project at 90^^ from branch, upper wall poorly developed, lower wall extended and lip-like. The opening is oval to dimidiate. Primary septa may be present to 2/3R, or may be just visible; secondary septa not usually visible, or a few may be present to 1 /4R. Coenosteum: Costate on radial corallites, openly reticulate to costate between. General appearance is light and spongy. Within Reef Distribution This is a reef flat species, occurring particularly on the inner and middle reef flat. It occurs with A. aspera, often in close association, as plate 60C shows. Identification Difficulties and History Brook (1893) described the radial corallities well and his descriptions should be consulted. The main problem with this species is seen when it occurs with A. aspera. The two species show similar colony shape variations, apparently in response to similar environmental conditions. Although the radial corallites are different in both shape and size, colonies sometimes appear to have a mixture of characteristics of both species. This problem was encountered on Redbill reef flat, at places where populations of the two species intermingled: some colonies suggest a hybrid appearance. It is possible that some of Vaughan’s A. pulchra specimens were A. aspera. Geographic Distribution Indo-Pacific: Cocos-Keeling Islands, Great Barrier Reef. Acropora aspera (Dana, 1846) (Plates 60, 61, 62) Madrepora aspera Dana, 1846, p.468, pi. 38, fig. 1, la, lb; Brook, 1893, p.62 (synonymy). Acropora aspera: Crossland, 1952, p.205, pi. 33, figs. 2, 3; Ncmenzo, 1967, p.65. Madrepora hebes Dana, 1846, p.468, pi. 35, fig. 5; Brook, 1893, p.l28 (synonymy). Acropora hebes: Vaughan, 1918, p.I74, pi. 73, figs. 2, 2a, pi. 74, figs. 1. 2, 2a, 2b, pi. 13, fig. 6; Hoffmeister, 1925, p.57. pi. 9, figs. 3a. 3b; Wells, 1950, p.36; Crossland, 1952, p.217; Wells, 1954, p.423, pi. 104, fig. 3; Stephenson and Wells, 1956, p.l4; Nemenzo, 1967, p.64, pi. 22. fig. 1. Madrepora cribripora Dana, 1846, p.470, pl.31, fig. 1, la-lc; Brook, 1893, p.l23. Madrepora manni Quelch, 1886, p.l50, pi. 9, fig. 1, la; Brook, 1893, p.63 (synonymy). Acropora ntanni: Fauslino, 1927, p.263, pi. 85, figs. 6, 7; Nemenzo. 1967, p.63. Acropora tuzonica Verrill, 1902, p.231, pi. 36c, fig. 4, pl.36F, fig. 9. ?Madrepora arabica Milne Edwards and Haime, 1860, p.l45; Brook, 1893, p.66 (synonymy). MaTIRIM E.XAMIM'D USNM; Fiji Islands. U.S. Expl. Exped., A. aspera holotypc 285; Fiji Islands, U.S. Expl. Exped., A. hebes syntypes 287, 286; Rewa River Mouth, Fiji Islands, U.S. Expl. Exped., A. cribripora holotypc 289. YPM; Manilla Bay, Luzon. Philippines A. luzonica holotypc 1809. PM: Mer Rouge, A. arabica holotypc 331. QM: Big Broadhurst Reef: S. side, surge channel on outer flat, 2 m, 21.x. 1973, C.W., GI0928; patch reef in lagoon, 28.iii.l973, C.W., G10929, GI0930; SW. side, reef slope, 18.X.1973, C.W., G10962. Bushy-Redbill reef: W. side, middle reef flat, 14.vi.l975, C.W., G 10947; N. side, inner flat-algal bank area, 6.vi.l975. C.W., G10937-41, G10943, G10960. G1096I; adjacent Redbill Is., reef fiat, 7.vi.l975, C.W., G 10944-6. Bowden Reef. SW. side, reef slope, 1 m, 15.vii.l972, C.W., GI0925. Heron Island; July 1973, C.W., GI0849-G10854; G 1091 3— G 1 091 7; G10921; W. side, inner reef flat. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 287 lLxii.1973, C.W., GI0860-2; W. side, reef flat: 15.xii.l973, C.W., G10886-92; 17.xii.l973, C.W., G10893, GI0896; N. side, reef Hat, ILxii.1973, C.W., G10855; N. side, reef flat (solid area), 14.xii.l973, C.W., G10883-5; N. side, inner reef flat, lLxii.1973, C.W., G10856. G10857; N. side, middle reef flat, lLxii.1973, C.W., G10858; S. side, outer reef flat, 12. xii.l973, C.W., G 10865-72; SW.side, outer reef Hat. 13. xii.l973, C.W., G10873-81; WSW. side, outer reef flat, 12,xii-1973, C.W., G 10864; SE. side, outer reef flat, 17.xii.l973, C.W.. GI0897-902. Low Isles. 1975, C. Limpus, G1093L Masthead Reef, reef flat, August 1974, J. Buhmann, G 10926. Michaelmas Cay, 1975, C. Limpus, G10932, G10933. Maer Island, Murray Island Group, N. side, outer reef flat, 17.vii.l974, G. Ingram, G10927. Bramble Cay, Oct. 1924, C. Hedley, G.B.R.C. 7, 93-7, 149-52, 168, 169, 21 1. Feather Reef, I Lviii.1924, ‘Geranium', G.B.R.C. 181. Ellison Reef, 25.vii.1924, Paradice, G.B.R.C. 2-6, 54-67, 176. Surprise and Flora Reefs, Sept. 1924, ‘Geranium’, G.B.R.C. 8, 9, 12, 13, 16, 19-21, 24, 28. Fiji Islands: Makaluvau Reef, 10. i. 1974, C.W., G10955, G 10957; Great Astrolabe Reefs, Feb. 1974, C.W.. G 10949-54. Field Diagnosis This is a basically arborescent species in which the colonies show considerable phenotypic flexibility. Different colonies or parts of a colony can have long, slender spreading branches with scattered radial corallites or shorter thicker branches, even to the extent of appearaing corymbose, with crowded corallites. Radial corallites are in two sizes and have poorly developed walls. Colours are commonly pale blue-grey, green-grey, or cream, less commonly bright blue. It is a characteristic of this species on the Great Barrier Reef that specimens, when collected, exude large quantities of mucous. Laboratory Diagnosis Branching pattern: Ranges from open branch- ing (at 90°) to closed branching, the branches being given off vertically to obliquely to give a corymbose appearance. Branch widths vary from 10 to 15 mm. Axial corallites: From non-exert to 3 mm exert. Outer diameter 3-5 to 4-5 mm; inner diameter 1-3 to 1-8 mm. Septation: Both septal cycles usually present, the primaries up to 2/3R, the secondaries up to 1/3R. Radial corallites: Two sizes of radial corallite are mixed. These are best regarded as large diameter’ and ‘small diameter’, although the ‘large diameter’ radials are also always more prominent. The ‘large diameter’ radials have round openings, upper wall undeveloped, and lower wall short and thickened, or extended horizontally as a lip, which may be rounded or pointed. The ‘small diameter’ radials are sub-immersed to immersed. In general colonies with short, thick branches tend to have the radial corallites crowded with short, thick lips, and colonies with long slender branches have radial corallites scattered, with thin pointed lips. On the larger radials primary septa are usually visible up to 1/3R; and secondary development varies, but often some septa are present, up to 1/4R. Coenosteum: Open reticulate with simple to laterally flattened spines between radials, costate on radials. Within Reef Distribution This is a reef flat species, occurring from the inner reef flat to outer flat, but not on elevated outer flat platforms. It can occur in very silty and coral-depauperate areas and shallow lagoonal situations. Identification Difficulties and History The two species ‘A. hebes' and A. aspera are both ‘well known’ and typical specimens of the two species appear very different from each other. However, the relationship of the two can be easily established in the field. The placing of less known synonyms is more difficult, and possibly there are other species names still to be included. Two major problems of identification occur with this species. The first is in separating some corymbose specimens from sturdy specimens of A. millepora, the second in separating some arborescent colonies from arborescent A. pulchra. (see discussions for these species). Geographic Distribution Indo-Pacific: ?Red Sea, Cocos-Keeling, Philip- pines, Great Barrier Reef, Fiji Islands, Samoa, Marshall Islands, Tongan Islands (C.W.). The ‘Acropora corymbosa’ Group A number of species have radial corallites with scale-like lips, and a disproportionately large number of species descriptions have been written to cover these species. In the following list there is barely a species which has not been placed in synonymy or compared with, or mistaken for some other species in the list, and of the ‘real’ species that exist, each has been placed under a number of the commonest 288 MEMOIRS OF THE QUEENSLAND MUSEUM names. The main species concerned are: A. corymhosa (Lamarck), A, efjlorescens (Dana), A. cytherea (Dana), A. spicifera (Dana), A. hyacinthus (Dana), A. surcidosa (Dana), A. millepora (Ehrcnberg), A. prostrafa (Dana), A. subulata (Dana), A. convexa (Dana), A. armata (Brook), A. reticulata (Brook). A, arcuata (Brook), A. rythereilo (Verrill). A conferta (Quelch), A. pectinata (Brook), A. recumhens (Brook), A. squamosa (Brook). The most widely applied name on this list is the oldest. A. corymhosa (Lamarck). Three con- tenders for this name are the species I am identifying as A. hyacinthus, A. cytherea and A. millepora. Older interpretations of this species tended to favour an A. cytherea-Wke appearance. A specimen regarded as the type (of A. corymhosa) was seen and described by Brook (1893) who points out the similarity between A. cytherea and this. Professor J. Wells has kindly shown me photographs of (apparently) this type, sent to him some time ago by the Paris Museum. Neither my own search of the Lamarck collection, nor a later search by Dr J. P. Chevalier could locate a type. My opinion is that the currently accepted view of this species (c.g. Wells 1954) is morphologically between A. millepora and A. hyacinthus. While it is possible that this is a true species, at least some of the specimens so identified are other species: for example Stephenson and Wells 1956 mention two specimens, of which one (G2626) is a small reef flat specimen of A. hyacinthus, and the other (G2623) a stunted A. millepora. The species has been mentioned by many other authors since 1893, e.g. von Marenzeller (1907), Vaughan (1918), Hoffmeister (1925, 1929), Thiel (1932), Crossland (1952), Ncmenzo (1967), Pillai and Scheer (1976). and is regarded as having a wide Indo-Pacific distribution. In the interests of taxonomic stability, I am not attempting to trace its synonyms on the basis of a regional study, but I feel a caution must be taken that the interpretation of this species by the various authors may not coincide. Hopefully, with extensive regional studies, this group will be given a more complete taxonomic treatment. The species which I interpret as A. hyacinthus and A. cytherea are dominant members of the reef-front assemblage. They arc the early colonizers and often influence the effective shape of some reef areas. Special problems are associated with the separation of these species at their overlapping limits. A. spicifera has not been present in my areas of study. A. millepora is a common reef-flat species which has usually (on the Great Barrier Reef) been identified as its synonym, A. squamosa. Acropora hyacinthus (Dana, 1846) (Plates 63, 64A-C, 65, 66A,B) Madrepora hyacinthus Dana. 1846, p.444, pi. 32, fig. 2; Brook, 1893, p.I07 (synonymy). Acropora hyacinthus: Th\e\. 1932. p.i23, pi. 1 6. fig. 2; 1933. p.20; Stephenson and Wells. 1956. p.16; Nemenzo. 1967. p.|l5, pi. 33, fig.I; Pillai and Scheer, 1976, p-29. Acropora hyacinthus (?part): Hoffmeister, 1925, p.64. pL]3, fig.3: Wells, 1954. p.42L ? Madrepora conferta Quelch, 1886. p.l64, pi. 10, fig.3; Brook, 1893, p.l08. Madrepora pectinata Brook, 1892, p.460* 1893 p 95 pl.27, fig.D, E. Madrepora recumhens Brook, 1892, p461- 1893 p.I06, pl.27. fig.F, Acropora pectinata: Vaughan. 1918, p.l72, pl.71, fig.I, la-lc, 2; Thiel, 1932, p.ll9, pi. 14, fig.4 (synonymy). Acropora corvmbosa (part): Stephenson and Wells 1956, p.l2- Madrepora surculosa var. turbinata Dana, 1846, p.446: Brook, 1893, p.200 (synonymy). Madrepora turbinata: Verrill, 1864, p.'42. Acropora turbinata: Verrill. 1902, p.242. Note: This .synonymy does not include all references to synonyms, as their interpretations are difficult to trace. Matfriai Examinkd USNM: Fiji Islands, U.S. Expl. Exped., A. hyacinthus holotype 246; Tahiti, U.S. Expl. Exped., A. surculosa (var. turbinata) synlype 251. YPM; Tahiti, A. surculosa var. turbinata type fragment 2017. BM: Fiji Reefs, Challenger, A. conferta holotype 1885.2.1.12; Thursday Is., Saville Kent, A. pectinata syntype 1892.6.8.155; low Woody Is., G.B.R . A. recumhens (var.) 1 892. 6. 8.161. QM: Big Broadhursl Reef; SW side, reef slope: Im, ! Lx. 1973, C-W . G9860, G986I; Im. 13.X.1973, C.W., G9862; l-3m. 22.X.197.3, C.W.. G9863; L5m, 11.x. 1973, C.W., G9859; 2-7m, I2.X.I973, C.W., G9864; 3m. 29.iii.1973, CW., G9872; 5-3m, 13.X.I973. C.W.. G9865:6 3m. 13.x. 1973, C.W.. G9866: 8Tm. 15.x. 1973 C.W.. G9867; Oct. 1973. C.W., G9868: patch reef in lagoon. 2 Lx. 1 973, C.W., G9869; E. side, reef slope, 4m. 2 Lx. 1973, C W.. G9870. Bowden Reef, opening in outer reef. 26.vii.1972, C.W., G9877. G9878: SW side, reef crest, 0-6m. I5.vii.l972, C.W.. GI0716. Bushy-Rcdbill Reef; S. side, outer flat, 3Lv.l975, C.W., G9874: NW.side, top of patch reef off reef edge, 19.xii.I972, E. Lovell, GI0727; W. of Redbill Is., edge of surge channel, 0-5m, I9.xii.I972, C.W., GI0232. WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF 289 Darley Reef, patch reef in lagoon, 24.jii.1973 C W G9876. Great KeppcI Island, E. side, fringing reef, Dec. 1975 R. Woolley, G 1 0723, G 10726. Heron Island: reef flat, W. side: 2.vii.l973, Y. Loya. G9875; 6.vii.l973, Y. Loya. G10720. July 1973, C.W.[ G107I8, GI0722; N. side, 9.viii.l950, W. Stephenson, G9684; reef flat. April 1954, E. Grant, G2680; N. side, 14.V.I954, W. Stephenson, G2681-3. Lady Musgrave Reef, W. side, outer slope, 16.vii.197L 1. Neuss, G6268. Tryon Reef, E. side, reef flat, 12.vii.l971. C.W., G6267; NW. side, reef slope, I8.xii.l971, I. Neuss G6478, G6480, G6494. Viper Reef, patch reef in lagoon, I m, 16.vii.l973 C.W., G10717. Bramble Cay, Oct. 1924, C. Hedley, G.B.R.C. 34-9 86-92. Coates Reef, 13.vii.l924, ‘Geranium’, G.B.R.C. 141, 195. Flora Reef. 21. v. 1924, Dr Paradice, G.B.R.C. 138. Flinders Reef. Moreton Bay, NW. margin, 9.viii-I972, E. Lovell, G6984; NW. slope, 2Lvi.l973, E. Lovell, G7297, G7298. Field Diagnosis Tabular to plate like colonies to over 3m diameter, formed by the developmental process described by Floffmeister (1925). Colonies are sturdy in texture, and the vertical branchlets have a rosette like appearance due to the regular arrangement of labellate corallites around them. Colours are blue with pink growing edge (common reef flat colouration), pale to dark brown, pink-, yellow-, green- or blue-brown. Laboratory Diagnosis Branching pattern: From a central to lateral stalk, branching is horizontal, with branches anastomosing to a greater or lesser extent. Secondary branchlets are single or in groups of up to seven, from 1 mm to 22 mm long, and from 3 mm to 8 mm wide (commonly around 10 mm long and 4 mm wide). In very shallow reef flat situations, the branchlets may be reduced to mere buds, but very wide (up to 8 mm). The longest branchlets are in lagoonal specimens. In this species and in A. cytherea, layers may be added to the original plate. Axial corallites: Up to 1-5 mm exert; outer diameter 1-4 to 2 0 mm; inner diameter 0-6 to LI mm. Septalion: primaries present to 2/3R, secondaries absent, or a few present to 1/4R. Radial corallites: Tubular appressed, with only lower (or outer) wall developed (2/3 circumfer- ence or less) to form a rounded lip. Radials are arranged evenly and closely around the branchlet, giving a rosette-like appearance, typical of this species. Septation may be strongly developed, ail primaries being present to I /4R, plus a few secondaries, or so weakly developed that only the directive septa can be distinguished. On horizontal branches immersed corallites occur. Coenosteum: Coslatc on radial corallites, reticulate with occasional laterally-flattened spines between. Within Reef Distribution This species occurs in lagoons, deep water reef flat situations, shallow outer reef flat, and on the reef slope to about 30 m (species no. 4 in Wallace 1975). Identification Difficulties and History Dana’s type of A. hyacinthus (illustrated by Hoffmeister 1925) is a young specimen just in the Vasiform’ stage of development. This compares closely with Juveniles of the species I have described, which is a sturdier species than A. cytherea. Much reef flat material of the species (well calcified) has probably been identified as A. corymbosa, while the interpretations of A. hyacinthus (e.g. by Hoffmeister) tend to be of a lighter-textured form. Whether his material was of A. cytherea or of a geographic variant of A. hyacinthus I am not able to judge. The deepest occurring colonies of A. hyacinthus are light textured with reduced radial corallite lips. These could be confused with A. cytherea from shallower depths but from a comparable depth A. cytherea has the distinctive ‘A. reticulata’ form. A. conferta (Quelch) has slightly smaller axial corallite dimensions than those given here, but on radial corallite characteristics it appears to be a synonym. This is one of the most successful Indo-Pacific species of Acropora: it is an early colonizer, and occurs with great abundance in some areas, Hoffmeister's detailed treatment of a species under this name, without parallel detailed treatment of A. cytherea (or 'A. reticulata'), leaves unsolved a taxonomic puzzle. A study of these two species on the Samoan reefs would be extremely useful. Geographic Distribution Widespread Indo-Pacific distribution from the Mascarene Archipeligo (G. Faure pers. comm.) to Tahiti. Exact localities cannot be quoted because of taxonomic confusion. Acropora cytherea (Dana, 1846) (Plates 63, 64A, D, 66C, D, 67) Madrepora cytherea Dana, 1846, p.441, pi. 32, fig. 3a, 3b; Brook, 1893, p.99 (synonymy). 290 MEMOIRS OF THE QUEENSLAND MUSEUM Acropora cytherea: Crossland. 1952, p.215. Madrepora efflorescens Dana. 1846, p.441, pi. 33, fig. 6; Brooic, 1893, p.35 (synonymy). Acropora efflorescens: Pillai and Scheer, 1976, p.26, pl.3, fig.3. Madrepora armala Brook, 1892, p.452; 1893, p.lOO, pi. 10, figs. A, B (synonymy). Madrepora reticulata Brook, 1892, p.461: 1893, p.68, pi. 4, figs. A, B. Madrepora reticulata var. cuspidata Brook, 1893, p.69. Acropora reticulata: Wells, 1954, p.422, pi.] 10, figs. 4-6, pi. 114, figs.1-6 (synonymy); Pillai and Scheer, 1976, p.28, pl.7, fig.l, Madrepora arcuata Brook. 1893, p.l02, pi. 12; Studer, 1901, p.395. Acropora cytherella Verrill 1902, p.253, pi. 36, fig. 7, pi. 36a, fig.7. pl.36F, fig.l. (synonymy). Acropora corymbosa 'cytherea Form’: von Maren- zeller, 1907, p.32. pl.l, figs.I. 2; pl.2, fig.3. Acropora hvacinthus (part): Hoffmeister, 1925, p.64; Wells, 1954, p.421. MaTI KIAI E\AMINt:D USNM: Tahiti, U.S. Expl. Exped., A. cytherea syntype 226; Samoa, A. hyacinthus (id. Hoffmeister), no. 14 Mayor Collection. YPM: Fiji Islands, U.S. Expl. Exped., A. cytherella holotype 2007; East Indies, A. efflorescens ?type 1799. BM: Singapore, A. arrttata syntypes 1850.1.16.1, 1857.4.6.1; Amirante Islands, A. reticulata syntype 1882.10.17.131; Navigator Island, A. arcuata syntypes 1862.1.27.5, 1875.10.2.9. QM: Big Broadhurst Reef; S W. side, reef slope: I m, 13. x. 1973, C.W.. G9841, G9842; 1-3 m, ll.x.1973, C.W., G9837, G9838; 2 m. 26.iii.1973, C.W., G9853; 2-3 m, I2.X.I973, C.W., G9839, G9840; 3 m, 28.iii.1973. C.W., G9856; 5 m, 27.iii.1973, C.W., G9855; 7 m, 13.X.I973, C.W., G9843; 8-1 m, 14. X.1973, C.W., G9845; 8-2 m, I4.X.1973, C.W., G9844; 8-3 m, I5.X.1973, C.W., G9847; 8-7 m, 15. X.I973, C.W., G9846; 9-7 m, 15.X.1973, C.W., G9849; 10-6 m, 6.x, 1973, C.W., G9848; SW. side, surge channel floor, 5 m, 20.X.I973, C.W., G9850. Bushy-Redbill Reef. NW. side, reef slope, 21.xii.l972, C.W.. G9857. Tryon Reef: NNE. side, outer slope. 21.xii.I97], I. Neuss, G6482; NW. side, outer slope, 6-10 m, 19.xii.l97I. I. Neuss, Ci6479. Viper Reef, patch reef in lagoon. 1 m from top, 16. vii.l972. C.W., G107I7. Flinders Reef, Moreton Bay: 1973, E. Lovell, G7290, G7291; NW. margin, 3-12 m, 9.viii.l972, E. Lovell, G6999, G7032; W. margin, 10 m, 10.iv.l972, E. Lovell, G7303, G7309. Solitary Islands, 1972, J. Veron et al., G7057. G7058. Field Diagnosis Tabular to plate-like colonies to over 3 m diameter, formed by the developmental process described by Hoffmeister (1925) for A. hyacinth- us. Colonies have a light crumbly texture. Axial corallites are usually obviously exert, and contrast with slight scale-like radial corallites. Horizontal branches are reticulated to fully anastomosed. Colours are cream, pale brown, pink-, yellow-, green- or blue-brown or -grey. Laboratory Diagnosis Branching pattern: From a central to lateral stalk, branching is horizontal, with branches anastomosing to a greater or lesser extent. Secondary branchiets are vertical and single or in groups of up to six, from long and slender (up to 20 mm length, width around 3 mm at this length), to short bundles of tubular axials approximately 5 mm long with occasional proliferation of radial corallites at their base. In the series of specimens from lesser to greater depths on the Big Broadhurst Reef, a transition occurs from long branchiets with non-proliferous tips (identifiable with ‘A, arcuata’ and A. armata), through shorter branchiets with proliferous tips, the proliferations often partially naked of radial corallites, to very short branchiets with few radial corallites (identifiable as ‘A. reticulata’). All specimens are openly reticulated and lightly calcified. Lagoonal colonies are more anastomosed and heavily calcified and the branchiets are similar to those of deep reef slope colonies. Specimens from the geographically extreme southern locations (Flinders Reef, Moreton Bay and the Solitary Islands) have the main branches strongly anastomosed, and bundles of short thick proliferous branchiets: they identify with A. cytherea sens, strict. Axial corallites: From 1-0 to 5-0 mm exert. Outer diameter 1-3 to 1-7 mm in Great Barrier Reef specimens. 1-6 to 2-5 mm in the Flinders Reef — Solitary Island specimens; inner diameter 0*7 to 10 mm. Septation: primary septa present, up to 2/3R; secondary septa often absent, never fully present, up to 1 /4R. Radial corallites: Tubular appressed with only lower (or outer) wall (1/2 circumference or less) developed, to form an elongate lip. In A. cytherea s.s. forms these lips are thickened: on the reef slope they are light structured and may be reduced to single or double points. Septal development of radials very poor: usually only directives can be seen, sometimes a few other primaries are present WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 291 as fine points. On horizontal branches immersed coralliles occur. Coenosteum: Costate on radial corallites, reticulate with laterally flattened spines in between. Within Reff Distribution On the Great Barrier Reef this is a reef slope species (species no. 5 in Wallace 1974), but occasional colonies occur in lagoonal and deeper water reef flat areas. On other reefs (e.g. Enewetak Atoll) it has a similar distribution (C.W.). Identification Difficulties and History Two special taxonomic problems are associated with this species: (1) In combining A. cytherea with A. reticulata, and (2) In distinguishing between lightly structured colonies of A. hyacinthus and colonies of A. cytherea with good branchlel development. A. cytherea s.s. occurs in geographically extreme locations (Tahiti, Solitary Islands, Moreton Bay). The temptation is to retain A. reticulata, which best describes the species in its main tropical range. The second problem can also be staled as a problem of interpretation of A. arcuata and A, armata. These and A. cytherea have been combined with A. hyacinthus by recent authors. On A. arcuata and A. armata my own observations are of a co-occurrence of these ‘species’ with definite A. hyacinthus at the same depth on the Big Broadhurst reef front, and their graduation into typical 'A. reticulata' with depth. Crossland’s works are important to the ecological documentation of this species: in 1928 he identified A. cytherea as A. hyacinthus, in 1931 reversing the identification. In 1952 he notes that both species are common in Tahiti. This species, as I have defined it, is easily separated from A. hyacinthus in the field by its light crumbly skeleton. The two species co-occur on the reef front, A. cytherea extending a little deeper, and A. hyacinthus extending more into reef flat areas. Geographic Distribution The species has a wide Indo-Pacific distribution from the Mascarene Archipeligo (G. Faure, pers. comm.) to Tahiti. Acropora millepora (Ehrenberg, 1834) (Plates 68, 81 D) Heteropora millepora Ehrenberg, 1834, p. 109. Madrepora millepora: Brook, 1893, p.Il6 (synonymy). Acropora millepora: Verrill, 1902, p.257; Thiel, 1932, p.l24, pi. 18, fig. I, pi. 19, fig.l; Nemenzo, 1967, p.94, pi. 28, figs.l, 2. Madrepora spathulaia Brook, 1891, p.469; 1893, p.l2I, pi. 32, fig.B. Madrepora squamosa Brook, 1892, p.463; 1893, p.l20, pl.20, fig. B; Acropora squamosa: Vaughan, 1918, p.l73, pi. 72, figs.l, 2, 2a, 3; Crossland, 1952, p.216; Stephenson and Wells, 1956, p.l8. Acropora sarmentosa: Vaughan, 1918, p.l7A, pi. 72, fig.4. MAThRiAL Examined BM: Claremont Is., Great Barrier Reef, Saville Kent: A. millepora (mentioned Brook, 1893) 1892.6.8.165; A. squamosa syntype 1892.6.8.163; Treasury Is., Solomon Islands, D.A. Guppy, A. spathulata holotype 1884.12.11.25. MNB: A. millepora holotype, 854 (photograph). QM; Big Broadhurst Reef: patch reef in lagoon, 28.111.1973, C.W., G1 1052-6; E. side (weather side), channel in outer flat, 21.X.1973, C.W., G1I055. Bushy-Redbill Reef: NW. side, outer reef fiat, 23.xii.1972, C.W., G11047, G11049; NW. side, reef slope, 23.xii.1972, C.W., G11046. Bowden Reef, SW. side, reef crest, 26.vi.1972, C.W., G11064. Darley Reef, patch reef in lagoon, 1 m, 18.vi.l972, C.W., G1I057; 3 m, 22.iii.1973, C.W., G11058; 4 m, 22.111.1973, C.W., G11059. Heron Island: N. side, reef flat, 14.xii.l973, C.W., GI0996-G1 1001; S. side, outer reef flat, 12.xii.l973, C.W., G 10973, G 10976-83; SW. side, outer reef flat, I3.xii.l973, C.W., GI0985-93; W. side, reef flat: 15.xii.l973, C.W., Gl 1002-5; 17.xii.l973, C.W., G1 1006-8; SE side, reef flat, 17.xii.l973, C.W., Gl 1009-16. Great Keppel Is., Dec. 1975, R. Woolley, G11061. Maer Island, Murray Island group, N. side, inner reef flat: 16.vii,1974, G. Ingram, G1I060; 18.vii.l974, G. Ingram, Gl 1062. Masthead Reef, reef flat, Aug. 1974, J. Buhmann, GI1063. Michaelmas Cay, 1974, C. Limpus, G11067. Low Isles, 1974, C. Limpus, G 11066, G 11068. Fiji Islands, Great Astrolabe Reefs, Feb. 1974, C.W., Gl 1033-43. Field Diagnosis Low corymbose or stalked corymbose colonies, commonly with neat round outline; branches terete or slightly tapering, covered by evenly sized and closely arranged radial corallites with flaring, scale like lips. Commonest colouration (on both the Great Barrier Reef and Fijian reefs) is a dull to brilliant green with dull to bright orange branch tips. The colonies can also have multiple colouration, being predominately blue or pink, 292 MEMOIRS OF THE QUEENSLAND MUSEUM with touches of other pastel colours and a grey sheen, or bright orange to pale cream. Laboratory Diagnosis Branching pattern; Branches are given off vertically to obliquely from a central to lateral region which may be consolidated into a stalk. In some colonies from sandy situations a growing point is barely recognizable, and the growth is bush like and relatively indeterminate. A small amount of secondary branching occurs, and the top of the colony is in a single plane. Branch width varies from 7 to 13 mm. Axial corallites: Barely exert. Outer diamter 24 to 3-9 mm; inner diameter 0-9 to 1*6 mm. Septation: primary cycle fully developed, up to i/2R; secondary cycle usually represented, but not all septa developed, up to 1 /4R. Radial corallites: No upper wall is developed, the lower half of the wall is expanded as a rounded lip, and the outer edges of this lip may Bare away from the opening of the corallite. The primary septa are often well developed, up to 2/3R, secondaries absent or a few present to 1 /4R. Coenosteum; Coslate on radial corallite lips, reticulate with simple spines between radials. Within Reei Distribution Reef flat, from sandy middle reef flat to consolidated outer flat, occasionally to a few metres below the reef top; tops of lagoonal patch reefs and fringing reefs. Identification Difficulties and History The usual name applied locally to this species is 4. squamosa, but the material from the Great Barrier Reef identified as A. millepora by Brook (apparently the last person to observe and describe Ehrenberg’s type) belongs to this species, and in fact forms a series from the same locality as his A. squamosa types. Verrill (1902) synonymised the two, but he has not been followed by later authors. Two species regarded as being similar to A. millepora (or A. squamosa*) may now be dismissed: A. sarmentosa was misinterpreted by Vaughan (1918), and is a clearly definable species; A. aspera (= A. hebes) with many synonyms is a highly versatile reef flat species which in its low corymbose form appears very similar to this species. A. millepora has a more restricted reef flat distribution than A. aspera, and extends closer to the reef crest. In aggressive interactions observed at Heron Island and on Bushy-Redbill Reef A. millepora tissues appear to be inclined to overgrow those of A. aspera. Small colonies of the species have- probably been identified as ‘4. corymbosa* by some authors. 4. imbricata (Ehrenberg) is another possible synonym. Geographic Distribution The most definite records of this species are only from the Great Barrier Reef, Solomon Islands, Fiji Islands (C.W.) and Marshall Islands (C.W.). Other less certain records extend the range to Ceylon, the Philippines, New Ireland, New Hebrides. Acropora delicatula (Brook, 1891) (Plate 69) Madrepora delicatula Brook, 1891, p.461; 1893, p.l09, pi. 28, figs. D, E. Acropora delicatula: Wells, 1954, p.420, plT15, figs.l, 2. non Acropora delicatula; Stephenson and Wells, 1956, p.l2. Material Examine^d BM; Solomon Islands, Guppy, 4. delicatula holotype 1884.12.11.23. QM; Big Broadhursl Reef, SW.side, reef slope: 2 m, 26.111.1973, C.W., G11442; 6 m, 25.iii.1973, C.W., Gll44i; 6 m, 27.iii.1973, C.W., G1I439; 8 m, 26.111.1973. C.W., G11436; 8-1 m, 14.x. 1973, C.W., GI1434; 8 6 m, 15.x, 1973, C.W., 011438; 13-4 m, 17.X.1973, C.W., G11437; 17 m, 23.X.1973, C.W., G1 1435. Bushy-Redbill Reef, NW. side, reef slope, 3-7 m, 21.X11.1972, C.W., G11440. Field Diagnosis Shallow caespitose or corymbose-plate colonies have slender vertical branchlets with small, scalc-like radial corallites. Colours are cre.*m, pale brown or greenish brown. Laboratory Diagnosis Branching pattern; From an attachment that is lateral or nearly so, branching is at first horizontal, then branchlets or bundles of branchlets are given off vertically upwards. The branchlet bundles can be as tall as 40 mm, and branchlet width is around 5 mm. A small amount of branching from the undersurface may occur. Axial corallites: 0-5 to I *5 mm exert. Outer diameter 1-5 to 2 mm; inner diamter 0-6 to 0*9 mm. Septation: primary septa well developed, up to 3/4R, secondary cycle usually absent, or a few septa may be present up to 1/4R. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 293 Radial corallites: Appressed tubular, nariform or dimidiate, the outer wall only developed, and extended as a Up which is correspondingly rounded, elongate or spade-shaped. The septal development varies, but commonly all primary septa are visible, up to I /4R, and two or three secondaries just visible. Coenosteum: Costate or rows of simple spines on radial corallites, neat rows of simple spines, or reticulate with scattered spines, between. Within Reef Distribution I have observed only a very few colonies, all in reef slope situations. The species is much more commonly represented from fringing reefs, and a series in J.C.U. is from the Lizard Island and Palm Islands reefs. Identification Difficulties and History This species is poorly documented, and working with a poor series I am unable to resolve some problems. A. subulata (Dana), may be a senior synonym. Stephenson and W'ells’ specimen from Low Isles (QM G2648) is A. tenuis. Geographic Distribution Great Barrier Reef, Solomon Islands, Marshall Islands. Acropora haimei (Milne Edwards and Haime, 1860) (Plates 70, 71) Madrepora haimei Milne Edwards and Haime, 1860, p.l51; Brook, 1893, p.77 (synonymy); von Marenzeller, 1907, p.51 pi. 16 figs. 45-48. Acropora haimei: Vaughan, 1918, p.l63, pi. 70, figs. 3, 3a. 3b, pi. 66. figs, 4. 5; Crossland, 1952. p.207. pl.33, fig. I, pi 35. fig. !: Ros.si. 1954, p.48; Stephenson and Wells, 1956, p.l4; Nemenzo. 1967. p.82. pi. 25. fig. 1. Material Examined QM: Big Broadhurst Reef: SW. side, reef slope: 9-8 m, 16.X.1973, C.W., G10216, G102I7; 9-6 m, 15.x. 1973, C.W., G10215; 8-7 m, 15.X.1973, C.W., G102I3, G102I4; 8-6 m, 15.X.1973, CW., GI0212; 8-3 m, 15.X.1973, C.W., G10211; 8 m, 14.X.1973, C.W., G102I0; 7-8 m, 23.X.1973, C.W., G10206-9; 7 m, 13.X.1973, C.W., G10205; 5-6 m, 23.X.1973, C.W., G10204; 6 m, 13.x. 1973, C.W., G10224; 5-7 m, 23.X.1973, C.W., G10225; 5 ni, 27.iii.1973, C.W., G10182, G10221; 5 m, 25.iii.1973, C.W., G10181; 4-2 m, 22.X.1973, C.W., G10222; 3-8 m, 12.X.1973, C.W., G10180; 1-6 m, 22.X.1973, C.W., G10223; 15 m, 12.X.1973, C.W., G10179; 1-3 m, ll.x l973, C.W., G10202; 1-3 m, 14.X.1973, C.W., G10203; 1 m, 26.iii.1973, C.W., G10194; SW. side, surge channel opening, 10 m, C.W., G10183; patch reef in lagoon, 21.x,1973, C.W., G10184, G10185. Bushy-Redbill Reef; NNE. side, reef slope, 3 m, 30.xii.l972, C.W., G10186, G10187, Gl 1303; NW. side, reef crest, 30.xii.l972, C.W„ G10195, G10227, G10228; W side, reef crest, 18.xii.l972, C.W., G10226. Bowden Reef, opening in SW. side of reef, 26.vii.1972, CW., G10188, G10I89, G10229-31. Field Diagnosis This is a medium sized compact arborescent species, which occurs as scattered turf like patches, bushes or thickets. It is distinguished from other arborescent species by very open radial corallites. Colour is usually cream or pale brown. L aboratory Diagnosis Branching pattern: Branching is open and irregular. Main branches are up to 15 mm wide and tapering. Axial corallites: Outer diameter 2-2 to 3-5 mm; inner diameter 0-8 to 1-2 mm. Septation: primary septa present up to 2/3R; secondary septa all present, or mostly present, up to 1/3R. Radial corallites: Tubular, extend at from 45° to 90° from branch; openings cochleariform; the upper part of the corallite wall (about 1/3 of the diameter) is thinner and shorter than the rest of the wall, and the thicker lower portion often flares slightly. Septal development is usually very marked: primaries can extend to R, and a full set of secondaries can be present. A good key to the identification of this species is the presence, on the proximal part of branches, of well formed cochleariform radials with stiong septal development. Coenosteum: Costate on ladials, reticulate with simple or laterally flattened spines in between. Identification Difficulties and History Although following Crossland’s interpretation, I am still not fully satisfied that A. haimei is the correct identification of this species. Milne Edwards and Hairne’s type cannot be located. Other authors, particularly von Marenzeller. describe a greater variety of colony shapes than I have seen, including shallow water reef flat forms which approach corymbose. To a worker with some experience in the field, this species becomes easy to distinguish from other arbores- cent species because of its large open radial corallites. Whhin Reef Distribution The species occurs as small patches on middle and outer reef flats, but achieves its greatest 294 MEMOIRS OF THE QUEENSLAND MUSEUM abundance on the reef slope (to about 10 m), in Heron Island, W. side, reef flat, 6.vii.l973, Y. Loya, surge channel openings and on sandy bottoms in G11457. deep lagoons and around broken reef- patch Masthead Reef, SW. side, upper reef slope, Aug. ai-goc 1974, J. Buhmann, G11419. Geographic Distribution Indo-Pacific: Red Sea, ?Diego Garcia, Maidive Islands, Ceylon, Mauritius, Singapore, Great Barrier Reef, Fiji Islands. Acropora tenuis (Dana, 1846) (Plates 72, 73) Madrepora tenuis Dana, 1846, p.451; Ortmann, 1888, p.l52; Brook, 1893. p.83 (synonymy). ? Madrepora eurystuma Klunzinger, 1879, p. 16, pi. L fig. 8, pi. 4, fig. 7, pi. 9, fig. 12. Madrepora eurystoma: Brook, 1893, p. 137 (synonymy). Madrepora macrostoma Brook, 1891, p.464; 1893, p. 105, pi. 19, fig. B. Madrepora bifaria Brook, 1892, p.453; 1893, p.llO, pi. 30, fig. A. Madrepora kenti Brook, 1892, p.458; 1893, p. 110, Pl- II, fig. B. Madrepora dilatata Brook, 1893, p.8I. Madrepora anihocercis Brook, 1893, p. 106, pl.l3, fig. C (synonymy). ?Acropora anihocercis: Nemenzo, 1967, p. 109. Materiai. Examined USNM; Fiji Islands, U.S. Expl. Exped., Madrepora tenuis holotype 259. BM; Koseir, Red Sea, A. eurystoma 1886. 10.5.5; Diego Garcia, G. C. Bourne, A. eurystoma 1891.4.9.1. (mentioned Brook, 1893); Mauritius, A. macrostoma holotype 1878.2.4.7; Java, A. bifaria holotype 1859.12.12.2; Thursday Island, Saville-Kent, A. kenti holotype 1892.6.8.202; Palm Island, Saville-Kent, A. anthocercis syntype 1892.6.8.235; Rocky Island, Saville-Kent, A. anthocercis syntypes 1892.6.8.236, 1892.6.8.237. QM: Big Broadhurst Reef, SW. side, reef slope; 1-9 m, 22.X.1973, C W., GII413; 3-3 m, 22.x.l973» C.W., G11429; 4 m, 24.iii.1973, C.W., 011431; 6 m, 27.111.1973, C.W., G11428; 7 m, 28.iii.1973, C.W., GI1430; 7-8 m, 14.X.1973, C.W., G11456; 8 m, 26.111.1973, C.W., G11433; 8 m, 14.X.1978, C.W., 011420; 8 1 m, 14.X.I973, C.W., G11424; 8-2 m, 14.X.I973, C.W., G11422, GII423; 8-6 m, 15.x. 1973, C.W., G1 1425; 9-2 m, 15.X.1973, C.W., G1 1421; 9-8 m, 16.X.1973, C.W., G11426, GU432. Bushy-Redbill Reef: W. side, outer reef flat, Dec. 1972, C.W., G114I7; NW. side, reef slope: 19.xii.l972, E. Lovell, G11412, G11415, G11416; NW. side, floor outside slope, 22.xii.1972, C.W., G1I427. Darley Reef, patch reef in lagoon, 22.iii.1973, C.W., G11414, G11418. Field Diagnosis Colonies are Thick plates’ or caespito- corymbose. The radial corallites are evenly arranged and have distinctive, lip like, flaring outer wall and strong septal development, which can often be seen with the naked eye. Once learnt in the field, this species is always easily recognised. The commonest colour is cream, less comrnonly colonies are greenish-blue or bright blue. Polyps are often partly extended during the day, and in cream colonies these are often bright orange (axial polyps) and bright purple (radial polyps). Laboratory Diagnosis Branching pattern: From a growing area which may be central to lateral, branching is horizontal, then secondary branchlets or groups of branchlets are given off vertically to obliquely, their tips in one plane. Branching may also occur from the lower surface but these lower branches are never as long as the upper. Branchlet widths are from 6 to 8 mm. Shallow water specimens are Thick plates’ of up to 110 mm vertical depth; deeper water specimens may be thin plates as little as 35 mm thick. Axial corallites: Up to 2 mm exert. Outer diameter: 1-9 to 3-0 mm; inner diameter 0-8 to 1-2 mm. Septation; both cycles usually developed, the primaries up to 2/3R, the secondaries up to 1/3R. Radial corallites: Tubular, ascending, the inner wall less developed (both in thickness and length) than the outer; the outer wall flaring, so that the opening appears large and round or slightly oval. Towards the middle of the branches these radials have a classic 'cochleariform' appearance. Both septal cycles are usually developed, and in some reef flat and lagoonal specimens these may fill the corallite. The size of the radials is very even over the corallum. With increasing water depth the corallite wall becomes less flaring, the ‘cochlear- iform’ appearance is lost, and the corallites are more scattered. Coenosteum: Either strongly costate or in rows of simple spines on the radial corallites, reticulate with simple spines between. Within Reef Distribution Reef flat (deeper water areas), lagoonal patch reefs, reef slope to about 10 m. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 295 Identification Difficulties and History The synonymy list for A. tenuis is large, and possibly still incomplete. The synonyms differ in features such as length of branches and degree of calcification. In the field, the colonies look quite similar to those of A. aculeus (Dana). Some field experience is necessary to distinguish them. The main differences are: (a) wide flaring lip and larger radial coralliles in A. tenuis: (b) colour pattern: A. tenuis is usually cream or pinkish brown while A. aculeus has a distinctive double colouration; (c) A. tenuis very often has the polyps expanded, as noted. The shape of the radial corallites shows some similarity to that in A. haimei, and strong similarity to that in A. striata Verrill, which does not occur in the Great Barrier Reef regions studied, but is a common species in the Marshall Islands. One specimen from Lizard Island in the J.C.U, with bottlebrush growth form is possibly A. striata. Geographic Distribution Indo-Pacific: Mauritius, Red Sea, Diego Garcia, Indonesia, Philippines, China Sea, Great Barrier Reef, Fiji Islands. Acropora tubicinaria (Dana, 1846) (Plate 74) Madrepora tubicinaria Dana, 1846, p.451, pi. 32, fig.7; Brook, 1893, p.l39 (synonymy). Acropora tubicinaria: Verrill, 1902, p.2I9; Wells, 1954, p.423, pi. 122, figs.3-5. Material Examined USNM: Fiji Islands, U.S. Expl. Exped., A. tubicinaria holotype 258. QM: Bushy-Redbill Reef: W. side, inner reef flat: 28.V.1975, C.W., G1 1071-6, G11080; Jan. 1973, C.W., G1 1077; 18.xii.l972, C.W., Gl 1081; NW. side, first reef crest, 22.xii.1972, C.W.. G11082; S. side, microatoll zone, 2.i.l973, C.W., G11078, G11079, G11083. Masthead Reef, August 1974, J. Buhmann, G11088. Great Keppel Island, Dec. 1975, R. Woolley, Gn089. Field Diagnosis Colonies are small rounded caespitose clumps (largest seen 25 cm diameter) in which the openings of the radial corallites are obvious. Colour is brown, occasionally with blue tips. Laboratory Diagnosis Branching pattern: From a central growing area branches are given off vertically to obliquely, and these branch again, sometimes infrequently, sometimes frequently. As colonies mature the base and basal parts of branches may die, and portions of the colony become separate from each other. Branches are from 8 to 10 mm thick, and may be truncate or strongly tapered. Axial corallites: Non exert. Outer diameter 1-8 to 3-2 mm; inner diameter 1-0 to 1-2 mm. Septation: both septal cycles developed, primaries up to 3/4R, secondaries up to 1 /4R. Radial corallites: Short tubular, usually upper part of wall less developed than lower, opening circular. Usually both septal cycles are developed, but just visible, although on parts of the corallum primaries can reach 1/2R and secondaries 1/4R. Coenosteum: On some colonies lines of simple spines are arranged both on and between radials. More commonly the coenosteum is costate or in lines of spines on the radials, and reticulate with simple spines between. Within Reef Distribution On Bushy-Redbill Reef, this species is common in the coral depauperate inner reef flat (recorded as A. digitifera by Wallace and Lovell 1977). With A. palifera and species of Porites it dominates this zone and it is rare on other parts of the reef. Identification Difficulties and History This species is rarely mentioned in the literature. On my evidence and that of Wells, it tends to specialize in certain reef zones (which may be characterized by poor coral cover), but is common in these areas. It is rare in collections brought to the QM for identification, and 1 have not yet seen it from fringing reefs. A. striata Verrill mentioned by Wells (1954) as a ‘related form’ does not occur on the Great Barrier Reef. Geographic Distribution Great Barrier Reef, Fiji Islands, Marshall Islands, Tahiti. Acropora aculeus (Dana, 1846) (Plates 75, 76) Madrepora aculeus Dana, 1846, p.450, pi. 32, fig. 6; Brook, 1893, p.I04 (synonymy). Acropora aculeus: Faustino, 1927, p.269; Nemenzo, 1967, p.lI4. Madrepora tubigera Horn, 1860, p,435; Verrill, 1864, p.4!; Quelch, 1886, p.l61; Brook, 1893, p.79 ( + further synonymies); Verrill, 1902, p.239, pi. 36, figs. I, 2-2b. pi. 36a, figs.l, 2-2b; pl.36f, fig. 8. 296 MEMOIRS OF THE QUEENSLAND MUSEUM Acropora tuhigera: Crossland, 1952, p.208. ''f Madrepora ruina Studer. 1878, p.533, pi. 2, figs. 6a, 6b; Brook. 1893, p.82. ‘ Acropora nano: Wells, 1950, p.39, pi. 10, figs. 3, 4. Acropora nana Nemenzo, 1967, p.85. Madrepora patula Brook, 1892, p.460; 1893. p.l 1 1, pi. 9, fig.E. Acropora paiula: Crossland, 1952, p.215; Stephenson and Wells, 1956, p.l6; Nemenzo, 1967, p.l02. ‘^Madrepora elegantvla Ortmann, 1889, p.507. pi. 12, ng.5. Madrepora elegantula: Brook, 1893, p.l 15. Mathrial Examinhd USNM; Fiji Islands. U.S. Expl. Expd., A. aculeus holotype 257; Cocos-Keeling, A. nana (mentioned Wells, 1950) 44322, VTM; A. tuhigera fragment of type 1483. BM: Port Denison, Great Barrier Reef, A. patula holotype? 1892.6.8.274. QM; Big Broadhurst Reef: SW, side, reef slope: I3.X.I973. C.W., G9093; 3 m, 28.iii 1973. C W., G9097; 5 m, 27.iii.1973, C.W., G9I07; 51 m, 23.X.1973, C.W., G9086. G9087; 6 m. I3.X.I973, C.W., G9092; 6 1 m, 13-X.1973. C.W., G9090; 7 m, 28.iii 1973. C.W.. G9106;8 m, 26.iii 1973, C.W., G9108; 8-2 m, 14 x. 1973. C.W., G9094: 8-5 m, 23.X.I973, C.W., G9100; 8-7 m, 15.x. 1973. C.W., G9095; 9-2 m, 23.X.1973, C.W., G9088; 10 rn, 20.X.I973, C.W.. G9099; 10 m, 23.X.1973, C.W.,G9101; I()-6 m. 16.x. 1 973, C.W., G9102; 13-3 m. 23.x.I973,C.W.,G9089;25 m, 1 7.x. 1973, C.W., 09091; 25 1 m. 18.X.1973, C.W.. G9I04; SW. side, surge channel Boor. 10 m, 20.x. 1 973. C.W., G9098, G9108. Bushy- Redbill Reef; W. side, middle reef flat: 8.vi-t975, CW, G9122-4: 14.vi.l975, C.W., G9125. G9126; W side, outer reef flat, 31. vi. 1975, C.W., G9I16; W. side, reef slope, 1-8 m, 3.vi.l975, C.W , G91 18, C;9120; NW. side, middle reef Hat, 30.xii.I972, C.W., G9114; NW. side, patch reefs; I9.xii.l972, E. Lovell. G9542; 15 m, 16.vi.l975. C.W., G9117; NW. side, reef slope, 3 7 m, 21.xii.l972, C.W., G9115; adjacent Redbill Is., reef crest: 19.xii 1972, CW, G9113; I.vi.l975. C.W., G9119; adjacent Redbill Is., reef slope, 20.xii.l972, C.W., G11463. Darley reef, patch reefs in lagoon: 3 m, 18.vii.l972, C.W., G9I10; 3 m, 22.iii.1973, C.W., G9I09; 4 m. 22.iii.1973, C W., G9111. Prawn reef, patch reef in lagoon, 15.vii.l972, C.W., 09112. Field Diagnosis This species occurs as Thick plates’ or caespito-corymbose units. The colony can be one plate with a loose central or side attachment, or a number of tiers of plates can develop. The single plates occur on the outer reef flat and deeper water parts of the reef slope: the layering develops in deep water middle reef flat areas and on the reef crest. Small round colonies can occur on the outer reef flat. Colours are blue, grey, green, or brown on lower parts of branches, with tips of branches yellow, lime green, pale blue or brown. Laboratory Diagnosis Branching pattern: From a usually lateral attachment, main branches are horizontal and secondary branchlets or bundles of branchlets are given off horizontally to obliquely on both sides of them. Branchlets are slender (3 to 7 mm). The ‘stout branches’ of Dana was either a slip of the pen or a reference to the main stem. Small shallow reef flat specimens grow as vertical branchlets directly from an encrusting base. Axial corallites: Outer diameter 1-8 to 2-4 mm; inner diameter 0-8 to 10 mm. Septation: primary septa well developed, up to 2/3R. secondaries usually partly but poorly developed, up to 1/4R.' Radial corallites- Tubular appressed to partly appressed, with round to slightly oval openings at 90*^ or more to ihe branch. Primary septa developed up to 1 /2R, and some secondaries visible. Radials are usually evenly sized and distributed on branches and show little variation. Wall can be thickened. Coenosteum: I.ines of simple spines on and between radials, or sometimes a spongy appear- ance between radials. Within Reef Distribution Middle and outer reef flats, reef crest, reef slope to 20 m, lagoonal patch reefs. Identification Difficulties and History Much of the variability in this species is related to colony size. Some idea of the scale of this variability can be seen by comparing Well’s A. nana specimen (1950, pi. 10, figs.3, 4) and my Plate 76, Fig.D. Other features such as thickness of wall, angle of radial lip, and completeness of radial septation, vary little, but are responsible for the various species in synonymy. A. elegantula (Ortmann) as interpreted by Brook (1893) (BM 1892.12.5.18) is a heavily calcified specimen of this species. Geographic Distribution Indo-Pacific: Cocos-Keeling Islands, Philip- pines, Great Barrier Reef, Fiji Islands, Samoa, Marshall Islands. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 297 Acropora cerealis (Dana. 1846) (Plate 77) Madrepora cerealis Dana, 1846, p.460, pi. 35, fig.2; Brook, 1893, (part) p.91 (synonymy). Acropora cerealis: Faustino, 1927, p.266, pi. 86, figs.l . 3; Nemenzo, 1967, p.83, pi. 25, fig.2, Madrepora hysirix Dana, 1846, p.476, pi. 40, fig. 1, pi. 31, fig. 5; Brook, 1893, p.l76 (synonymy). Acropora hystrix: Wells, 1954, p.425; pi. 125, figs. 1-4. Madrepora tizardi Brook. 1893, p.89, pl.Il, figs. C, D. (synonymy). Acropora tizardi: Wells, 1954, p.425, pi. 1 25, figs. 5, 6; Nemenzo. 1967. p.l03 Material Examined USNM: Sooloo Sea, U.S. Expi. Exped., A. cerealis synlype 269; East Indies, U.S. Expl. Exped., A. cerealis syntype 270, Fiji Islands, U.S. Expl. Exped., A. hvstrix holotype 298. YPM: Fiji Islands, U.S. Expl. Exped., A. hystrix fragment of type 2039. BM: Tongatabu, J. J. Lister, 1891.3.6.9; Amboina, Challenger, 1885,2.1.1; A. cerealis (Brook 1893 mentioned specimens); Tizard Bank, A. tizardi syntype 1889.9.24.1 15. QM: Big Broadhurst Reef, SW. side, reef slope: 4-5 m, 12.X.1973, C.W., G9532; 5-1 m, 14.X.I973, C.W., G9539; 6 m, 13.x. 1973, C.W., G9535; 6-3 m. 23.X.1973, C.W., G9538, G9531; 8-6 m. 15x1973, C.W., G9533; 8*7 m. 23.X.1973, C.W., G9529; 10 m. 23.X.I973, C.W., G954I; 14*2 m, 23.x4973. C.W., G9530; 13.X.I973. C W., G9537; Oct., 1973, C.W., G9540. Bowden Reef, slope of opening in SW. side, 3 m, 26.vii.1972, C.W., G9519. G9520. Bushy-Redbill Reef, NW. side, reef crest, 22.xii.1972, C.W., G9543. Darley Reef, patch reefs in lagoon: 5 m, 24.iii,1973, C.W., G9524; 1 m, 22.iii.1973, G9525, G9528; 3 m. !8.vii.l972, C.W., G9526. Viper Reef, patch reefs in lagoon, 3 m, 6.vii.l972, C.W.. G9521. G9523. Field Diagnosis Colonies can be untidy caespitose clumps in lagoonal situations, through caespito-corymbose on the upper reef slope, to corymbose plates on the deeper reef slope. Radial corallites are narrow tubo-nariform, with tendency for outer edge to be hooked upwards ('A. tizardi’}. with elongate oblique opening (‘A. hystrix) tubo-nariform tending to nariform (A. cerealis s.s.). The resultant appearance is a colony similar to A. nasuta, but with ‘spinier’ appearance and slender, usually more complexly branching, branchlets. Colour is usually cream to pale brown. Laboratory Diagnosis Branching pattern; From a central to lateral attachment, main branches are given off vertically (mainly in lagoonal colonies) to horizontally. Secondary branching is on all sides of vertical branches or vertically to obliquely upwards from horizontal branches. There is a tendency for further secondary branching. Branchlet widths are around 10 mm, but much of this width is given by the spreading radial corallites. Axial corallites: Outer diameter 1-8 to 21 mm, inner diameter 0-7 to 0-9 mm. Septation: primary septa well developed, up to 3/4R, secondaries absent or a few present to 1 /4R. Radial corallites: The shape of the radial corallites is nariform to tubo-nariform, but the opening varies from rounded to slightly oval (lagoonal specimens) to accentuated elongate. The lower wall may be slightly thickened or may be extended as a ‘hook’. In lagoonal specimens, radials tend to be scattered. In reef-slope specimens with elongate radials these tend to be arranged in neat rows along the branches. Septal development varies, but primaries are always strongly visible, and secondaries usually at least partially visible. Coenosteum: Costate or dense lines of spines on corallites. reticulate with simple spines in between. Within Reef Distribution Deep water reef flat areas, reef slope (to 15 m in present study), lagoonal patch reefs. Identification Difficulties and History Wells (1954) commented on the similarity between A, tizardi and A. hystrix. Unfortunately A. cerealis sens, strict, is the most difficult of the synonyms to place correctly. Without the accentuated oblique radial corallile opening it approaches A. nasuta in general appearance. Deeper reef slope specimens are very lightly structured, and are easily confused with specimens of A. tenuis from similar localities Geographic Distribution China Sea, Philippines, Great Barrier Reef, Fiji Islands, Marshall Islands, Tongan Islands. Acropora nasuta (Dana, 1846) (Plate 78) Madrepora nasuta Dana, 1846. p.453, pi. 34, rig.2; Brook, 1893, p.73 (synonymy). Acropora nasuta: Verrill, 1902. p.257; Hoffmeister, 1929, p.364; Wells, 1954, p.424. pi. 1 13, figs. 5, 6; pi. 124, figs. 1-3; Nemenzo, 1967, p.88, pi. 26, ng.3. 298 MEMOIRS OF THE QUEENSLAND MUSEUM Madrepora nasuta var. crassilahia Brook, 1 893, p.74. Acropora nasuta crassilahia: Wells, 1954, p.425, pi. 1 24, fig. 4. Acropora nasuta var. crassilahia: Nemenzo, 1967, p.89, pi. 26, fig.2. Madrepora effusa Dana, 1846, p.455; Brook, 1893, p.76 (synonymy). Acropora effusa: Verriil, 1902, p.229, pi. 36, fig.l6, 16a, pl.36B, fig.7, 7a. Madrepora cymhicyathus Brook, 1893, p.86 (synonymy). Acropora cymhicyathus: Hoffmeister, 1925, p.63, pi. 13, figs.2a, 2b; Wells, 1954, p.425; pi. 124, figs. 5-7; Stephenson and Wells, 1956, p. 12. Matf.rial Examined USNM: Tahiti, U.S. Expl. Exped., A. nasuta holotype 260. YPM: Point Pedro, Ceylon, A. effusa holotype 8147. QM: Big Broadhurst Reef: patch reef in lagoon: 21.X.1973, C.W., 011212; 28.iii.1973, C.W.. G11227; SW. side, outer reef fiat, 22.iii.!973, C.W„ G11209; SW. side, reef crest: 1-5 m, Il.x.1973, C.W., GI1216; 1-5 m, 22.X.I973, C.W., G11213; 2 m, ll.x.1973, C.W.. G1 1215, 011274; SW. side, reef slope: 7 m, 28.iii.1973, C.W., G11224; 7-6 m, 13.X.1973, C.W., G11217. Bowden Reef, SW. side, reef crest, 1 m, 15.vii.l972, C.W., G1I21I. Bushv-Redbill Reef; adjacent Redbill Is., reef flat: 18.xii.r972, C.W., GI1204; 7.vi.l975, C.W., 011222; 12.vi.l975, C.W., 01 1223; W. side, middle reef fiat; 5.vi.l975, C.W., GI1226; 27.vi.l975, C.W., 01 1221, 01 1228; NW. side, first reef crest, 22.x. 1973, C.W., 011214. Parley Reef, patch reef in lagoon, 3 m, 22.iii.!973, C.W., GI1218. Heron Island: W. side, outer reef flat, 7.vii.l973, C.W., 011209; W. side, reef flat, .luly 1973, C.W., 011205; W. side, reef slope, 7 m, 7.vii.l973, Y. Loya, 011203. Palm Islands, 1939, T. C. Marshall, 011206-8. Field Diagnosis Corymbose to tabular colonies, the branchlets covered evenly and neatly with nariform corallites with elongate openings. Colour is most commonly cream to pale brown, usually with pale blue tips, but can also be yellowish or greenish. Laboratory Diagnosis Branching pattern: Branches arise from an encrusting plate, or a sturdy stalk. Secondary branches are vertical in the centre of the colony, and more curved towards the edges; they are not usually very proliferous. Axial corallites: Outer diameter 2 0 to 3*0 mm; inner diameter 0-5 to 0-9 mm. Septation: primary septa present, up to 3/4R, secondary septa anything from all absent to all present up to ' 1/4R. Radial corallites: Nariform, dimidate, or tubo-nariform, opening at 90" to branch or less. The walls of the corallites may be thickened. Primary septa present up to 1/3R, secondary septa poorly developed, usually less than a full cycle present, up to 1/4R. Coenosteum: Laterally flattened or forked spines are arranged densely on radial corallites, sometimes joining as costae. Between radials coenosteum is reticulate. Within Reef Distribution Occurs on most parts of the reef flat, reef crest, upper reef slope (to about 8 m in studied areas). This is sometimes the only Acropora (other than A. palifera) occurring in shallow lagoonal patch reefs in inner lagoons adjacent to coral cays. Identification Difficulties and History The close similarity of A. nasuta and A. cymhicyathus has been commented on by other authors. Although not well represented in the literature, this is a common Acropora on the Great Barrier Reef, occurring in a variety of habitats. Colonies are often very regularly formed, and easily recognised, although it can sometimes be confused with more regular colonies of A. cerealis. It is a good photographic subject, usually being identified in popular texts as ‘A. surculosa'. Geographic Distribution Indo-Pacific: Ceylon, Great Barrier Reef, Fiji Islands, Samoa, Marshall Islands, Tahiti. Acropora diversa (Brook, 1891) (Plates 79, 80A, B) Madrepora diversa Brook. 1891. p.46I; 1893, p.l41, pi. 16, fig.B. Acropora diversa: Wells, 1954, p.424, pl.l 17, figs. 3-6; Stephenson and Wells, 1956. p.l3. Madrepora concinna Brook. 1891, p.460; 1893, p.I65, pi. 17. Acropora otteri Crossland, 1952, p.229, pl.43, figs.l, 2, pl.44, figsJ, 2. Material Examined BM: Diego Garcia, G. C. Bourne, A. diversa holotype 1891.4.9.4; Mauritius, A. concinna syntypes 1878.2.4.3, 1878.2.4.8; Great Barrier Reef Expedition, A, otteri syntypes 1934.5.14.17; 1934.5.14.76; June Reef, outer moat A. otteri (? also a syntype) 1934.5.14.315. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 299 QM: Big Broadhurst Reef: SW. side, outer reef flat* 22.X.1976, C.W.. GM232. GI1233; 18.x 1973, C.W., G1 1261, Gl 1263, G1 1269, Gn279, 011280; 22,x.l973, C.W., G1I272, G11285; SW. side, reef slope: 1-3 m, 22.X.1973, C.W., G11245; 1-5 m. 18.x. 1973, C.W., G11266; 3 m, 27.iii.l973, C.W., 011254; 3-3 m. 22.X.1973, C.W., 011237; 4-5 m, 23.X.1973. C.W., G1 1273; 51 m, 23.x. 1973, C.W., G11236, G11270; 5-7 m, 23.x. 1973, C.W., G1I241; 6 m. 27.iii.1973, C.W., 011252; 7 m, 28.iii.1973, C.W., 0112.59; 7-1 m, 13.X.1973, C.W.. 011276; 7-8 m, C.W., 011267; 8 m, 28.iii.1973, C.W., 011246, 011258; 8-1 m, i4.x.l973, C.W., 011231; 8-2 m, 14.x. 1973, C.W., 011230, 01 1243, G1 1285; 8-6 m. 15.x. 1973, C.W-, 01 1240; 8-7 m, 15.X.1973. C.W., 01 1264; 9-6 m, C.W., 01 1265; 9-7 m. 15.X.1973, C.W.. 011271; 9-8 m, 16.x. 1973, C.W., 011239; 10-6 m, 23.x. 1973, C.W.. 011275; 12-5 m, I7.X.1973, C.W., 011242; 14-2 m, 23.X.1973, C.W., 01 1235; SW. side, reef crest; 1-5 m, 11.x. 1973. C.W., 011278; 1-6 m. 22.X.1973, C.W., 011284; 22.x 1973. C.W., 011234. Bowden Reef, SW. side, reef crest; 0-75 m, 1 5.vii.l972, R. Pearson, Gl 1 268; 1 m, 15.vii.l972, C.W., 011248, G11251, 011260; SW. side, reef slope: 26.vii.1973. C.W., 011238. Bushy-Redbill Reef, NW. side, reef slope: Jan. 1973, C-W , 011249; 19.xii.l972, E. Lovell. 01 1257; GI 1262; 011282. Darley Reef, patch reef in lagoon: 1 m, 24.iii.1973, C.W., G11250; 5 m, 19.vii.l972, C.W., 011244, 011256; 24.iii.1973, C.W., 011253, 011255. Field Diagnosis Corymbose, caespito-corymbose, tabulate to plate-like colonies have a mixture of tall and short tubular radial corallites which is usually distinctive. A variety of colours occur, vis. cream, yellow-brown, blue-grey, blue-brown. Laboratory Diagnosis Branching pattern: From an attachment region which is central to lateral, branches are given off upwards vertically to obliquely. These may be as short as 20 mm to as long as 70 mm, and may branch proliferously or rarely. Branch widths vary from 7 to 20 mm. Axial corallites: 1 to 3 mm exert. Outer diameter 2-4 to 3-2 mm (slightly larger in A. diversa type); inner diameter 0-8 to 1-1 mm. Septation: both cycles usually present, the first cycle up to 3/4R, the second up to 1/3R. Radial corallites: Tall and short corallites are mixed, in a pattern which may be regular (tall radials arranged in vertical rows) or very irregular. The form of the tall and short radials is similar — tubular, with round, or slightly oval openings, sometimes tubo-nariform. The outer wall may be thickened. The septal development is usually poor, primary septa being present, up to 1/3R, but usually less than this, the secondary cycle usually only partly present, up to 1/4R. Coenosteum: Simple pointed to laterally flattened spines are densely arranged on the radial corallites, sometimes forming costae. Between the radials their arrangment is less dense. Within Reef Distribution Outer reef flat, reef crest and reef slope to about 15 m, lagoonal patch reefs. Identification Difficulties and History Amongst my material there is great variability in radial corallite features such as length, shape of opening, ratio of short to long, as well as the expected colony-shape variability. Both in the field and in the laboratory I find areas of overlap and difficulty in distinguishing some specimens from A. nasuta and others from A. variabilis. A study concentrating on these three species would be profitable. Geographic Distribution indo-Pacific: Mauritius, Diego Garcia, Great Barrier Reef, Marshall Islands. Acropora variabilis (Klunzinger, 1879) (Plate 80C, D) Madrepora variabilis Klunzinger, 1879, p.17, pl.l, fig.lO, pl.2. figs.I, 5, pi. 5, figs.l, 3, pl.9, fig.l4; Brook, 1893, p.l61. Acropora variabilis: von Marcnzeller, 1907, p.49, pi. 15, figs. 4B44; Vaughan, 1918, p.l81, pi. 80. figs.2. 3, 3a, 3b; Faustino, 1927, p.276; Wells, 1950, p.38; 1954, p.428, pi. 128, figs.l, 2, pi. 130, figs. 1, 2; Rossi, 1954, p.52; Stephenson and Wells, 1956, p.l9; Scheer and Pillai, 1974, p.23, pi. 8, fig-2; Pillai and Scheer, 1976, p.31. Acropora variabilis var. pachyclados: Crossland, 1952, p.222, pi 38, figs.l, 6. Matf.riai Examined MNB: Koseir, Klunzinger, A. variabilis var. pachyclados 2118; Koseir, Klunzinger, A. variabilis var. cespiiofoliata 2120 (Klunzinger mentioned specimens, examined as photographs only). QM- Big Broadhurst Reef, SW. side, outer reef flat: 11.X.I973, C W.. GII291; 18.x 1973, C.W.. 011287-90. Field Diagnosis Colonies may be small round clumps of vertical to oblique branches or stalked corymbose colonies to about 30 cm across. Branches are covered by appressed tubular corallites with round openings. The sizes of the radial corallites may be similar or extremely variable. Colour may be brown. 300 MEMOIRS OF THE QUEENSLAND MUSEUM lavender-brown, greenish-brown, or yellow or cream with purple corallites. Laboratory Diagnosis As this species has been well described by other authors, and my series is poorly representative of the species, further description is not given. Refer to Vaughan 1918 for tables of measurements of skeletal features. Within Reef Distribution Shallow outer reef flat (particularly where an elevated platform is present), reef crest. A large series in the JCU comes from the fringing reefs of the Palm Islands and Lizard Island. Identification Difficulties and History Although this species is poorly represented in my areas of study, evidence suggests it may be common (a) on windward outer reef flat platforms (as small round clumps) and (b) on reef edges of the fringing reefs of continental islands (as caespito-corymbose colonies). The series in JCU from Lizard Island and the Palm Island group is in the second category. An allied species is A, valida (Dana) (see Hoffmeister, 1925, p.6()). This latter species may also occur on the Great Barrier Reef, but is not represented in the QM collections, except by two specimens collected in the Fiji Islands, and closely comparable with Dana's type. Geographic Distribution Indo-Pacific' Red Sea, Nicobar Islands, Cocos- Keeling, Philippines, Great Barrier Reef, Marshall Islands. Acropora hiimilis (Dana, 1846) (Plates 81, 82, 83) Modrepora humilis Dana, 1846, p.483, pl.31, fig. 4, pi. 41, fig. 4. Acropora humilis: Wells, 1954, p.425, pi. 1 00, fig. 1. pLl26, figs.l^, pi. 1 27, figs.3, 4, pL128. figs. 3-5 (synonymy), Rossi, 1954. p 50; Slephen.son and Wells, 1956, p.l5; Pillai and Scheer, 1976, p.32. This species has been given extensive taxonomic treatment by Wells 1954. He combined 17 species (some with additional synonyms designated by earlier authors), and concluded that three broad forms could be recognised. These he considered to be characteristic of different reef localities and related, in particular, to water level. Although a large suite of specimens is on hand at the QM, their description is withheld until a further study concentrating on this complex species can be carried out. For general purposes, this species is probably the best known and most easily recognised Acropora. Some additional informa- tion is added below on Well’s first facies, forma alpha {A. samoensis, A pelewensisj. Acropora humilis (Dana, 1846) forma a Wells, 1954 MatfriaI- Examined QM‘ Big Broadhurst Reef, SW. side, reef slope: 8 6 m, 14.X.1973, C.W , G1 132, Gl 135; 8-6 m, 15.X.1973, C.W., 011131; 8-7 m. 15.X.1973, 011130; 9-8 m, 16.x. 1973, G11134; 10-4 m, 16.x, 1973, C.W.. G11133; 12-5 m, I7.X.I973. C.W., G11I37. Bowden Reef, SW side, upper reef slope, 26.vii.1972, C.W., G11I89. Darlev Reef, patch reef in lagoon: 24.iii.l973, C.W.. Gl 1 128; 3 m, 22.iii.l973, C.W., GI 1129; 22.iii.1973, 5 m, C.W., GI16IL Field Diagnosis Colonies may be low arborescent (with shrub-like growth) caespito-corymbose. or plate- like. Branches are terete (non tapering) or slightly tapering, with wide axial corallites, and large thick-walled radial corallites. Colours are cream-brown, sometimes with pale blue tips, or pale blue or lavender. LABORArORY DIAGNOSIS Branching pattern: Superimposed on the colony shape described above is usually a high degree of budding. The branches may be as broad as 20 mm, but in the plate like colonies (which occur on reef slopes) and particularly in deeper-water specimens, branches can be as narrow as 8 mm. Axial corallites: About 2 mm exert. Outer diameter 3-0 to 5 0 mm; inner diameter 11 to 1-6 mm. Septation: both cycles present, primaries up to 3/4R, secondaries up to 1 /2R. Radial corallites: Tubular or appressed tubular, with round or oval openings, or nariform, the outer wall thickened. Primary septa are usually present, up to 1 /3R; secondary septa not usually fully developed, but some present up to 1/4R. Coenosteum: Costale to reticulate both on and between radial corallites, with spines that are laterally flattened and sometimes slightly elaborated at the tip. Within Reef Distribution This form extends down the reef slope and sloping surfaces of lagoonal patch reefs. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 301 Identification Difficulties and History Deep water specimens, because of the narrow and proliferous branches, are barely recognisable as A. humilis. and begin to have a similar general appearance to A. sarmenlosa. Geographic Distribution A. humilis has a wide Indo-Pacific distribution. Acropora digitifera (Dana, 1846) (Plate 84) Madrepora digitifera Dana, 1846, p.454; Brook, 1893, p.75. Acropora digitifera: Verrill, 1902, p.228, pi. 36, fig. 12, pl.36B, fig.3; Vaughan, 1918, p 175, pi. 13, fig, 7; pi. 76, figs. I, la, 2; Wells, 1954, p-427, pi. 127, figs.l, 2; 1955. p.9; Stephenson and Wells, 19^6, p.l3 (synonymy). Acropora fraierna Verrill, 1902. p.247, pi 36, fig. 18, pi 36B, fig.9. Materia! Examined YPM: A. digitifera type (fragment, ex. Boston Soc. Nat. History) 4192; Tahiti, U.S. Expl. Exped., A fraierna holotype 2032. QM: Big Broadhurst Reef, SW. side, outer reef flat: lLx.1973, C.W., G11170; 18.X.I973, C.W., GI1169, G11171, Gni72; 22.X.I973. C.W., G11168, 011187. Bushy-Redbill Reef, adjacent Redbill Is., reef flat, I8.xii.I972, C.W., GII176; 2.vi.l975, C.W G11173, GI1I74; 5.vi.l975, C.W., 011175; 10.vi.]975, C.W., G11188; Il.vi.l975, C.W., G11177. Heron Island, W. side, reef flat. July 1973, C.W., G11180. Oyster Cay, C. Limpus, 1975, G11179 Fiji Islands: Great Astrolabe Reefs, Yaucuve Leva fringing reef, Jan. 1974, C.W., G11183, G11184, G1II86; Vaga Bay, Beqa, patch reefs, 15. i. 1974, C. J. Wallace, G1 1 182. Field Diagnosis Colonies are corymbose, with central to lateral attachment, and tapering branches. Appearance is of a diminutive, ‘neat’ A. humilis. Colour is commonly cream to pale brown, with or without blue branch tips. Laboratory Diagnosis Branching pattern; From a growing area which may be central to lateral, and more or less stalk-like, main branches grow horizontally, and give off secondary branches or bundles of branches vertically to obliquely. These may taper to a point, or be terete, and are from 8 to 20 mm greatest width Axial corallites: Non exert. Outer diameter 2-8 to 3-8 mm; inner diameter 0-8 to 11 mm. Septation: both cycles developed, primaries up to 2/3R, secondaries up to 1/4R. Radial corallites: A size gradation occurs from branch tip to proximal and small corallites are interspersed with large. Shape is dimidiate, or tubular with oval to dimidiate opening, radials spreading at 90° from branch. Outer wall is thickened. Coenosteum: Costate or with regular lines of spines on corallites, spongy with spines in between. Within Reef Distribution Limited to the shallow middle reef and outer reef flat pavement (‘A. digitifera' from inner reef flat as interpreted by Wallace and Lovell 1977 is A. tubicinaria.) Identification Difficulties and History On some reefs (e.g. Enewetak, Marshall Islands) this species is well marked and easily recognised. On the Great Barrier Reef, where A. digitifera occurs with A. humilis. some colonies cannot be definitely assigned to one or other species on morphological grounds. As Stephenson and Wells (1956) point out, the main difference is in dimensions. A further analysis of one must include the other Geographic Distribution Great Barrier Reef, Moreton Bay (Queens- land), Fiji Islands, Marshall Islands, Tahiti. Acropora multiacuta Nemenzo, 1967 (Plate 85) Acropora multiacuta Nemenzo, 1967, p.l33, pl.39, figs.l, 2. 3; Scheer and Pillai 1974, p 24, pi. 6, fig-4. Material Examined USNM: (donated): Darley Reef, shallow patch reefs in lagoon, 1 m, 24.iii.l973, C.W. BM: (donated): Darley Reef, shallow patch reefs in lagoon, I m, 24.iii.l973, C.W. JCU: Philippine Islands, 1975: 3 m, M. Pichon. 2509/75; 5 m, M. Pichon, 2525/75. QM: Darley Reef, shallow patch reefs in lagoon: 1 m, 19.vii.l972, C.W., G6721, G6722; upper surface, 23.iii.l973, C.W., G10465, G10470, G10471: upper surface, 24.iii.I973, C.W., G10466, GI0467. GI0469; 1 m, 24.iii.I973, C.W., G10464; 3 m, 22.iii.1973, C.W., G 10468. 302 MEMOIRS OF THE QUEENSLAND MUSEUM Field Diagnosis Irregular caespitose to caespito-corymbose colonics from an encrusting central to lateral base, with axial corallites sturdy and prominent, sometimes to the extent of the entire branch or one side of the branch being naked of radial corallites. Colour is whitish blue with pale blue polyps. The largest colony seen to date is 19*5 cm diameter. Laboratory Diagnosis Only twelve Great Barrier Reef specimens have been examined and these show much variation, particularly in the extent of radial corallite presence on the branches, the amount of secondary branching, and the length of the main branches. Some dimensions such as the width of the branches are not included in the diagnosis for this reason. Branching pattern; From the encrusting base, the main branches arise vertically to obliquely, and may be variously curved. The longest branch in the present collection is 10 cm. At the base of main branches smaller branches may occur. These do not always alter the colony shape, but in the side attached specimens it appears they would contribute to the development of a bracket shape as the colony matured. Some specimens have prolific incipient branching along the main and secondary branches. Axial corallites: (On the main branches) outer diameter 3-5 to 6-5 mm; inner diameter 10 to 2-0 mm. Seplalion; both cycles present, occasionally a third cycle partially developed, primaries up to 1/3R, secondaries up to 1 /4R. On the basal and incipient branches axials arc narrower and sometimes compressed so that the opening is oval. ThCvSe are described as radial corallites by Nemenzo. Radial corallites: Scattered, nariform, tubo- nariform, or partly appressed tubular; often oriented with opening down or across the branch. Septa not developed at all, or primaries just visible. Coenosteum: Densely echinulate on both radial corallites and inter-corallite areas; the spines laterally flattened, occasionally some pseudocos- tatc development on the axial corallite. Within Reef Distribution The only populations seen have been on the upper surface and edges of patch reefs, just below low water. The colonies occur on the surface or in depressions in the irregular reef surface, and the main branches may be curved to maintain an overall vertical orientation. The naked areas of branches are always upwards. Identification Difficulties and Notes Nemenzo used the species grouping Alticyathus for this species and for his A. fastigata (1967, p.l34). The latter may be a synonym, or may be A. digitifera. I have not been able to locate Nemenzo’s types in the UP collections. On the Great Barrier Reef, the species has only been seen from the Darley Reef lagoon, and it cannot yet be fully categorised. Geographic Distribution Indo-Pacific: Nicobar Islands, Philippines, Great Barrier Reef. Acropora clathrata (Brook, 1891) (Plates 86, 64C). Madrepora clathrata Brook, 1891, p.459; 1893, p.49, pi. 5, pl.6, Hg.A, B. Madrepora orbicularis Brook, 1892, p.460; 1893, p.37, pi. 2 (synonymy); Madrepora vasiformis Brook, 1893, p.37, pi. 26, fig. A (Synonymy). Acropora vasiformis: Pillai and Scheer, 1976, p.27, pi. 3, fig. 4. Acropora tutuilensis (part) Hoffmeister, 1925, p.71, plate 19, figs, la-le. Material Examined BM: Mauritiu.s, A. clathrata holotype 1893.4.7.78; Ceylon, A. orbicularis holotype 1883.3.24.7; Rodriguez, A. vasiformis holotype 1876.5.5.92. USNM: Pago Pago Harbour, Tutuila, Samoa, A. tutuilensis no. 1; ?nos. 2, 3. (Mayor collection) (mentioned Hoffmeister, 1925). QM: Big Broadhurst Reef, SW. side, reef slope: 5 m, 25.111.1973, C.W., G9753; 8 m, 26.iii.1973, C.W., G9752; I m, 27.iii.1973, C.W„ G9748; 3 m, 27.iii.I973. C.W., G9749; 4 m, 27.iii.I973, C.W., G975I; 5 m, 27.111.1973, C.W., G9750: 1 m, 13.x. 1973, C.W., G9742, G9746; 6-3 m, 13.x. 1973, C.W., G9735; 7 m, 13.X.1976. C.W., G9743; 71 m, 13.X.1973, C.W.. G9740, G9741; 7-3 m, 14.X.1973, C.W., G9739; 8 m, I4.X.1973, C.W., G9744, 8-1 m, 14.X.1973, C.W., G9738; 19-7 m, 15.X.1973, C.W., G9737. 4 m. 20.X.I973, C.W., G9745; 10 m, 20.x. 1973, C.W., G9759; 1*9 m, 22.X.1973, C.W., G9747; 3*3 m, 22.X.1973, C.W., G9736. Bowden Reef: slope of opening in SW. side: 0-5 m, 26.vii.1972. C.W., G9760; I m, 26.vii.1972, C.W., G9761; SW. side, reef slope, 1 m, 15.vii.l972, C.W., G9762. Bushy-Rcdbill Reef: NW. side, reef slope, Dec. 1972, C.W., G9754; NNE. end, 30.xii.l972, C.J.W., G9755; S. end. reef slope, 3 m, 3.vi.l975, C.W., G9756; reef WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 303 slope adjacent Redbill Is., 0-5 m, l.vi.l975, C.W., G9758; 1 m, 2.V.1975, C.W., G9757. Parley Reef: patch reef in lagoon, 3 m, 18.vii.l972, C.W., G9763. Field Diagnosis Horizontal plate (reaching approx. 1-2 m diameter), developing by unilateral expansion from a vasiform shape with a single attachment, which becomes lateral as the colony develops. The plate has a flat appearance, due to the orientation of the secondary branchlels in the plane of the plate or obliquely to it. Common colours are pinkish brown, apple green and yellowish green. Laboratory Diagnosis Branching pattern; From the attaching stalk, branching is oblique to horizontal. Secondary branches arise in the same plane as primary branches, or obliquely to them, forming a single layer or a series of interlacing layers. All branches, except those towards the edge zone, are of similar diameter. Anastomosis of branches varies, so that anything from an open network to a solid plate (with branches either fused to the plate or projecting obliquely from it) is possible. Axial corallites: Outer diameter 1-5 to 2*2 mm, inner diameter 0-6 to 0-9 mm. Septation: primary septa present, up to 1 /3R, secondaries usually not visible, or a few present to less than 1/4R. Radial corallites: A number of shapes are possible, and specimens may possess all, some, or only one of the possible types, viz. tubular, with round, oval, or dimidiate openings, tubo-nariform, nariform, rostrato-nariform, dimidiate, sub- immersed or immersed. Immersed corallites usually only occur along lines of fusion of branches. The other types project from the branch, at from 90° to 45°, occasionally less. There is a tendency for differential thickening or extending of the longer wall sometimes with bizarre extensions such as long points on nariform and horns on dimidiate corallites. Plate 86 shows: (D) all uniform size, nariform or dimidiate with differential outer wall development. (B) mixed size, tubular with round, oval, and dimidiate openings, immersed and sub-immersed. (F) similar to B, but corallites more appressed and scattered. Septation: septa are never well developed; usually only the directive septa are visible, or a few other primaries present as small spines. As the corallites are rarely appressed, they appear well spaced and there is always some coenosteum between. Coenosteum; On unthickened corallites this may be visibly porous and is usually costate, on thickened radials it is composed of compactly arranged flattened spines, sometimes pseudocos- tate. Between corallites it is reticulate. Identification Difficulties and History This coral occurs with the other large horizontal plate Acropora {A. hyacinthus and A. cytherea), from which it is easily separated by its flattened appearance and by the lack of scale-like corallite lips. Identification difficulties occur at the laboratory stage, where so many different combinations of colony features and corallite size and shape are seen. This variety is demonstrated in the synonymy: A. clathrata s.s. has narrow branches, openly reticulated, and radial corallites are tubular, with a variety of sizes and opening-shapes; 'A. orbicularis’ and ‘A. vasifor- mis’ are almost solid plates, ‘A. vasiformis' with rostrate thickening of many radial corallites, A. orbicularis’ with unthickened radials in a variety of shapes. Hoffmeister’s no. 1. specimen of A. tutuilensis is a partly fused plate of this species. His no. 4 is probably A. rotumana. All the type specimens can be compared with specimens in the QM series. A. stigmataria (Milne Edwards and Haime) (see Brook 1893, p.5U), may be a senior synonym but its type cannot be located for examination. Within Reef Distribution The species appears to be restricted to sloping surfaces or good water cover. Very large specimens can occur around low water mark on the edge of surge channels, on the upper reef slope, and on the floor of surge channel openings (see Plate 64C). Geographic Distribution Indo-Pacific: Mauritius, Rodriguez, la Reunion (G. Faure pers. comm.), Ceylon, Seychelles (?), Great Barrier Reef, Samoa. Acropora divaricata (Dana. 1846) (Plates 87, 88) Madrepora divaricata Dana, 1846, p.477, pl.41, fig. 2; Milne Edwards and Haime, 1860, p.l40; Brook, 1893, p.64. Madrepora tenuispicata Studer, 1880, p.20, figs. la, lb; Brook, 1893, p.96. 304 MEMOIRS OF THE QUEENSLAND MUSEUM Acropora lenuispicata: Piliai and Scheer, 1974, p.455 fig. 4b. '1 S-iadrepora complanata Brook, 1891 p 459- 1893 p.70, pl.4, fig.C. Acropora complanata. Pillai and Scheer, 1976. p.28. pi. 7. fig,2, ?Madrepora complanata var. informis Brook 1893 p.7L Material Examined USNM: Fiji Islands, U.S. Expl. Exped., A. divaricaia holotype 299. YPM: Fiji Islands, U.S. Expl. Exped., A. divaricata fragment of type 2008. BM: Seychelles, H M.S. Alert, A. complanata syntypes 1882.10.17.140, .147, .148; Macclesfield Bank, 13 fathoms, A complanata var. informis syntypes 1892.10.17.71, .72, .73. Hessisches Landmuseum: Acropora tenuispicata (colour transparencies only) (mentioned Pillai and Scheer 1974). QM; Big Broadhurst Reef: SW. side, reef slope; 12-4 ni, 23.x. 1973, C.W.. G9I62; II m, 16,x.l973, C.W., G9166; 9-8 rn, 16.X.1973, C.W.. G9167; 9-6 m, 15.x. 1973, C.W., G9164; S-7 m, 15.x. 1973, CW, G9170. G9I73; 8-6 m, I5.X.1973, C.W., G9I74, G9175; 83 m, 15.x. 1973, C.W., G9I63, G9172; 81 m] I4.X 1973, C.W., G9161. G9168; 8 m, 14 x 1973, C.W., G9160. G9I65. 09178; 7-8 m, 14.X.1973, C W., G917U G9I76, G9177; 7-6 m, 28.iii.1973, C.W., G9180; 7 1 m, 13 X.1973, C.W., G9169; 7 m, 28.iii.1973, C.W., G9181, G9188, 6 m, 26.iii,1963, C.W., G9I79; 6 m, 25 iii 1973 C.W . G9184. Bushy-Redbii! Reef: W. side,, reef slope; 8 in. 3.viT975, C.W., G9186; 2 m, 3.vi.l975, C.W., G9185; NW. side, reef patches, 12 m, I4.vi.l975. CW G9187. Darley Reef: patch reefs in lagoon; 7 m, 22.iii.1973, C.W, G9182; 3 m, 18.vii.l972, C.W., G9I84- 22.iii.l973, C.W., G10219. Fiji Islands, Great Astrolabe Reefs (Kadavu), W. side of Yaukuve Levu, fringing reef, l.ii,1974, C.W., G9781 G9782. Fiflu Diagnosis Occurs as bracket like colonies with central to lateral attachment, ‘corymbose’ in having all branches reaching up to a horizontal plane, and ‘caespitose’ in having divaricate branching within the boundaries of the colony shape. Reaches approximately 50 cm diameter. Branches appear rough because of projecting radial corallites. Colour is usually a drab dark brown or dark brown with blue tips. Laboratory Diagnosis Although the species is easily recognised in the field, skeletal fragments can be confusing. Branchlet dimensions and radial corallite shape are variable amongst colonies in a single population, and pieces taken from different parts ol the same corallum may appear different because of their orientation. Branching pattern; From a single area of attachment, branching is central to lateral. Periferal branches contribute to an oblique undersurface; inside these is a network of short branchlets at wide angles, the final branchlets being erect or nearly so, and ending in a horizontal plane. Branchlet widths vary from 7 rnm to 15 mm. Axial corallites; Outer diameter 2-3 to 3 0 mm; inner diameter 0-8 to 1-1 mm. Septation: primary septa present, up to 1/2R, secondary septa poorly developed, but usually some present, up to 1/4R. Radial corallites: Shape and size of the radial corallites changes along the branches. On upper branchlets they are prominent (up to 3 mm long), usually extending at from 45^^ to 90°. They are usually tubular on branch tips, passing through tubo-nariform to nariforin, then rounded to sub-immersed proxirnally. The prominent radials are sometimes extended by rostrate development (see Plate 88 C, D). Within the sequence from distal to proximal, radials are usually evenly graded and neatly arranged, but they can be unevenly graded, so that branches appear ragged (Plate 88 A), and downward directed radials can occur anywhere along the branch. Within Reef Distribution This coral occurs in the higher diversity (for Acropora) parts of the reef, where no particular colony shape predominates: middle reef slope, deeper outer reef Oats, patch reefs in deep lagoons and leeward broken reef areas. The species, although common, has a drab appearance when alive, and is easily missed when subjective observation techniques are used. Identification Difficulties and History 1 he radial corallites of this species are similar to those of the flat plate species, Acropora clathrata- I have not found specimens Acropora divaricata deep enough to form flat plates but am confident that A. complanata (Brook), a flat plate species dredged from deep water, represents deep water flattening of A divaricata. For A. tenuispicata (Studer) I have drawn on Pillai and Scheer’s (1974) interpretation of this species. The type is not in the MNB, where the remainder of the Studer types are located. The combination of a determinate bracket with divaricate branching WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 305 is unique in the Acropora. The shape has not been emphasised in previous descriptions. A series in the JCLI from Lizard Island fringing reefs shows more rounded colony shape, with slender branches, often naked of radial corallites on their upper surface. Geographic Distribution I ndo- Pacific: Seychelles, Singapore, Fiji Is- lands, Great Barrier Reef. Acropora sarnientosa (Brook, 1892) (Plate 89) Madrepora sarmentosa Brook, 1892, p.462; 1893, p.I27, pi. 22 Acropora sarmentosa: Nemenzo, 1967, p.90, pi. 26, fig-4. non Acropora sarmentosa: Vaughan, 1918, p. 174, pi. 72, rig.4, pi. 73, fig.l. Acropora rosaria form I: Crossland, 1952, p.224, pl.40, fig.3. Material Examined BM: Port Denison, Saville Kent, A. sarmentosa syntype 1892.6.8.228. UP: Little Balatero Cove, Puerto Gaelera, Oriental Mindoro, Nemenzo 259; Muelle, Puerto Gaelera, Oriental Mindoro, Nemenzo 324 (mentioned Nemenzo, 1967). QM: Big Broadhurst Reef. SW. side, reef slope: 17m, 24.X.1973, C.W., G9064, G9065; I5-7m. 23.X.I973, C.W., G9062; 15-4m, I7.X.1973, C.W.. G9056; 9-7m, I5.X.1973, C.W., G9059; 8 Im, 14.x. 1973. C.W., G9055; 8m, 28.iii.l973, C.W., G9054; 7-6m, 13 x. 1973, C.W., G9057; 6-8m, 20.x. 1973, C.W., G9058; 41m, 22.x. 1973, C.W., G9061. Bowden Reef, slope of opening in SW. side, l-2m, 23.viii.1972, C.W., G9066. Bushy-Redbill Reef, NW. side, reef slope: 30.xii.l972, E. Lovell, G9071; 3-7m, 2l.xii.l972, E. Lovell, G9073; NW. side, reef crest, 30.xii.l972. C.W., G9070; NW side, patch reefs; 12m, 15, vi. 1975, C.W., G9075; 12m, 14.vi.l975, C.W., G9060: SW. side, reef crest, 24.V.I975, C.W , G9063; outer reef flat adjacent Redbill Island- Lvi.1975, C.W., G9072; 22.xii.l972, E. Lovell, G9074. Darley Reef, patch reefs in lagoon. 6m, 22.iii.1973, C.W., G9067; 4m, 22.iii.1973, C.W., G9068; 2.5m, 24.iii.1973, C.W., G9069. Heron Island, W. side, reef flat, 6.vii.l973, Y. Loya, G9704. Ellison Reef: dredged 9 fathoms, 25.vii 1924, Dr Paradice, G9709; 24.vii.1924, C. Hedley, G9707. Fiji Islands, Great Astrolabe Reefs: Qasilabe fringing reef, 4^6m. 4.ii.l974, C.W., G97I0; Yaukuve Levu, W. side, fringing reefs, 8m, l.ii.l974, C.W., G9705. Field Diagnosis Colonies of this species usually have few (commonly 2 or 3) thick rounded branching units, consisting of a central horizontal to oblique branch or branches, with vertical branchlet bundles evenly distributed, but longer on the upper side. Colonies of more than 50 cm across are unusual, and attachment is usually from the side. The overall appearance is smooth — axial corallites are not exert, radial corallites are large, evenly distributed, uniformly sized and not projecting; secondary branching patterns are regular and branchleis terete. The usual colouration on the Barrier Reef is two-toned, and cryptic — most often a dull greenish grey or -brown, with pale brown or pink tips to the branchleis. This colouration occurs also in the Fijian reefs. On patch reefs in deep lagoonal situations, the colony can assume a sturdy rounded shape with central attachment. Laboratory Diagnosis Branching pattern; From a side attachment, branching is horizontal or oblique, with two to several mam branches. Vertical to acute branchleis occur at regular intervals along the main branches; these usually branch again, one to several limes, and are shorter and narrower on the under surface. Upper surface branchleis are 5 to 9 mm wide. Axial corallites: Outer diameter 3 0 to 4 0 mm, up to 7 mm in lagoonal specimens; inner diameter 10 to 2-0 mm. Septation: 12 septa or slightly less, usually well developed (primaries to 3/4R, secondaries to 1/2R)- Radial corallites: All of similar size, neatly and evenly arranged around the branchlet, appressed tubular. (Brook described them as ‘swallow-nest shaped’). In shallow water specimens the wall is thick, and corallites are densely packed around the branchlet. With increasing depth, radial corallites become more scattered, thinner walled, and the wall may flare a little, and appear lip like. Coenosleuni: Spines which may be laterally flattened or slightly elaborated are arranged evenly both on radials and between: sometimes radial walls are costate. Within Reef Distribution Reef slope from crest to limits of depth of Acropora distribution (species no. 18 in Wallace 1975); deeper water diverse reef flats and lagoonal patch reefs; sporadically in shallow outer reef situations where there is space for outward growth; fringing reefs. The species is never very 306 MEMOIRS OF THE QUEENSLAND MUSEUM abundant, but is usually present in most reef habitats. Identification Difficulties and History In the field, lower reef slope specimens of this species and of A.Jlohda may appear similar. The two-toned colouration, thicker branchiets and smoother general appearance of A. sarmentosa can distinguish it. Vaughan (1918) suggested that A. sarmentosa may be a growth form of A. squamosa (= A. milleporaj. His specimens were not A. sarmentosa but were indeed ‘A. squamosa’. His misidentificalion of this species may account for it being overlooked in much of the literature. Both Brook and Nemenzo describe their specimens well. Geographic Distribution Indo-Pacific: Philippines, Great Barrier Reef, Fiji Islands. Acropora florida (Dana, 1846) (Plates 90, 91, 92) Madrepora florida Dana, 1846, p.466, pi. 37, fig.], non Madrepora florida: Brook, 1893, p.53. Madrepora gra\>ida Dana. 1846, p.470. Brook, 1893, p.59 (synonymy). Acropora gravida: Neinen/o, 1967, p.l07, pi. 31, fig. 3; Scheer and Pillai, 1974, p.l8, pl.5, fig.l; Pillai and Scheer, 1974, p.453. Madrepora mirabilis Quelch, 1 886, p. 1 59, pi. 1 0, fig. 5; Brook, 1893, p.i25. Madrepora cornpressa Bassett-Srnith, 1890, p.452; Brook, 1893, p.60, pi. 33, fig.F. Madrepora affinis Brook, 1893, p.60, pi. 28, fig.F (synonymy). Acropora affinis: Crossland, 1952, p.205, pi. 34, fig.l; Nemenzo, 1967, p.77. Acropora vermiculata Nemenzo, 1967, p 108 pi 31 fig.4. ■ ’ Material Examined USNM: Fiji Islands, U.S. Expl. Exped., A. florida holotype 282. YPM: Fiji Islands. U.S. Expl. Exped., A. florida fragment of type 2002. BM: Tizard Bank, China Sea, 5 fthms, A. cornpressa holotype 1889.9.24.117; Darnley Island, J.B. Jukes, A. affinis syntype 1846.7.30.29; Banda, Challenger, A. mirabilis holotype 1885.2.1.14; Claremont Is., G.B.R., Savilie Kent, A. ornata var. (id. Brook) 1892.6.8.112. QM: Big Broadhurst Reef: S W side, reef slope: 15-3ni, i7.x.l973, C.W., G8649; lL3m, 17.x, 1973, C.W., G8653; 9 3m, I6.X.I973, C.W., G8647; 9-3m, 17.x. 1973, C.W., G8646; 8-6m, 17.X.1973, C.W., G8b48, G8654; 8-6m. i6.x.l973. C W,, 08661; 8ni, 28.iii.l973, C.W., G8664; 7-9m, I5.X.I973, C.W„ 08650; 7-4m,’ 15.x. 1973, C W . G8656, G8662; 6 6m, 14 x. 1973, C W., (18655, G8657, G8667; 6m, 27.iii 1973, C W.. G8665; ' 6m, 25.iii.I973, C W., G8666; 5-8m. )3.x.I973, C.W..' i G865I; 5-2m, I3.X.I973. C.W., G8645; 3m. 25,iii.I973’ 1 C W., G8669; 2*3m. 1 Lx. 1975, (' W., G8644; 2m ! 25.iil.1973, C.W., G866I, 08663, G8668, G8670; S W. ■ side, surge channel, 8-5m, 20.x 1973, G8658, G8659; S ! W. side, outer reef fiat, 18.X.I973, G8652. ’ ^ Low Isles, June 1974, C. Limpus, G8672. Murray Islands (Maer Is.), 18.vii.l974, G. Ingram G867L Bowden Reef, S. end, reef slope, 3-3m 15vii 1972 C.W., G6726. Darley Reef, lagoonal patch reef, 3-5m, 19.vii 1972 C.W.. G6725. Viper Reef, lagoonal patch reef, l-5m, 16viil972 C.W G6724. Fiji Islands, Makaluvau Reef, outer reef flat 10.1,1974, C.W., G10273. Field Diagnosis This is cl sturdy open arborescent species, in which the surface of the branches is covered by short secondary branchiets. It is highly variable in two aspects, ihe shape of the colony and the density and prominence of the secondary ; branchiets. The colony shape changes with depth I and slope of the attaching surface. In shallow ' water and flat substrate it forms a rounded, open j colony with central attachment; on sloping j surfaces the attachment is more lateral, and the I branches lend to extend horizontally. With ! increasing depth the branches become flatter. The secondary branchiets vary from evenly distributed with even lengths to scattered and variously sized and may even be undeveloped on some branches. The most irregular colonies occur in fringing reef situations. Laboratory Diagnosis Branching pattern: Sturdy main branches divide sparsely to form an upright bush. The shape of the colony and the cross-sectional shape of the branch become Hatter with increasing water depth. The branches usually proliferate towards the branch lips into shorter branches. The surface of the branches is covered with short branchiets. Except on completely vertical branches the branchiets arc shorter on the under-surface (to absent on horizontal branches). Axial corallites: Outer diameter 2-0 to 3 0 mm; inner diameter 0 8 to 1-4 mm Septation; primary septa present, up lo 2/3R; secondary septa usually all developed, or at least 3 present, up to 1/2R. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 307 Radial corallites: Evenly sized and distributed, I appresscd tubular with round opening, with wall which may flare slightly. Septation: primary septa present, up to 1 /2R, some to all secondaries present, up to 1 /4R. Coenosteum; Costatc or reticulate on radial I corallites, reticulate in between, with scattered j simple spines. Within Reef Distribution ' Entire reef slope from reef crest to limits of Acropora growths; deep water lagoonal patch reefs, fringing reefs. Occasionally on middle or . outer reef flat. It is well known as an early I recolonizer on the fringing reefs near Townsville, N.Q. (D. Tarca, pers. comm.). ' Identification Difficulties and History ; The variability of the species is expressed in the I synonymy. Brook misunderstood A. florida: his specimens in the BM are possibly A. rotumana. I A. florida s.s. is atypical of the species as it appears on the Great Barrier Reef in that the corallites are larger and the branchlets poorly developed: G6725 is closest to this. The species is best described by A. affinis, which has evenly I sized and distributed branchlets; its similarity to A. gravida was apparent to Brook. A. cotnpressa is named from a flat plate specimen of this species. A. mirabilis is named for a single aberrant specimen which is probably this species under some environmental stress. Geographic Distribution Indo-Pacific: Singapore, Celebes, Philippines, Strait of Malacca, Nicobar Islands, Great Barrier Reef, Fiji Islands, Enewetak Atoll (C.W.). The ‘Acropora echinata’ Group I t The species Acropora echinata, A. suhglabra. ! A. carduus and A. longicyathus, with their ^ synonyms, have in common their shape, radial corallite structure and coenosteal structure. The growth forms are comparable within the group, and nothing exactly similar is found outside the group. The branches are commonly labelled "bottlebrushh secondary branchlets, or bundles of branchlets, are given off evenly around the main branches, giving a round brush like unit. The growth of a colony is indeterminate, the units being capable of openly branching or closely proliferating, giving a variety of form in collected specimens. The colonies tend to occur on rubble or sandy floors, and the bases of branch units are usually dead. If the colonies occur in deep sloping conditions, they are small and approach a flattened form. The radial corallites are round tubular appressed or partly appressed, tending to develop into axial corallites. Their numbers relative to the number of axials are lower than in most other groups. In all species, the number of radial corallites per axial decreases distally on the branch unit, so that long naked axials may occur around the base. The spines of the coenosteum have multiple tips, and the coenosteal appearance on and between radials is similar. The main differences among the species are in the corallite dimensions and the amount of elaboration of the coenosteal spines. These species occur only in the sheltered, deeper parts of reefs, where almost any Acropora species can survive. They appear to have poor differentia- tion of radial corallites from axials, and if this is accompanied by a similar lack of functional differentiation it may account for their lack of colonizing success. ■Acropora echinata (Dana, 1846) Madrepora echinata Dana, 1846, p.464, pi. 36, fig.l, la; Brook, 1893, p.I85 (synonymy). Acropora echinata: Vaughan, 1907, p.l58, pis. 49, 50; Wells, 1954, p.423, pi. 135, figs. 1-4, pi. 136, figs 1-6 (synonymy); Nemenzo, 1967, p.l26; Pillai and Scheer, 1976, p.33, pl.ll, figs.l, 2. Material Examined USNM: Fiji Islands, U.S. Expi. Exp., A. echinata syntype 275. UP: Batangas Channel, Puerto Galera, Oriental Mindoro C1064 (mentioned Nemenzo, 1967). I have not seen this species in the central and southern Great Barrier Reef. A single specimen in James Cook University was collected at Lizard Island on a sandy bottom. A large specimen in the Queensland Museum bearing no locality data has been presumed to be from the Great Barrier Reef. The species is well described and illustrated by other authors. Further notes are given with A. subglabra. Geographic Distribution Philippines, Suiu Sea, Great Barrier Reef, Fiji Islands, Samoa, Marshall Islands. Acropora subglabra (Brook, 1891) (Plate 94A, B) Madrepora subglabra Brook, 1891, p.470; 1893, p.l86, pi. 29, fig.c (synonymy). 308 MEMOIRS OF THE QUEENSLAND MUSEUM Acropora suhglahra: Thiel. 1933. p.24. M. subglahra var. rugixsa Brook, 1893, p.l87. Acropora suhglahra var. rugosa: Nemcnzo, 1967, p.l25, pi. 35, fig.2. Madrepora procumbens Brook, 1893, p. 188, pi. 29, fig.d. Acropora procumbens; Th\c\, 1932, p.l30, pi. 10, fig.2; Ncmenzo, 1967. p.I27-128, pi. 35, fig.l. Material Examined BM: A. procumbens syntype 1843.3.6.131; South Seas, A. subglabra syntype 1841.12.11.1. UP: A. subglabra var. rugosa C731 (mentioned Nemenzo, 1967). QM: Lizard Island fringing reef, June 1973, R. Pearson. G 1 07 13, G 107 1 4. Field Diagnosis Sprawling, shrubby bottlebrush colonies, of very slender proportions, colour pale brown or pinkish brown. Laboratory Diagnosis Branching pattern: Main branches may have any orientation from vertical to horizontal. Secondary branchlets distributed evenly around main branches, up to 30 mm long, undersurface branchlets being shorter than upper surface branchlets in horizontal branches. Axial corallites: Outer diameter 0-9 to 1-5 mm; inner diameter 0-5 to 08 mm. Septation: primaries complete, to 3/4 R, secondaries absent or some present to less than I /4R. Radial corallites; Scattered appressed tubular, tubo-nariform or nariform, up to 3 mm long. Non-appressed tubular radials are incipient axials. Coenosteum: Lines of elaborate spines both on and between radial corallites. Within Reef Distribution The species apparently does not occur in the Central and Southern Great Barrier Reef province, but is present further north, in situations below the reef flat. Identification Difficulties and History This species requires field study. Brook’s description of A. procumbens was on the basis of ‘contracted (axial corallite) apertures’, but in fact the internal axial diameter of the type is only an average 0-1 mm less than that of the A. subglabra type. A. echinata (Dana) is a similar species of slightly larger dimensions, and it is probable that this is a (senior) synonym. Geographic Distribution Indo-Pacific: Singapore, Philippines, Banda, Great Barrier Reef. Acropora carduus (Dana 1846) (Plates 93A, 94C, D) Madrepora carduus Dana, 1846, p.464, pi. 36, fig. 2; Brook, 1893, p.I78 (synonymy). Acropora carduus: Faustino, 1927, p.277, pi. 93, figs.l, 2; Nemenzo 1967, p. 123-4, pi. 34, fig. 3. Material Examined USNM; Sooloo Sea, U.S. Expl. Exped., Acropora carduus paratype? Ill (N.B. Rathbun (1887) records 278 from Fiji as type). QM: Big Broadhurst Reef, S W. slope: 6 m, 25.111.1973, C.W., G10729. G10730; 6-3 m. 23.X.1973, C.W., GI0738; 7-8 m, I4.X.1973, C.W., G10733; 8 m, 26.111.1973, C W, G10731; 8 m, 14.X.I973, C.W., GI0734; 9-6 m, 15.x. 1973, C.W., G10736; 9-8 m, 16.x. 1973, C.W.. GI0737; 12-4 m, 23.x. 1973, C.W., GI0732; 16-2 m, 23.X.I973, C.W., G10735. Bowden Reef. SW. side, reef slope. 2 m, 26.vii.i972, R. Pearson, G10741. G10742. Bushy-Redbill Reef: NW. side, sandy floor around patch reefs. 15 m, 15.vi.l975, C.W., G 10739; W. side, reef crest, 2 m, 3.vi-l975, C.W . G 10740. Fiji Islands, Great Astrolabe Reefs, Jan. 1974, C.W G 10744. Field Diagnosis Arborescent to shrubby, bottlebrush branched colonies of dimensions intermediate between those of A. subglabra and A. longicyathus. Colour; cream, pale brown, or pink-brown. Laboratory Diagnosis Branching pattern: Growth indeterminate, main branches covered by short branchlets, which are evenly distributed, more or less equal in size and extending at 45° to 90° from the branch. Total diameter of this ‘bottlebrush' is 15 to 30 mm, with branchlets up to 12 mm long and 5 mm wide. Main branches may be proliferous, giving a shrubby appearance, or sparsely branching, giving an arborescent appearance: usually both types of branching occur within a colony. In deep water (e.g. specimen G 10735) the branchlets develop on the upper surface of the colony only, and the colony is effectively reduced to a small plate. Axial corallites: From non exert (budding at tip) to 7 mm exert. Outer diameter 1-0 to 2 0 mm; inner diameter 0-5 to 0-8 mm. Septation: primary WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF 309 septa well developed, up to 3/4R, secondary septa absent, or some present, to 1/4R. Radial coralliles; On branchlets, scattered appressed tubular, sometimes approaching nar- iform, with round openings. On some colonies radials crowded, partly appressed tubular and extending out from branchlets. On the main branches radials are sub-immersed to immersed, or in some cases tubular appressed. Primary septa poorly to well developed to I/2R, secondaries usually absent except in immersed corallites, where a few may be present. Radial corallites are best represented on distal parts of branches. At the bases of branches most have developed into long tubular axials. Coenosteum: finely echinulate on radial corallites and between. Occasionally spines are arranged in rows or even joined as costae. Within Reef Distribution Reef slope, from below the area of rough water surge to about 20 m; deep water reef flat areas, sandy floors of lagoons and reef-edge patch reefs. Identification Difficulties and History Being a deeper water species, A. carduus is poorly represented in older collections, and hence presents few historical problems. > It occurs with A. longicyathus. and they can be easily separated by obvious differences in corallite dimensions. Geographic Distribution Indo-Pacific: Philippines, Sulu Sea, Great Barrier Reef, Fiji Islands. Acropora longicyathus (Milne Edwards and Haime, 1860) (Plates 93B, 95) Madrepora longicyathus Milne Edwards and Haime, 1860, p.l48: Brook. 1893, p.l87 (Synonymy). Acropora longicyathus: Nemenzo, 1967, p.l28, pi. 35, figs. 3, 4. Madrepora prolixa Verrill, 1866, p.22. Acropora prolixa: Verrill, 1902, p.237, pi. 36, figs. 3, 3a, pl.36A, Figs. 3, 3a; pl.36F, fig.l4; Hoffmeister, 1925, p.65, pi. 16; Crossland, 1952, p.226. Material Examined: PM: A. longicyathus type 303A. USNM: Ousima, A. prolixa syntype 414. QM; Big Broadhursl Reef, S W. side, reef slope: 6m, 25.iii.1973, C.W., G10763; 7m, 27.iii.l973, C.W., G10767; 8m, 26.iii.1973, C.W., G10762, G10764; 9m, 15.X.1973, C.W., G10751, G10752; 9-6m, 15.x. 1973, C.W., GI0753; 9 7m, 15.X.1973, C.W., G10754; 9-8m, 16.x. 1973, C.W., G10755; 10-3m, 16.X.1973, C.W., G10756; I0-6m, 16.X.1973, C.W.. G10757, G10758; 12m, 16.x. 1973, C.W.. G10759; 12-5m, 17.X.1973, C W.. G10760; surge channel, 10m, 20.X.1973, C.W., GI076L Bushy-Redbil Reef: W. side, outer flat, 31.V.1975, C.W.,G]0745; reef slope near Redbill Is., 3m, l.vi.l975, C.W., G10746; W. side, patch reefs (sandy floor), 8m, 3.vi.l975, C.W., G10747, G10748; N W. side, patch reefs (sandy floor) I2m, 14. vi. 1975, C.W., G10749, G10750. Field Diagnosis Arborescent to shrubby, bottlebrush branched colonies of sturdy dimensions. Colour: cream, pale to dark brown, or blue-brown. Laboratory Diagnosis Branching pattern: Growth indeterminate, main branches covered by evenly distributed proliferous branchlets or bundles of branchlets. These are directed at 45° to 90° to the branch, and may be as short as 5 mm all over branch, or all several centimetres long and much branched. Greatest total branch width is 80 mm, smallest (except for branch tips) 40 mm. Deep-water colonies can have little undersurface development, and approach a plate like shape. Axial corallites: Sometimes exert (to 10 mm) near base of branch, more often one side naked of radial corallites, other side with 3 or 4 from their outer rim. Outer diameter 2-1 to 2*8 mm; inner diameter 0-8 to 13 mm. Septation; primary cycle present to 3/4R, secondary cycle present, or at least partly developed up to 1 /4R. Radial corallites: On branchlets, appressed or partly appressed tubular with round openings. When fully appressed, radials are scattered, only a few to each axial corallite: however, branchlets can have radials touching. In the first case, radials on main branches are immersed or (more usually) sub-immersed; in the second, main branch radials are usually similar to those of branchlets. Coenosteum: Neatly echinulate on and between corallites, the spines laterally flattened with simple to forked or more elaborate lips. Within Reef Distribution Reef slope, from below the area of rough water surge to about 20m; deep water reef flat areas, sandy floors of lagoons and leeward patch reefs. Identification Difficulties and History The two species A. longicyathus and A. prolixa have been combined by other authors: Verrill’s 310 MEMOIRS OF THE QUEENSLAND MUSEUM specimens were small branch tips. This species is not well represented in collections. Geographic Distribution Philippines, Ousima, Samoa, New Guinea, Great Barrier Reef. The ‘Acropora .squarrosa’ Group The group of species A. elseyi. A. rosaria, A. squarrosa and A. granulosa has in common (1) dense cchinulate cocnosleum of elaborate spines, (2) large tubular to nariform radial corallites, and (3) a tendency for some branchlcts to have an upper surface naked of radial corallites. The group is linked to the \4. echinata' group which has, however, less differentiated radial corallites. The synonyms adopted here may be controver- sial. Most of the species involved seem to be well documented in the literature, but they still present extraordinary identification problems. The synonymies are presented as hypothe.ses requiring further testing. The oldest name for the group, A. squarrosa (Ehrenberg) has been treated by a number of authors (c.g. von Marenzeller 1907, Vaughan 1918, Wells 1954), but the interpretations are various and sometimes confusing. A. granulosa. with its synonyms here listed, links this group to the ‘A. echinata' group. A. nncrophthalma, as here interpreted, is included in the group because of its cocnosteal structure. Acropora microphthalma (Verrill, 1869) (Plate 96) Madrepora microphthalma Verrill, 1869, pp.83, and 102 . Acropora microphthalma: Verrill, 1902, p.232, pl.36C, fig.l, 36E, fig. 1 5. Acropora microphthalma: Wells, 1954. p.429, pi. 126, figs. 7 9 (synonymy), non Acropora microphthalma: Stephenson and Wells, 1956, p.lO. Acropora laevis (part) Crossland, 1952, p.230. Mate-rial Examini-d YPM: Ryuku Islands, A. microphthalma holotype 774 (fragment). QM: Big Broadhurst Reef, SW. side, reef slope; 3m, 28.111.1973, C.W.. G8684; 4m, 13.x. 1973, C.W., G8688, G8689; 6m. 25.iii.1973, C.W., G8685; 6-4m, 14.x. 1973, C.W„ G8693; 7-8m, 15.X.1973, C.W., G8691; 8m. 26.111.1973, C.W., G8683, G8686; 8m, 28.iii.1973, C.W., G8682; 91m, 16.X.1973, C.W., G8690; 9-4m, 16.X.1973, C.W., G8692. Bowden Reef, SW. side, reef crest, 0-3m, 24.vii.I972, C.W., G8680. Field Diagnosis An aborcsccnt species forming small clumps (up to 100 cm across) the branches having slender dimensions, and radial corallites being small and crowded. Colour is most commonly a whitish- cream. Although because of its size it is not a conspicuous species, it is the most delicate arborescent coral in the Great Barrier Reef area, and is easily recognized. Laboratory Diagnosis Branching pattern: Branching is open arbores- cent, with many short branchlcts being given off towards branch tips at 45*^ to 90° to the main branches. The greatest branch width measured in the collections is 14 mm, the smallest (excluding branchlcts) 5 mm. Axial corallites: Outer diameter 1-8 to 2-3 mm; inner diameter 0-8 to 1 00 mm. Septation: primary septa strongly developed, up to 3/4R, secondaries absent, to prc.scnl up to 1/4R. Radial corallites: Extend at about 45°, short tubular with round to oval opening to tubo- nariform, outer wall slightly thickened. First septal cycle well developed, up to 2/3R, second cycle usually partially represented. Radials arc uniform in size and evenly distributed, giving a fine-grained appearance to the branches. Cocnosleum: When spines are well developed, they have elaborate tips and are distributed both on corallites and between. However, lightly calcified coralla may have simple pointed spines and a generally spongy appearance. Within Reef Distribution Middle reef flat, deeper reef flat areas, sandy floors around patch reefs, reef slope to surge channel floor. Identification Difficulties and History Interpretation of A. microphthalma from VerrilTs poor series is difficult and previous identifications have linked it with A. exilis. The A. microphthalma of Stephenson and Wells (1956) (no. G2697, QM) is in my opinion A. nasuta. The species as here interpreted is distinctive, particularly in the field, and the fragment I have seen of VerrilTs type compares well with the tips of the larger dimensioned specimens. At least one specimen of Crossland's A. laevis series (BM 1934.5.14.67) is this species. Geographic Distribution Ryuku Islands, Great Barrier Reef, Fiji Islands (C.W.), Enewetak Atoll (C.W.). WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 311 Acropora elseyi (Brook, 1892) (Plate 97 A, B.) Madrepora elseyi Brook, 1892, p.456; 1893, p. 172, pl.ll, figs. E, F. Acropora elseyi: Crossland, 1952, p.223; Pillai and Scheer, 1976, p.31, pL9, fig.l. Madrepora exilis Brook, 1892, p.457; 1893, p.l72, pi. 10, figs. C, D. Acropora exilis: Crossland, 1952 (part), p.223, pi. 39, fig. 4; Stephenson and Wells, 1956, p.l3, pi. 1(b) (synonymy). Material Examined BM: N. Australia, J. Elsey, A. elseyi syntypes 1857.11.18.214, 215, 216, 217, 218; Thursday Is., Saville-Kcnt, A. elseyi 1892.6.8.241-4, 6; Rocky Is. Saville-Kcnt, A, elseyi 1892.6.8.247 (mentioned Brook, 1893). Port Denison, Savillc-Kent, A. exilis syntypes, 1892.6.8.103, 104, 106. QM: Big Broadhurst Reef, E. side, side of reef flat channel, 2Lx.i973, C.W., G10802. Bushy-Redbill Reef: W. side, middle reef flat, Im, 27.V.1975, C.W., GI0793; G10794; W. side, outer reef flat, 8.vi.l975, C.W., G10797; N W. side, first reef crest, 23.xii.1972, C.W., GI0798; adjacent Redbill Is., reef crest, 19.xii.l972, C.W., G 10795; adjacent Redbill Is., reef slope, 19.xii.l972, C.W., G10796. Field Diagnosis Low bushy to caespitose colonies with bottlebrush-lype branches. Colour usually bright yellow, yellow-brown or lime green-brown. Laboratory Diagnosis Branching pattern: Main branches surrounded by regularly placed short branchlets which may be of equal or unequal length. Main branches up to 15 mm wide, branching units up to 100 mm wide, branchlets up to 8 mm wide. Axial coralliles: From non-exerl to 2 mm exert. Outer diameter 1-6 to 3-2 mm; inner dimaeter 0*6 to 10 mm. Septation: primary septa well developed, up to 3/4R, secondary septa absent or few present to less than 1/4R. Radial corallites: On both main branches and branchlets, radials are tubular with round openings, becoming round tubular distally. Radials are usually evenly distributed on branches, and almost touching, sometimes upper surface of small branchlets is naked of corallites. Septation: primaries well developed, up to 1/2R, secondaries absent or a few just visible. Coenosteum: Neatly and densely costate both on and between corallites. Within Reef Distribution This is one of the few Acropora occurring in the shallow longitudinal patch reefs perpendicular to the reef edge, forming the windward edge of lagoons, and is characteristic of these areas. It also occurs in deeper reef flat areas, and reef crest and upper reef slope on some leeward reef areas. It is a common species of fringing reefs of continental islands. Identification Difficulties and History Both species in synonymy were described from the Barrier Reef, A. elseyi being slightly sturdier than ‘A. exilis’. The species will be better understood when studied on fringing reefs. Geographic Distribution Indo-Pacific: Maidive Archipelago, Great Barrier Reef. Acropora rosaria (Dana 1846) (Plate 97 C, D) Madrepora rosaria Dana. 1846, p.465, pi. 36, fig. 3. Madrepora rosaria var. diffusa Brook, 1893, p. 1 80. Acropora rosaria: Vaughan, 1918, p.l84, pi. 82, figs. 2, 2a, 2b; Weils, 1954, p.428, pi. 130, figs. 3, 4; Stephenson and Wells, 1956, p.l8. Acropora rosaria (part): Crossland, 1952, p.224, pi. 40, figs.l, 4. Madrepora syringodes (part) Brook, 1892, p.463; 1893, p.l77, pi. 33, fig.E. Material Examimt:) USNM: Fiji islands, U.S. Expl. Exped., A. rosaria holotype 281; paratype 933. QM: Palm Islands, 1939, T. C. Marshall, G9192, G9193, G9I94, G9I95, G9196, G9197, G9200, G9202. Field Diagnosis Not determined. Laboratory Diagnosis Branching pattern: Usually ‘bottlebrush’ branching units arc formed, a central branch of up to 20 mm diameter bearing branchlets at even intervals all around. Axial corallites: Exert 10 to 2 0 mm. Outer diameter 2-5 to 3-8 mm; inner diameter: 0-8 to 2-4 mm. Septation: first cycle well developed, up to 3/4R, second cycle usually present, or most septa present, up to 1/3R. Radial corallites: Tubular, partly or fully appressed, or nariform. Walls thick, primary septa developed up to 1/3R and secondary septa partly developed, up to I/2R. Coenosteum: Densely arranged spines, some- times laterally flattened, both on and between corallites. 312 MEMOIRS OF THE QUEENSLAND MUSEUM Within Reef Distribution The specimens described were collected on the fringing reefs of the Palm Island group. Identification Difficulties and History This species requires further study. The only material easily identifiable with A. rosaria available to me was collected in habitats not comparable with those I have studied. I am thus unable to pursue the possibility of a close relationship between this species and A. squarrosa. I consider it probable that the figured syntype of Brook's A. syringodes (BM 1892.6.8.209 from Palm Islands) is A. rosaria. Interpretations of A. syringodes are difficult to follow and seem to span a number of species, and it is likely that the latter species, as described, is a mixture. Geographic Distribution Great Barrier Reef, Fiji Islands, Samoa, Marshall Islands. Acropora squarrosa (Ehrenberg, 1834) (Plates 98, 99, 100) Heteropora squarrosa Ehrenberg, 1834, p.l!2. Madrepora squarrosa: Brook, 1893, p.65 (synonymy). Acropora squarrosa: von Marenzeller, 1907, p.46, pi. 1 4, figs. 36-39; Vaughan. 1918, p.l84, pi. 83, figs. 2, 2a, 2b; Wells. 1954, p.427, pi. 129, figs. 1, 2; Rossi, 1954, 1954, p-52; Nemenzo, 1967, p.69, pl.21, fig.4; Pillai and Scheer 1976, p.31. Acropora murrayensis Vaughan, 1918. p. 183, pi. 82, figs.l, la. lb; Nemenzo, 1967, p.71, pi 23, fig.2. Madrepora syringodes (part) Brook, 1892, p.463; Brook, 1893, p. 177 (not illus.). Madrepora cancellata Brook, 1893, p. 166, pi. 32, fig.C. Acropora cancellata: Crossland, 1952, p.225, pl.41, figs.3, 4. Material Examinrd USNM: Murray Islands, A. murrayensis holotypc. BM: A. syringodes 1893.4.7.163 (mentioned Brook, 1893); Louisadc Archipelago, 15 fathoms, A. cancellata holotype 185L9.29.39. QM: Big Broadhurst Reef, SW. side, reef slope: 1-3 m, 23.X.1973, C.W., GI0839; 1 -9 m, 23.x.i973, C.W., G10840; 6-3 m, 13.X.1973, C.W., GI0826, G10827; 6-3 m. 23.X-1973. C.W.. G1084I; 7 m, 13.x. 1973, C.W., G10825; 7-8 ni. I4.X.I973. C.W.. G10828, G10830; 8 m. 14.X.I973, C.W'., G10829; 81 m, 14.X.1973, C.W.. GI0824; 8-2 m, 14.X.1973, C.W., G10831; 8-7 m, 15.X.1973, C.W., G10832; 9 m, I5.X.1973, C.W., G10833, G10834; 9-2 m, 25.X.1973, C.W., G10842; 10-6 m, 16.X.I973, C.W., G10836; 11-4 m, 25.X.1973, C.W., G10843: 12*9 m, 23.X.1973, C.W., G10845; 13-4 m, 17. X.1976, C.W., G10837; 30 m, I8.x.i973, C.W., G10838; E. side, reef slope, 4 m, 2Lx.l973, C.W.. G 1 0844. Bushy-Rcdbill Reef: Reef crest adjacent Redbill Is.: 18. xii.l972, C.W., G10768, G10770; 19.xii.l972, C.W., G10769; Lvi.I975, C.W., G10771; reef slope adjacent Redbill Is.: 20.xii.l972, E. Lovell, G10772, G10773: 20.xii.I972, C.W., G9I89; W. side, middle reef flat; 1 m, 8.vi.l975. C.W., GI0774, G10775; I4.vi.l975, C.W., G10776; NW side, first reef crest: 22.xii.I972, E. Lovell. G10777; 22.xii.1972, C.W., GI0778, G10779; NE. side, D. Hadley. I. i. 1973, G10780. Bowden Reef, SW\ side, reef slope: 24.vii.1972, C.W,, GI0789; 26.vii.1972. C.W., GI0788. Darley Reef, patch reef in lagoon: Im, I8.vii.l972, R Pearson, GI0782, GI0785; 1 m, 22.iii.I973, C.W., G10783, GI0784, G10786, GI0787. G10790; 3 m, I8vii.l972, C.W.. GI0846; 3 m, 24.iii.1973, C.W., G10791; 4 m, 22.iii.1973, C.W., G10792; 6 m, 22.iii.1973, C W.. G10847; 7 m, 22.iii. 1973, C.W., G 10846. Viper Reef, patch reef in lagoon, 2 m, 16.vii.l972, C.W., G1078L Field Diagnosis Colonies can be bottlebrush branching, caespitose to plate like, some of the variation being due to depth gradients. The surface of the branches has a general ‘globular’ appearance due to rounded, thickened corallites, and a whitish glow behind the general colouration due to dense coenosteum showing through semi-transparent tissue. Common colours are pale blue, lavendar, cream-brown and yellow-brown. Laboratory Diagnosis Branching pattern: Bottlebrush branching colonies have tapering branching units, and the branchlets extend widely from the main branch (90° or just less). In caespitose colonies there is a tendency tow'ards development of a bottlebrush unit (incipient branchlets developed evenly around the branches) and even in plate-like colonies the short upright branches bear indications of incipient branchlets along their length. In most specimens of this species branchlets which are naked of radial corallites on their upper surface can be seen. Axial corallites; From barely exert to 2*5 mm exert. Outer diameter 2-6 to 3-8 mm; inner diameter 0-7 to 1-4 mm. Septation: primaries usually well developed (up to 3/4R), secondaries completely absent to fully present, up to 1/3R. Radial corallites: Tubular appressed to nariform, with round to oval openings at 90° or less to branch. Wall is thickened so that corallite WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF 313 may appear round tubular, or distorted rostrate hooks or horns may be developed. Septa are usually poorly developed; primaries can usually be just detected, and sometimes a few second- aries. Coenosteum: Laterally flattened spines with elaborated tips are arranged densely and evenly over both the corallites and the inter-corallite region. Identification Difficulties and History The material examined shows extraordinary variation in colony shape, branch size, and radial corallile distribution. However, I am unable to separate the collection into units on any of these characters. The evidence suggests a species where the role of the radial corallite is very flexible. The ‘easiest’ identification in this group is A. murrayensis s.s,, which was well described by Vaughan. This occurs particularly on sandy floor situations in lagoons and deep water reef flat areas. On the reef crest and upper reef slope it can also occur, but with shortened ‘botllebrush’ branches. The caespitose to plate-like forms occur mainly on sloping surfaces, and on deep parts of the reef slope small flat plates approaching A. granulosa are formed, these being identifiable with A. cancellata s.s. At least one of Brook’s syntypes of A. syringodes (BM 1893.4.7.163) from unknown locality, compares closely with caespitose specimens in the present collection. The biggest problem is in interpreting A. squarrosa s.s, which I have taken (following Vaughan 1918) to be caespitose, approaching corymbose, with very regularly arranged radial corallites and relatively thick branches. I suspect A. austera specimens are sometimes identified to this species. Geographic Distribution Indo-Pacific; Red Sea, Seychelles, Maldives, Minicoy, Philippines, Louisade Archipelago, Great Barrier Reef, Marshall Islands. Acropora granulosa (Milne Edwards and Haime, 1860) (Plates 101, 102) Madrepora granulosa Milne Edwards and Haime, 1860, p.l56; Brook, 1893, p.l89 (synonymy). Madrepora speciosa Quelch, 1886, p.l63, pi. 10, fig.l; Brook, 1893. p.l91. Madrepora clavigera Brook, 1892, p.455; 1893, p. 183, pi. 9, figs. A, A'. Acropora clavigera: Crossland, 1952, p.226, pi. 40,, fig. 2, pl.42, fig.3. Madrepora rayneri Brook, 1892, p.461: 1893, p.l91, pi. 8, fig.A. Acropora rayneri: Wells, 1954, p.431, pi. 134, fig. 6, pi. 137, figs. I, 2, pi. 139, figs.l, 2. Material Examined PM: Acropora granulosa holotype 328a. BM: Acropora clavigera holotype 1851.1 1.14.28. Fiji, Acropora rayneri syntypes, 1862.2.4.30, 1862.2.4.44. QM; Big Broadhurst Reef, SW. slope: Oct., 1973, C.W., G1I483; 5 m, 27.iii.1973, C.W., G11476; 7 m, 26.iii.l973, C.W.. 011474; 7 m, 27.iii.1973, C.W., GI 1477, G1 1493; 8 m, 28.iii.1973, C.W., G1 1492; 8 m, 24.X.1973, C.W., G1I481; 10 m, 23.x. 1973, C.W., GI1494; 10-4 m, 16.X.1973. C.W., G11487; 17-2 m, 17.X.1973, C.W., GH480; 25 m, 17.x. 1973, C.W., 011485. Bushy-Rcdbill Reef: adjacent Redbill Is., reef crest, I m, Lvi.1975, C.W., G11490; 20.xii.l972, E. Lovell, Gl 1491; NW. side, patch reefs, I5.vi.l975, 12 m, C.W., G11488. Darley Reef, patch reef in lagoon, 2-5 m, 19.vii.l972, C.W., G6723. Feather Reef, reef slope, 10 m, 24.x. 1972, R. Pearson, G11479, G11482. Field Diagnosis Colonies are side-attached thin plates, with anastomosing horizontal branches and short vertical branchlets or groups of branchlets. Axial corallites are long, tapering or rounded, and may be sinuous. Radials are relatively few and .scattered. Laboratory Diagnosis Branching pattern; described above. Axial corallites: From barely exert to as long as 10 mm bare of radials. Outer diameter 1-2 to 2-5 mm; inner diameter: 0-6 to 2-2 mm. Septation; primaries well developed, up to 3/4R, secondaries absent or poorly represented, to less than 1/4R. Radial corallites: Appressed tubular to nariform with round to slightly oval opening, directed at 90^ or less to branch. Radials, except at growing edge of colony, are few, and inconspicuous, the appearance of the colony being dominated by the axial corallites. Coenosteum; Spines with pointed or laterally flattened tips are densely and evenly arranged on and between corallites. Within Reef Distribution Reef slope and sides of lagoonal patch reefs, usually at depths greater than 20 m, but can be much shallower on very steeply sloping surfaces, for example G 11490 is a specimen from 1 m on the side of a sleep and narrow surge channel opening. 314 MEMOIRS OF THE QUEENSLAND MUSEUM Identification Difficulties and History The conibinalion of the various species in synonymy seems barely possible from Ihe type material, and also from much of my own material which identifies strongly with one or another species. A more acceptable combination would be two species, A. speciosa = A. rayneri and A. granulosa = A. clavigera. However some large specimens show a mixture of characteristics of all four species. The differences amongst the described species are in dimensions and the degree of tapering of the axial corallilc; ‘A. rayneri' is very slight, and A. speciosa' sturdier, both with tapering axials, A. granulosa s.s. and ‘A. clavigera' both have thick, rounded, non-tapering axials. In all the material there is little difference in inner axial diameter, axial septation, radial corallite shape and coenosteal spines. The locations of the specimens give no clue to the morphological differences. All categories except A. speciosa' (one lagoonal specimen) co-occur on the reef slope. Their different appearances (whether genotypically or phenotypically deter- mined) cannot be explained by reef slope gradients, and are probably related to rnicrohabi- tat features. Despite the variety in the species, extreme care must be taken in assigning a specimen to this species, as deeper water specimens of many other species appear (as cleaned specimens) similar to this species. Geographic Distribution Indo-Pacific: Mascarene Archipeligo (G. Faurc, pers. comm.), Louisade Archipelago, Great Barrier Reef, Fiji Islands, Marshall Islands. Acropora austera (Dana, 1846) (Plates 103, 65C) Madrepora austera Dana, 1846, p.478; Brook, 1893, p.56 (synonymy); Verrill, 1902, p.266, pi 36, fig. 10, pi. 36B, fig. 1. Materiai. Exmaim d YPM: Acropora austera (fragment of type) 4190. QM: Big Broadhursl Reef, S W. side, reef slope: 7-8 m, I4.X.1973. C.W., G 108 1 3, G 10828; 8-0 m, 14.X.I973, C.W., G10814; 8-2 m, 14.x. 1973. C.W., G10815; 9-0 m, 15.x. 1973. C.W., G10816. Bowden Reef. SW. end. reef slope, 26.vii.I972, C.W.. G10817. Bushy-Redbill Reef: adjacent Redbil! Is., reef crest: 2.vi.l975. C.W.. G10806, GI0807; 4.vi.l975, C.W., G 10808; S. side, reef slope, 3 m, 27.xii.1972, C.W., G10809, G10810; W, side, patch reef, 8 m, 15. vi. 1975, C.W., G10811; N W. side, patch reef, 19.xii.l972, E. Lovell, G 108 12. Darley Reef, patch reef in lagoon, 24.iii.1973, C.W., G10818, G10819. Viper Reef, patch reef in lagoon, 16.vii.l972, C.W., G 1 0820. Fiji Islands, Great Astrolabe Reefs, .Ian. 1974, C.W., GI0821, G10822. Field Dlagnosis Patchy, irregularly branching colonies of bushy, bottlcbrush or caespitose appearance; large, irregular-length rounded radial corallitcs with very large openings. Colour cream to pale brown or dirty yellow. Extended polyps may be bright orange (axials) and purple (radials). Laboratory Diagnosis Branching pattern: Main branches are up to 40 mm width. These give off secondary branches of such irregular spacing, angle, and length, that the colony may appear caespitose, bottlcbrush, or low arborescent. Although growth is apparently indeterminate, colonies do not often reach more than about 1 m across, occurring usually in areas of high density cover. Axial corallites: From barely exert to 3 mm exert. Outer diameter 2-4 to 3-8 mm; inner diameter 1-0 to 1*5 mm. Septation: all septa usually present, primaries up to 2/3R, secondaries up to 1 /2R. Radial corallites: On secondary branches radials are tubular, nariform or tubo-nariform, of mixed length. Similar corallites on main branches tend to become appressed, then rounded, distally. The shape of the opening approaches a square, and the lower (or outer) wall is sometimes thickened accentuating the angularity of the opening. Primary and most or all secondary septa are developed and, as Dana comments, these usually slope towards the centre deep in the corallite. Coenostcum: Reticulate with elaborated spines on and between radial corallites. Within Reee Distribution Reef crest and upper slope, top and sides of lagoonal patch reefs. This species seems to occur particularly where there is a bend or edge on the reef surface (see plate 65C). Identification Difficulties and History This species is neglected in the literature, although my experience has shown it to be widely distributed. Geographic Distribution Indo-Pacific: Singapore, Philippines, Great Barrier Reef, Fiji (C.W.), Enewetak Atoll (C.W.). WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 315 Acropora brueggemanni (Brook, 1893) (Plate 50C, D) Madrepora brueggemanni Brook, 1893, p. 145, pi. 24. (synonymy). Acropora brueggemanni: Crossland, 1952, p.221. Madrepora brueggemanni var. uncinata Brook, 1893,, p.l46, pi. 35, fig. E. Acropora brueggemanni var. uncinata: Nemenzo, 1967, p.55. Material Examined BM: Singapore, A. brueggemanni syntype 1878.4.1.1. JCU: Palm Island Group, T. Done, 4 specimens; Lizard Island Group, M. Pichon and T. Done, 6 specimens. QM: Big Broadhurst Reef, SW. side, reef slope, 5 m, 27.iii.l973, C.W., G11495; 6 m, 27.iii.1973, C.W., G1 1496-8. Palfrey Islet, Lizard Island group, 30.vii.l977, P. Hutchings, G1 1499-506. Field Diagnosis Sturdy arborescently branching colonies occur as either small clumps or extensive thickets. Axial corallites are obviously large and bulbous and there may be several axials at or near the branch tip. General appearance approaches that of A. palifera (see Wells 1954, p.430), but a definite branching pattern is achieved. Colour is pale brown to pale apple green. Laboratory Diagnosis Branching pattern: The branches may be round or irregular in cross section, from 15 to 30 mm diameter, and either tapering or truncate. The angle of branching is usually wide. Axial corallites: Outer diameter 2-0 to 8 0 mm; inner diameter 0-8 to 1-4 mm. In general, when there is a single axial corallite, this is wider than the members of a bundle of axials. Septation: Both septal cycles usually developed, the primaries up to 3/4R, secondaries up to 1/3R. Radial corallites: From short (barely emergent) to appressed tubular, with round openings. Primary septal cycle usually well developed, up to 1 /2R; secondary cycle poorly developed, up to 1/4R. Coenosteum: Dense arrangement of elaborated spines both on and between radials. Within Reef Distribution In my work, I have encountered this species only as a rare member of the upper reef slope assemblage. It is very common on the fringing reefs of the Lizard Island and Palm Island Groups, usually on sandy or poorly consolidated substrates. IDENTIFICATION DIFFICULTIES AND HISTORY Both Brook and Crossland note an approach to the characteristics of subgenus Isopora in this species. Strong affinities with A. palifera can be seen in the radial corallite structure, coenosteal texture, and tendency to multiple axial corallites. It is considered in the present paper in order to avoid field-identification problems. Geographic Distribution Singapore, Philippines, Great Barrier Reef. ACKNOWLEDGMENTS I wish to express my gratitude to the following: Prof. Jan Verwey (whose taxonomic decisions on Acropora were made many years ago but whose search for perfection still keeps him from publication) for his generosity in discussing his work, and Mrs Verwey for her hospitality. Prof. John Wells for encouragement, constant assis- tance and hospitality. Other workers in coral taxonomy, in particular Maya Wijsman-Best, Richard Randall, Brian Rosen, Michel Pichon, Gerard Faure, John Veron, and members of the N.S.F. Enewetak Taxonomic Workshop (1976) for advice and discussion. Micky Watkins and Hugh Hope who took us to the reef. Robert Pearson whose unpublished observations con- tributed to the ground work of the study, and Colin Limpus who made large contributions of specimens and observations. Museum colleagues who helped in the field (Roily McKay, Terry Tebblc, Dianne Gleeson, David Joffe); in separate field collecting (Glen Ingram, Jan Buhmann) and back-up work (Alan Easton and staff, photo- graphy; Peter Berryman and staff, artwork); many members of technical and clerical staff, and Bruce Campbell and Lester Cannon for advice on the manuscript. Peter Beveridge and family and Bruce Carlson for hospitality in Fiji. The following Museum Curators and their staff for assistance with specimens and hospitality: Dr K. Reutzler and Dr F. Bayer (USNM), Dr P. F. S. Cornelius (BM), Dr J. P. Chevalier (PM), Dr W. Hartman (YPM), Dr D. Kuhlmann (MNB), Dr M. Wijsman-Best (RNHL), Dr Abad-Santos (LP), Dr Van Soest (Zoological Museum Amsterdam). Neville Coleman for considerable assistance with coloured underwater records. D. and M. Tarca of Townsville. Many other friends in the field, but in particular Ed Lovell, Robin Elks, Harvey Walsh and Carl Wallace. John Hardy and Bob Grimmer of the University of Queensland MEMOIRS OF THE QUEENSLAND MUSEUM Electron Microscope Unit. Michel Pichon and Terry Done for hospitality at James Cook University. The study was supported by grants from the Advisory Committee on Crown-of-Thorns starfish and Queensland Museum Funds. LITERATURE CITED Bassett-Smitu. P. W., 1890. Report on the corals from Tizard and Macclesfield Banks, China Seas. Ann. Mag. Nat. Hist. (6) 6: 353-74. Boschma. H., 1961. Acropora Oken, 1815 (Anthozoa, Madreporaria): proposed validation under the plenary powers. Bull. Zool. Nomencl. 18(5); 334-5. Brakei . W. H., 1977. Corallite variation in Porites and the species problem in corals. Proc. 3rd Inter. Coral Reef Symp. Miami, Florida: 457-62. Brook. G., 1891. Descriptions of new species of Madrepora in the collections of the British Museum. Ann. Mag. Nat. Hist. (6) 8; 458-71. 1892. Preliminary descriptions of new species of Madrepora in the collections of the British Museum. Part 11. Ann. Mag. Nat. Hist. (6) 10: 451-65. 1893. The genus Madrepora. British Mus. (Nat. History) Cat. Madreporarian Corals, 212 pp., 35 pis. China. W.E., 1963. Opinion 674: /tcroporo Oken, 1815 (Anthozoa, Madreporaria): validated under the plenary powers. Bull. Zool. Nomencl. 20(5): 329-30. Crossland, C., 1928. Notes on the ecology of the reef-builders of Tahiti. Proc. Zool. Soc. London 1928: 717-35, 5 pis. 1931. The reduced building-power and other variation in the Astrean corals of Tahiti, with a note on Herpetolitha Umax and Fungia sp. Proc. Zool. Soc. London 1931: 351-91, 22 pis. 1952. Madreporaria, Hydrocorallinae, Heliopora and Tubipora. Sd. Reps. Great Barrier Reef Exped. 1928-29 6: 85-257. 56 pis. Dana, J. D,, 1846. Zoophytes; U.S. Exploring Exped., 7: 740 pp.; atlas, 61 pis. (1849). Ehrenberg. C. G., 1834. Beitrage zur physiologischen Kenntniss der Corallenthiere im Allgemeinen und besanders des Rothen Meeres. Abh. K. Akad. Wiss. Berlin 1832: 250-380. Faustino. L. a., 1927. Recent Madreporaria of the Philippine Islands. Monograph 22, Bur. Sci. Maniia, 1-310, 100 pis. Gardiner. J. S., 1898. On the perforate corals collected by the author in the South Pacific. Proc. Zool. Soc. London: 251-lb, 1 pis. Hoffmeister. j. E., 1925. Some corals from American Samoa and the Fiji Islands. Pap. Dept. Marine Biology Carnegie Inst. Washington 22: 1-90, 23 pis. 1929. Some reef corals from Tahiti. Jour. Washington Acad. Sci 19: 357-65, pls.l, 2. Horn. H., I860. Description of new corals in the Museum of the Academy. Proc. Acad. Nat Sci Philadelphia 1860: 435-36. Kllnzinger, C. B., 1879. Die Korallentheire des Rothen Meeres. Zweiter Theil; Die Stienkorallen. Erster Abschnitt; Die Madreporaceen und Oculin- aceen, pp. 1-88, 10 plates (Berlin). Lamarck. J. B. P. de, 1816. Histoire naturelle des Animaux sans vertebres. Vol. 2, pp. 1-568. (Paris). Marenzeller. E., von.. 1907. Expeditionen S.M. Schiff ‘Pola’ in das Rote Meer. Zool. Ergeb. 26. Riffkorallen. Denkschr. Akad. Wiss. Wien 80* 27-97, pis. 1-29. Maxwell, W. G. H., 1968. ‘Atlas of the Great Barrier Reef. 258 pp. (Elsevier; The Netherlands). Mayor, A. G., 1924. Preface to: some posthumous papers of A. G. Mayor. Pap. Dep. Marine Biology Carnegie Inst. Washington 19: v-vii. Milne Edwards. H. and Haime. J., I860. Histoire naturelle des coralliaires, Vol.3, 560 pp. (Paris). Moore. R. C., Hill, Dorothy, and Wells. J. W., 1956. Glossary of morphological terms applied to corals. PP.F245-251 in Moore, R, C., (ed.) ‘Treatise on Invertebrate Paleontology’ part F Coelenterata (Geol. Soc. Amer.). Nemenzo. F., 1967. Systematic studies on Philippine shallow-water Scleractinians: VI. Suborder As- trocoeniida (Montipora and Acropora). Nat. Appl. Sci. Bull. Univ. Philippines 20(1); 1-141 (text); (2): 144-223 (plates). Oken, L., 1815. Steinkorallen. Lehrb. Natureesch 3nV 59-74. ' ^ Ortmann, a., 1888. Studien uber Systematik und geographische Verbreitung der Steinkorallen. Zool. Jahrb. Abt. Syst. 3: 143-788. 1889. Beobachtungen an Steinkorallen von der Sudkuste Ceylons. Zool. Jahrb. Abt. Svst 6* 631-70. PiLLAL C. S. G., and Scheer. G. 1974. On a collection of Scleractinia from the Strait of Malacca. pp.445-64 in Proceedings of the Second Interna- tional Coral Reef Symposium Vol 1. (Great Barrier Reef Committee: Brisbane). 1976. Report on the stony corals from the Maidive Archipelago. Zoologica 126: 1-83, 32 plates. Potts, D. C., 1976. Growth interactions and morphological variants of the coral Acropora palifera. pp. 79-88 in Mackie, G. O. (Ed.) ‘Coelenterate Ecology and Behaviour’ (Plenum Publishing Corp: New York). 1977. Differentiation in coral populations. Atoll Res. Bull: in press. Quelch, j, j,, 1886. Report on the reef-corals. Challenger Reports Zool. 16: 1-203, 12 pis. Rath BUN. R., 1887. Catalogue of the species of corals belonging to the genus Madrepora contained in the U.S. National Museum. Proc. U.S. Nat. Mus 10: 10-19. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 317 Ricarty Menendez. F. O. and Friedman, G. m., 1977. Morphology of the axial corallite of Acropora cervicornis. Pp 453-6 in Proceedings of the Third International Coral Reef Symposium (University of Miami, Florida). Rossi. L., 1954. Spedezione subacquea italiana nel Mar Rosso. Richerche zoologiche V. Madreporarii, Stoloniferi e Milleporini. Riv. Biol colon. 14: 23-72, pis. 1-10. Roughley. T. C., 1936. ‘Wonders of the Great Barrier Reef. 282 pp. (Angus and Robertson: Sydney). Scheer. G. and Pillai, C. S. G., 1974. Report on the Scieractinia from the Nicobar Islands. Zoologica 122: 1-75 33 plates. Stephenson. W., and Wells. J. W., 1956. The corals of Low Isles, Queensland, August 1954, Paps. Dept. Zool. Vniv, Queensland 1(4): 1-59, 7 plates. Stl'Der. T., 1878. Zweite Abtheilung der Anthozoa polyactinia, welche wahrend der Reise S.M.S. Corvette Gazelle un die Erde gesammelt wurden. Monatshr. K. Akad. Wissensch. Berlin 1878: 525-50, 5 pis. 1880. Beitrage zur Fauna der Steinkorallen von Singapore. Mitt, der Natur. Gesellschaft. Bern (1880): 15-53. 1901. Madreporarien von Samoa, den Sandwich Inseln und Laysan. Zool. Jahrb Abt. Sept. 14(5): 388-428, pis. 23-31. Thiel. M. E., 1932. Madreporaria. Zugleich ein Versuch einer Vergleichenden Oekologie der gefunden Formen. Mem. Mus. Roy d’hist. Nat. Belgique 2(12): 1-177. 1933. Ueber Einige Korallen von den Philippinen nebst Bernerkungen ueber die Systematik der Gattung Acropora. Bull. Musee Royal d’Hist. nat. Belgique 9(36): 1-37. Umbgrove. J. H. F., 1939. Madreporia from the Bay of Batavia. ZoologischeMededeelingen 22: 1-64, 8 plates. 1940. Madreporaria from the Togian reefs (Gulf of Tomini, North Celebes). Zoologische Mededeelin- gen 22: 265-310, 15 plates. Vaughan. T. W., 1907. Recent Madreporaria of the Hawaiian Islands and Laysan. Bull. U.S. Nat. Mus. 49(a): 1-427, pis. 1-96. 1918. Some shoal- water corals from Murray Islands, Cocos-Keeling Islands and Fanning Islands. Pap. Dep. mar. Biol. Carnegie Inst. Washington 9: 51-234, pis. 20-93. Verrill, a. E., 1864, List of the polyps and corals sent by the Museum of comparative zoology to other institutions in exchange, with annotations. Bull. Harvard Coll. Mus. Comp. Zool. 3: 29-60. 1866. Synopsis of the polyps and corals of the North Pacific Exploring Expedition. 1853-1856, III. With descriptions of some additional species from the West coast of North America. Comm. Essex Inst. 5: 17-50, 2 plates. 1869. Synopsis of the polyps and corals of the North Pacific Exploring Expedition, 1853-1856, IV. Comm. Essex Inst. 6: 51-178, 3 plates. 1901. Variations and nomenclature of Bermudian, West Indian and Brazilian reef corals, with notes on various Indo-Pacific corals. Trans. Connecticut Acad. ArtsSci. 11 : 163-68. 1902. Notes on corals of the genus Acropora (Madrepora Lam.) with new descriptions and figures of types, and of several new species. Trans. Connecticut Acad. Arts Sci. 11 : 207-66, 7 plates. Wallace. Carden C., 1975. Distribution patterns of the coral genus Acropora on the reef slope: a preliminary report. Proc. Crown-of-Thorns starfish Seminar, Brisbane. 6 September 1974: 81-107. (Australian Government: Canberra). Wallace. Carden C., and D.\le. M. B., 1977. An information ansis approach to distribution of the coral genus Acropora on the reef slope. Atoll Research Bull: in press. Wallace. Carden C. and Lovell, E. R., 1977. Topography and coral distribution of Bushy and Redbill Islands and surrounding reef. Great Barrier Reef, Queensland. Atoll Research Bull. 194: 1-22, 4 plates. Wells. J. W., 1936. The nomenclature and type species of some genera of recent and fossil corals. Am. Jour. Sci. (5) 31 : 97-134. 1950. Reef corals from the Cocos-Keeling Atoll. Bull. Raffles Mus. 22: 29-48, pis. 9-14. 1954. Recent corals of the Marshall Islands. Geol. Survey Prof. Paper 260 - 1 : 385-486, pis. 94-185. 1955. Recent and subfossil corals of Moreton Bay, Queensland. Pap. Zool. Vniv. Queensland 4(10): 1-23. 1956. Scieractinia. pp.F328-F478 in Moore, R. C. (ed.) ‘Treatise on Invertebrate Paleontology’ part F. Coelenterata. (Geol. Soc. Amer). 318 MEMOIRS OF THE QUEENSLAND MUSEUM INDEX TO SPECIES Page Plates A. abrotanoides 280 47 A. aculeus 295 75, 76 A. aspera 286 48, 60a, 81 A. austera 314 65. 103 A. breuggemanni 315 50 A. carduus 308 93. 94 A. cerealis 297 77 A. clathrata 302 53. 64, 75, 86 A. cyiherea 289 44, 53, 64, 66, 67, 70 A. delicatula 292 69 A. digitifera 301 48, 65, 84 A. divaricata 303 87, 88, 93 A. diversa 298 79, 80 A. echinata 307 A. elseyi 311 97 A. florida 306 64, 90, 91, 92, 93 A. formosa 282 51, 52, 53, 64, 65, 75 A. grand is 281 50 A. granulosa 313 101, 102 A. haimei 293 53, 70, 71 A. horrida 284 55, 56 A. humilis 300 44. 65, 81, 82, 83 A. hyacinthus 288 44. 53, 64, 65, 66, 70, A. intermedia 280 48, 49 A. longicyathus 309 93. 95 A. microphthalma 310 96 A. millepora 291 43, 68, 81, 84 A. multiacuta 301 85 A. nasuta 297 78 A. pulchra 285 58, 59, 60 A. robusta 278 44, 45, 81 A. rosaria 311 97 A. rotumana 279 46 A. sarmentosa 305 89 A. splendida 283 53, 54, 90, 93 A. squarrosa 312 98, 99, 100 A. subglabra 307 94 A. tenuis 294 43, 72, 73 A. tubicinaria 295 74 A. variabilis 299 44, 80 A. vaughani 285 57 WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF 319 Fig. 3: Diagrammatic representation of Plate 43C indicating features used in the description of species. MEMOIRS OF THE QUEENSLAND MUSEUM Plate 43 Features of Acropora displayed by scanning electron microscopy. A. Side view of a branch tip (of A. mitlepora) from which tissues have been removed to display skeleton. B. Side view of a branch tip (of A. tenuis) from which skeleton has been removed, to display soft tissues. The specimen is in a retracted state, so that tentacles are withdrawn. C. Face view of skeletal branch tip of A. millepora. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 43 MEMOIRS OF THE QUEENSLAND MUSEUM Pi.ATi-; 44 Acropora robusta (Dana, 1846) on Bushy-Rcdbill Reef. A. Large colony on SW. reef crest showing distorted humps, some vertical cones, and some free branching. Other corals in the picture are Pocillopora. B. (Left hand side) colony on SW. reef crest with vertical cones and free branching. Other Acropora in the picture are A. hyacinthus (upper centre), A. variabilis (centre) and several A. humilis. C. Small colony on SE. (weather) side of reef (arrow), with A. cuneata (centre), A. humilis (upper centre) and Pocillopora. MEMOIRS OF THE QUEENSLAND MUSEUM Pl.ATF. 45 Acropora robusta (Dana. 1846) A, B. Free branching portion of colony, G 10191 . Big Broadhurst reef. SW. slope, 1-5 m, colour green, pink tips. C, D. Sturdy vertical humps from colony, G 10259, Bushy-Redbill Reef, W. slope. 10 m. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 45 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 46 Acropora rotumana (Gardiner, 1898) A, B. Specimen from branch lip, G10235, Big Broadhurst Reef, SW. side, outer reef flat, colour pale green. C, D. Two branches of colony, G10263, Bowden reef, SW. side,’ reef crest, colour pink-brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 46 MEMOIRS OF THE QUEENSLAND MUSEUM Pl,ATl^ 47 Acropora ahroianoides (Lamarck, 1816) A, B. G 10239 Bushy-Redbill reef, NW. side, reef crest, colour yellow-brown. C, D. G 10238 Big Broadhurst reef, upper surface of a patch reef in the lagoon, colour yellow-brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 47 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 48 Acropora intermedia (Brook, 1891) on Bushy-Redbill Reef. A. Large thicket in deep water middle reef flat on leeward (W.) side. B. Small patches with A. digitifera (centre) on reef crest, SW. side. C. Small patch (left) with A. aspera on inner middle reef flat, SW. side. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 48 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 49 Acropora intermedia (Brook, 1891) A. B. G1 1400 Big Broadhurst Reef, SW. slope, 2 m, colour bright blue. C, D. G11295 Big Broadhurst Reef, SW. slope, 2 m, colour pale green. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 49 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 50 A. B. Acropora grandis (Brook, 1892) G11298 Darley reef, patch reef in lagoon, 4 m, colour creamy brown. C, D. Acropora brueggemanni (Brook, 1893) G 11498 Big Broadhurst Reef, SW. slope, 6 m, colour brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 50 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 51 Acropora formosa (Dana, 1846) on the reef. A. Large thickets on sandy bottom in deep middle reef flat, Bushy-Redbili Reef, leeward (W.) side. B. Thicket (foreground) with diverse Acropora assemblage in background, base of upper reef terrace. Big Broadhurst Reef SW. side. WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 51 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 52 Acropora formosa (Dana, 1846) A, B. G11339, Big Broadhurst reef, 2 m, colour pink-grey. C, D. G11381 Bowden Reef, floor of opening in SW. side, 5m. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 52 MEMOIRS OF THE QUEENSLAND MUSEUM Plath 53 Acropora splendida Nemenzo, 1967 on Big Broadhurst Reef, SW. side. A. Colony on gently-sloping upper reef terrace (about 7 m) surrounded by flat-plate species. The distinctive ‘arborescent- bracket’ shape can be seen. B. Colony in a mixed assemblage at about 10 m (centre of plate). This colony is flatter than the previous, and its shape is somewhat disguised by another arborescent colony (A. formosa, lower left and right). The small flat colony upper left of A. splendida is A. clathrata: the large plate above this is A. hyacinthus, and above it to the right is A. cytherea. The small dense bushy colony (upper left) is A. haimei. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 53 MEMOIRS OF THE QUEENSLAND MUSEUM Pl.ATF 54 Acropora splendida Nemenzo, 1967. A, B. Specimen from inner part of large bracket, G8699, Big Broadhurst Reef, SW. slope, 6 m, colour brown, pale blue tips to branches. C, D. Specimen from outer part of bracket, G8701, Big Broadhurst Reef, SW. slope, 7 m, colour olive green, pale green tips to branches. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 54 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 55 Acropora horrida (Dana, 1846) A. Shrub like formations (centre) with soft corals in deep water leeward (W) middle reef flat on Bushy-Redbill Reef. B. ‘Bottlebrush* branch from C, below. C. Sprawling arborescent formation on sandy bottom, 15 m depth, ofUreef floor. Bushy-Redbill Reef, NW. side. * 4 % WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 55 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 56 Acropora horrida (Dana, 1846) A, B. Piece of slirub-like colony, G9078, Bushy-Redbill Reef, NW. side, middle reef flat, colour lavender-grey. C, D. Piece of colony, open arborescent, proliferous with short branchlets, 09077, Bushy-Redbill Reef, NW. side, floor outside slope, 8 m, colour pale blue. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 56 MEMOIRS OF THE QUEENSLAND MUSEUM pLATi-; 57 Acropora vaughani Wells, 1954. A, B. G 10270, Big Broadhurst Reef, SW. slope, 7 m, colour pale brown. C, D. G 1027 1 Big Broadhurst Reef, SW. slope, 8 m, colour pale brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 57 MEMOIRS OFTHEQUEENSLAND MUSEUM Pl,ATE 58 Acropora pulchra (Brook, 1891) A, B. G1 11 18, Heron Island reef flat. Low corymbose colony with slender branches. C, D. Gill 16 Heron Island, W. side, reef flat, colour cream-brown, blue tips. Corymbose colony with thick branches. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 58 MEMOIRS OF THE QUEENSLAND MUSEUM P\ .\Ti[ 59 Acropora pulchra (Brook, 1891) A, B. Glini Heron Island Reef flat. Compact arborescent colony. C, D. Specimen from reef flat adjacent to Redbill Island. Open arborescent colony. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 59 MEMOIRS OF THE QUEENSLAND MUSEUM Pl,ATE 60 Acropora aspera (Dana, 1846) on the reef. A, B. Microatoll type development of corymbose colonies on middle reef flat. Heron Island reef, W. side. (Scale divisions are 2 cm). C. Large thicket (left half of picture) in contact with thicket of A. pulchra (right hand side) on inner middle reef flat, Bushy-Rcdbill Reef, SW. side. rfM- WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 60 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 61 Acropora aspera (Dana, 1846) A, B. G1096I, Bushy-Redbill Reef, N. side, inner flat-algal bank area, colour brown, blue tips. C, D. G 10930 Big Broadhurst Reef, patch reef in lagoon, colour pale green. (Specimen typical of ‘A. hebes'). WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 61 \ MEMOIRS OF THE QUEENSLAND MUSEUM Plate 62 Acropora aspera (Dana, 1846) A, B. G 10929 Big Broadhurst Reef, patch reef in lagoon, colour pale green. C, D. G 10921 Heron Island reef flat, colour pale brown. (Specimen typical of A. aspera 5 . 5 .) WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 62 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 63 Large flat plates (Acropora hyacinthus. A. cytherea and A. clathrata) and arborescent species, on Big Broadhursl Reef. (N.B. species cannot be distinguished). A. On upper reef terrace. B. On surge channel floor. Note the change in arborescent to flat plate species ratio between this and the reef terrace in the background. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 63 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 64 Large flat plate species on the reef. A. Acropora hyacinthus (upper left), A. cytherea (lower left), A. clathrata (centre). With A. florida (left of centre) and arborescent species on the surge channel floor (approx. 10 m) at Big Broadhurst Reef. B. A. hyacinthus with other Acropora in the Darley Reef lagoon. Note oblique additions to the original flat plate. C. A. clathrata (centre) with A. hyacinthus (right) and A. formosa (left) on the reef crest at Bushy-Redbill Reef. D. Three juvenile A. cytherea colonies on the reef crest at Bushy-Redbill Reef. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 64 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 65 Acropora hyacinthus (Dana, 1846) on Bushy-Redbill Reef. A. Deep middle reef flat area on leeward (W.) reef (1-5 m at low tide^ with A. formosa. B. Outer reef flat, SW. side. Several colonies with A. digitifera (centre and lower centre) and A. formosa. C. Reef crest, SW. side. Two colonies (arrows) with A. austera (centre left), A. digitifera (centre) and A. humilis (centre right). WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 65 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 66 Acropora hyacinthus (Dana, 1846) and A. cytherea (Dana, 1846) A, B. A. hyacinthus portion of colony, G9865 Big Broadhurst Reef, SW. side, reef slope, 5-3 m, colour pink-brown. C, D. A. cytherea portion of colony, G9856 Big Broadhurst Reef, SW. side, reef slope, 3 m, colour pink-brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 66 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 67 Acropora cytherea (Dana, 1846) A. G7291 Flinders Reef, Moreton Bay, SE. Queensland. B. G9855 Big Broadhurst Reef, SW. slope, 5 m, colour pale pinkish brown. C. G9849 Big Broadhurst Reef, SW. slope, 9*7 m, colour yellow. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 67 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 68 Acropora millepora (Ehrenberg, 1834) A, B. G11064 Bowden Reef, slope of opening in SW. side, 2 m, colour orange. C, D. G11049 Bushy-Redbill Reef, NW. side, outer reef flat, colour pale brown, yellow tips. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 68 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 69 Acropora delicatula (Brook, 1891) A, B. GI1441 Big Broadhurst Reef, SW. slope, 6 brown. C. G11435, Big Broadhurst Reef, SW. slope, I i, colour pale m. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 69 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 70 Acropora haimei (Milne Edwards and Haime, 1860) A. Compact arborescent clump on gentle reef slope on SW. side of Big Broadhurst Reef, about 5 m depth, (centre of picture) with flat plate species A. hyacinthus and A, cyfherea, and other arborescent species. B. Turf like patch on middle reef flat at Bushy-Redbill Reef, SW. side, with soft corals, Seriatopora (lower left) and Acropora palifera (lower right). WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 70 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 71 Acropora haimei (Milne Edwards and Haime, 1860) A, B. G 10222 Big Broadhurst Reef, SW. slope, 4-2 m, cream. C, D. G 102 1 7 Big Broadhurst Reef, SW. slope, 9-8 m, colour colour cream. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 71 20mm 10mm 30mm MEMOIRS OF THE QUEENSLAND MUSEUM Pl.ATE 72 Acropora tenuis (Dana, 1846) on the leeward (W.) side of Bushy-Redbill Reef. A. Corymbose plate (arrow) on outer middle reef flat. With sponges and soft coral. B. Layered colony in middle reef flat area, sandy bottom. With A. palifera, Pocillopora and Seriatopora. C. Layered colonies on reef crest. m WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Platc 72 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 73 Acropora tenuis (Dana, 1846) E. G11418 Darley Reef, patch reef in lagoon, colour pale brown. GI1428 Big Broadhurst Reef, SW. slope, 6 m, colour pink-grey. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 73 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 74 Acropora tubicinaria (Dana^ 1846) A, B. G11072, Bushy-Redbill Reef, W. side, inner reef flat, colour pale brown. C, D. G 11077, Bushy-Redbill Reef, W. side, inner reef flat, colour pale brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 74 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 75 Acropora aculeus (Dana, 1846) on the leeward side of Bushy-Redbill Reef A. Plate like colony on middle reef flat (arrow) with soft corals and sponges. B. Layered colony on reef crest (bottom of picture), with A. formosa (left), lA. haimei (right), A. palifera (centre), A. clathrata (plate) and other corals. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 75 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 76 Acropora aculeus (Dana, 1846) A, B. From Big Broadhurst Reef, SW. slope, about 10 m. Small, thin plate specimen. D. Large tiered plate colony in deep water middle reef flat area, Bushy-Redbill Reef, W. side. C. Portion of a large tiered plate. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Platf 76 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 77 Acropora cerealis (Dana, 1846) A, B. G9519 Bowden Reef, slope of opening in SW. side, 3m, colour blue. C. D. G9529 Big Broadhurst Reef, SW. slope, 8-7m, colour cream. E, F. G9526 Darlcy Reef, patch reef in lagoon, 3m, colour pale brown-lavender. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 77 ^ o MEMOIRS OF THE QUEENSLAND MUSEUM Plath 78 Acropora nasuta (Dana, 1846) B. Colonies in the deep middle reef Hat of Bushy-Redbill Reef, . side with sandy bottom and soft corals. 11224 Big Broadhurst Reef, SW. slope, 7m, colour grey. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 78 MEMOIRS OF THE QUEENSLAND MUSEUM Pi.ATi-: 79 Acropora diversa (Brook, 1891) A, B. Gl 1246 Big Broadhurst Reef, SW. slope, 8m, colour purple, cream tips. C, D. G11245 Big Broadhurst Reef, SW. slope l-3m, colour cream. WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 79 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 80 A, B. A. diversa (Brook, 1891) G 11236 Big Broadhurst Reef, SW. slope 5-1 m, colour blue-brown. C, D. A. variabilis (Klunzinger, 1879) G11290 Big Broadhurst Reef, SW. side, outer reef flat, colour cream. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 80 MEMOIRS OF THE QUEENSLAND MUSEUM Plate-. 81 Acropora humilis (Dana, 1846) and A. millepora (Ehrenberg, 1834) on the reef. A. Reef edge at Big Broadhurst Reef, SW. side, looking back across the reef flat. A. rohusta and A. hyacinthus in the foreground: most of the other colonies are A. humilis. B. A. humilis colony on sandy bottom, deep middle reef flat, Bushy-Rcdbill reef, W. side. This colony could be regarded as forma a — . forma />*. C. Outer reef flat on windward side (SE.) of Bushy-Redbill Reef. Colony in centre may be regarded as A. humilis forma ; . and that to its left as A. humilis forma P . To the right is another low A. humilis. and other corals are A. palifera and Pocillopora. D. A. millepora (centre) on reef flat adjacent to Redbill Island; to its left is A. humilis forma V , and another small A. millepora colony. To the right is A. aspera. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 81 MEMOIRS OF THE QUEENSLAND MUSEUM Pi.Aii- 82 Acropora humilis (Dana, 1846) A, B. G1 161 1 Darley Reef, patch reef in lagoon, 5m, colour pale brown, lavender tips. Forma a Wells. C, D. G11189 Bowden Reef, slope of opening in SW. side. Forma (\ Wells. WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 82 MEMOIRS OF THE QUEENSLAND MUSEUM Pl.ATH 83 Acropora humilis (Dana, 1846) (Specimens not mentioned in text). A. B. 011 150 Big Broadhurst Reef, weather (E.) side, 2m, colour blue. A. humilis forma v Wells. C. D. Gil 142 Big Broadhurst Reef, SW, slope, 2m. Forma ^ » forma ft Wells. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 83 MEMOIRS OF THE QUEENSLAND MUSEUM Plath 84 Acropora digitifera (Dana, 1846) A. Colonies on outer reef flat adjacent to Redbill Island (small colony centre left is A. milleporaj B, C. G1 1 169 Big Broadhurst Reef, outer reef flat, colour cream, blue tips. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 84 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 85 Acropora multiacuta Nemenzo, 1967. All specimens from Darley Reef, patch reef in lagoon, all pale blue. A. G1047!. B. G10468. C. D. G6721. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 85 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 86 Acropora clathrata (Brook, 1891) A, B. G9762, whole colony, Bowden Reef, SW. side, reef slope Im, colour lavender-brown. C, D. G9753, portion of colony, Big Broadhurst Reef, SW. side, reef slope, 5ni, colour yellow-brown. E, F. G9750, portion of colony. Big Broadhurst Reef, SW. side, reef slope, 5m, colour brown, with mauve tips to branchlets. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 86 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 87 Acropora divaricata (Dana, 1846) (C is a whole colony, others are portions of colonies). A. G9162 Big Broadhurst Reef, SW. slope, 124m, colour brown, blue tips. B. G9181 Big Broadhurst Reef, SW. slope, 7m, colour brown, blue tips. C. G9182, Darley Reef, patch reef in lagoon, 7m, colour brown, lavender tips. D. G9173 Big Broadhurst Reef, SW. slope, 8-7m, colour brown. E. G9174 Big Broadhurst Reef, SW. slope, 8-6m, colour brown, blue tips. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 87 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 88 Acropora divaricata (magnified views of specimens in plate 87) A. G9162, see Plate 87A. B. G9181, see Plate 87B. C. G9173, see Plate 87D. D. G9174, see Plate 87E. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 88 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 89 Acropora sarmentosa (Brook, 1892) A. B. G9057, Big Broadhurst Reef, SW. slope, 7-6 m, colour cream-brown. C, D. G9059, Big Broadhurst Reef, SW. slope, 9-7 m, colour green, pink-grey tips. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 89 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 90 Acropora florida (Dana, 1846) on Big Broadhurst Reef. A. In mixed assemblage on floor of surge channel at 10 m (centre). Rounded, open-arborescent colony. Branches are round in cross-section, secondary branchlets are evenly distributed and sized. Coral in bottom left corner is A. splendida. B. Colony on reef slope, at about lOm (arrow). Branches here are more or less horizontal, and are flattened in cross-section. Secondary branchlets arc differentially developed on the upper surface. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 90 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 91 Acropora florida (Dana, 1846) A, B. G8651. portion of colony. Big Broadhurst Reef, SW. slope, 5- 8m, colour brown. C. G8665, whole colony. Big Broadhurst Reef, SW. slope, 6m, colour pink-brown. D. G8657, portion of colony. Big Broadhurst Reef. SW. slope, 6- 6m, colour green-brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 91 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 92 Acropora JJorida (Dana, 1846) A, B. G8655 Big Broadhurst Reef, SW. slope, , 6-6m, colour pale. pinkish brown. (This specimen resembles A. florida C, D. G8647 Big Broadhurst Reef, SW. slope, 9-3m (flat plate specimen). WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 92 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 93 Acropora echinata group on Big Broadhurst Reef, SW. side. A. On surge channel floor at 10m. A. carduus (Dana, 1846). (arrow) with a small table (7^4. cytherea) and A. Jlorida (surrounding). B, On gentle reef slope, about 12m. A. longicyathus (Milne Edwards and Haime, I860) (arrow). Above this is A. splendida. and A. divaricaia, upper right. WALLACE; THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 93 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 94 A, B. Acropora subglahra (Brook, 1891) G10714 Lizard Island fringing reef, colour pinkish brown. C, D. Acropora carduus (Dana, 1846) G10731 Big Broadhurst Reef, SW. slope, 8m, colour brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 94 MEMOIRS OF THE QUEENSLAND MUSEUM Pi. ATI-: 95 Acropora longicyathus (Milne Edwards and Haime. I860) A. C. G 10767 Big Broadhurst Reef, SW. slope, 7 m, colour brown. Portion of colony from similar location. B. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 95 MEMOIRS OF THE QUEENSLAND MUSEUM Pi. ATE-; 96 Acropora microphfhalnia (Verrill. 1869) A, B. G8685. Big Broadhurst Reef, SW. slope, 6m, colour brown. C, D. From Big Broadhurst Reef, SW. slope. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Pi.ATt: 96 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 97 A, B. Acropora elseyi (Brook) G 10793, Bushy-Redbill Reef, W. side, middle reef flat, 1 m, colour yellow. C, D. Acropora rosaria (Dana) G9202, Palm Islands, 1929, T. C. Marshall. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 97 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 98 Acropora squarrosa (Ehrenberg, 1834) on the reef. A. Bottlebrush branching colony on reef crest, Bushy-Redbill Reef, W. side, colour blue (arrow). B. Bottlebrush branching colony, (with shortened brushes) on edge of hard/soft coral patch, middle reef flat, Bushy-Redbill Reef, W. side, colour yellow-brown (arrow). C. Caespitosc ‘incipient bottlebrush' colony on lagoonal patch reef, Darley Reef (arrow). D. Bottlebrush branching colony on sand, deep middle reef flat, Bushy-Redbill Reef, W. side, colour lavender-blue. E. Bottlebrush thicket in deep water reef flat at Wistari reef, near Heron Island, (photo courtesy D. Ross Robertson). F. Caespitose colony on lagoonal patch reef, Darley reef, colour pale blue. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 98 MEMOIRS OF THE QUEENSLAND MUSEUM Pl.ATE- 99 Acropora squarrosa (Ehrenberg, 1834) A, B. G10782 specimen comparable to 'A. murrayensis’ Darley Reef, patch reef in lagoon, 1 m, colour lavender. C, D. G 10842 specimen approaching A. syringodes’ (part). Big Broadhurst Reef, SW. side, reef slope, 9-2 m, colour cream. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 99 MEMOIRS OF THE QUEENSLAND MUSEUM PiATi: 100 Acropora squarrosa (Ehrenberg, 1834) A, B. G 10846 low cacspitosc specimen. Dariey Reef, patch reef in lagoon, 3 m, colour patches ol pale yellow, lavender and blue. C, D, E. G10845 Hal plate specimen (D is undersurface). Big Broadhurst Reef, SW. side, reef slope, 12-9 m, colour pale blue-brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 100 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 101 Acropora granulosa (Milne Edwards and Haime, I860). Specimens with blunt axial corallites. A, B. Gl 1487, Big Broadhurst Reef, SW. side, reef slope, 10-4 m, colour cream-grey. C, D. GI1492, Big Broadhurst Reef, SW. side, reef slope, 8 m, colour brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 101 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 102 Acropora granulosa (Milne Edwards and Haime, 1860). Specimens with tapering axial corallites. A, B. G6723 (comparable with 'A. speciosa'}. Darley Reef, patch reef in lagoon, 2*5 m, colour pale brown. C, D. GI 1476 (comparable with ‘A. raynerV). Big Broadhurst Reef, SW. side, reef slope, 5 m, colour pale brown. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 102 MEMOIRS OF THE QUEENSLAND MUSEUM Plate 103 Acropora austera (Dana, 1846) A, B. G 10828 Big Broadhursl Reef, SW. slope, 7-8 m, colour cream. C, D. G10810 Bushy-Redbili Reef, S. side, reef slope, 3m, colour lavender. WALLACE: THE GENUS ACROPORA IN THE GREAT BARRIER REEF Plate 103 I CONTENTS Pag( Bartholomai. Alan The Macropodidae (Marsupialia) from the Allingham formation, northern Queensland. Results of the Ray E. Lemley Expeditions, Part 2 127 Bartholomai. alan The rostrum in Palorchestes Owen (Marsupialia: Diprotodontidae). Results of the Ray E. Lemley Expeditions, Part 3 ]45 CovACEVicH. J. and Ingram. G. J. An undescribed species of rock dwelling Cryptoblepharus (Lacertilia: Scincidae) 151 Archfr, Michall The nature of the molar-premolar boundary in marsupials and a reinterpretation of the homology of marsupial cheekteeth 157 Archlr. MICHAEL and Brayshaw. Helen Recent local faunas from excavations at Herveys Range, Kennedy, Jourama, and Mount Roundback, north-eastern Queensland 165 Cannon. L. R. G. Pterastericola vivipara n. sp., a parasitic turbellarian (Rhabdocoela: Pterastericolidae) from the Crown-of-Thorns starfish, Acanthaster planci 179 Stephenson. W., Cook, S. D., and Newlanos. S. J. The macrobenthos of the Middle Banks area of Moreton Bay 185 Greenwood, J. G. A new species of Pontellopsis (Copepoda, Calanoida) from Moreton Bay, Queensland 213 Jeli . Peter A. Asihenopsis Whitehouse, 1939 (Trilobita, Middle Cambrian) in northern Australia 219 Bruc E. A. J. Paranchistus pycnodontae sp. nov., a new pontoniine shrimp associated with an ostreid bivalve host 233 Jones. H. I. Gastrointestinal nematodes from aquatic Australian snakes 243 Philups, W. j. Some parasitic barnacles (Rhizocephala: Sacculinidae) from portunid crabs in Moreton Bay, Queensland 255 Gh l. j. C. H. The old burial vault at North Quay, Brisbane 265 Wallace. Carden C. The coral genus Acropora (Scleractinia: Astrocoeniina: Acroporidae) in the central! and southern Great Barrier Reef Province 273