Balanoid Barnacles of the Genus Hexaminus (Archaeobalanidae: Elmiinae) from Mangroves of New South Wales, Including a Descriptio
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AUSTRALIAN MUSEUM SCIENTIFIC PUBLICATIONS Anderson, D. T., J. T. Anderson and E. A. Egan, 1988. Balanoid barnacles of the genus Hexaminius (Archaeobalanidae: Elminiinae) from mangroves of New South Wales, including a description of a new species. Records of the Australian Museum 40(4): 205–223. [16 December 1988]. doi:10.3853/j.0067-1975.40.1988.155 ISSN 0067-1975 Published by the Australian Museum, Sydney naturenature cultureculture discover discover AustralianAustralian Museum Museum science science is is freely freely accessible accessible online online at at www.australianmuseum.net.au/publications/www.australianmuseum.net.au/publications/ 66 CollegeCollege Street,Street, SydneySydney NSWNSW 2010,2010, AustraliaAustralia Records of the Australian Museum (1988) Vol. 40: 205-223. ISSN 0067 1975 205 Balanoid Barnacles of the Genus Hexaminius (Archaeobalanidae: Elminiinae) from Mangroves of New South Wales, including a Description of aNew Species. D.T. ANDERSON, J.T. ANDERSON AND E.A. EGAN School of Biological Sciences, University of Sydney, Sydney, NSW 2006, Australia ABSTRACT. Hexaminiusfoliorum n.sp. is described and separated from H popeiana Foster on differences in adult and larval anatomy, supported by differences in cirral activity, copulatory activity and breeding. Structure, function and reproduction in Hfoliorum are more specialised than in H popeiana. The differences are related adaptively to the occupancy by H popeian.a of a variety of hard substrata, but not mangrove leaves, and the confinement of H foliorum to the ephemeral habitat of immersible mangrove leaves. Hexaminiusfoliorum may be a specialised descendant of H popeiana. ANDERSON, D.T., J.T. ANDERSON & E.A. EGAN, 1988. Balanoid barnacles of the genus Hexaminius (Archaeobalanidae: Elminiinae) from mangroves of New South Wales, including a description ofa new species. Records of the Australian Museum 40(4): 205-223. Foster (1982) revised the taxonomy of the high found at their study site (Iron Cove, Port Jackson, shore estuarine barnacles of New South Wales, NSW) on two distinct surfaces, rocks and the lower hitherto referred to as Elminius modestus Darwin leaves of the mangrove Avicennia marina. The (Pope, 1945). The majority of the four-plated settlement of this species on mangrove leaves was not specimens were referred by Foster to a new species, mentioned by Foster (1982), nor, indeed has this part Elminius covertus. Foster also observed that E. of"the mangrove surface been recorded as a habitat covertus is intermingled with a six-plated for other balanomorphs in Australia, although a archaeobalanid which he described and named as number of species are known to inhabit mangrove Hexaminius popeiana. An archaeobalanid subfamily trunks and pneumatophores (Hutchings & Recher, Elminiinae was erected by Foster in recognition of 1982; Achituv, 1984). The observation was therefore the close relationship between Hexaminius and of sufficient interest to prompt Egan and Anderson Elminius, a concept further supported by Buckeridge to make parallel studies during 1982-1984 of the (1983) and by Egan & Anderson (1985), who seasonal breeding of populations of H popeiana described the larval development of H popeiana and from rocks and mangrove leaves at the Iron Cove E. covertus. The larvae of E. covertus are close to but site, while utilising the mangrove leaf individuals as a different from those of E. modestus. The larvae source of nauplii for larval culture. Initially it was described by Egan & Anderson (1985) under the assumed that the differences in external appearance name H popeiana are more like those of the Conopea between the mangrove leaf and rock populations group of archaeobalanines, but also share sufficient were a consequence of a greater level of erosion of the features in common with the larvae of Elminius to rock-dwelling individuals. However, a study of lend further weight to the Elminiinae as a distinct seasonal cycles during 1982-1984 showed that the subfamily. two populations had different breeding patterns. The During their study of larval development, Egan mangrove leaf population, as illustrated by Egan & and Anderson noted that H popeiana were to be Anderson (1985), bred throughout the year, with 206 Records of the Australian Museum 40 11') o ....>- ..D E Q) ....c.... 11') '+-o o Z --------1982------ ------ 1 98 3 --------- - 1 984 - o No embryos f2J Embryo stages 11 - IV • Embryo stage V Fig.I. Percentages of individuals of rock-dwelling Hexaminius popeiana brooding embryos of different developmental stages, in monthly samples from Iron Cove, Port Jackson, NSW from 1982 to 1984. Sample size is given at the top of each column. The embryo stages are those of Sandis on (1954). some reduction in the autumn months. The rock H. popeiana Foster, inhabits surfaces of a permanent population, in contrast, built up through the winter or semipermanent nature on the upper shore of to a spring breeding peak, and showed little or no sheltered estuarine locations, including rocks, breeding during the summer and autumn (Fig. I). wooden structures and the trunks of mangrove The question therefore arose, whether the external bushes. The other species, newly described here, is differences between the leaf-dwelling and rock- confined to the leaves of Avicennia marina. Much of dwelling animals were indicative of separate the present paper is devoted to a description of the breeding populations. We have now made a thorough new species. We also provide a more extended morphological comparison of animals from the two description of H. popeiana than was given by Foster substrata, utilising animals from a number of sites (1982). Many of the features which differ between along the New South Wales coast, some of which the two species were not mentioned in the initial were also used as collection sites by Foster (1982). In description by Foster. view of the apparent association of one of our The description of the larval stages of H. popeiana breeding populations with the leaves of Avicennia given by Egan & Anderson (1985) was based on marina, we took particular care also to collect larvae obtained from mangrove leaf populations, specimens from the trunks of mangroves as well as and is therefore referable to the new species. As from the leaves, and from rock surfaces in each supplementary evidence of the differences between locality. the species, we include in this paper a comparative This comparison has shown that there are two account of the larvae of H. popeiana (s.s.), which are species of H examinius in the estuaries of the central similar in general but different in detail from those of coast of New South Wales. One ofthem, referable to H. n. sp. We conclude our account with an analysis of Anderson, Anderson & Egan: Hexaminius 207 the cirral activities and copulatory activities of H. complete shell 190). PARATYPES hermaphrodite, sexually popeiana and H. n. sp. These activities also differ in mature, AM P (spec. D25, Fig. 2e, colI. 14 May 1984, 189), ways that support the morphological analysis. from Sisters Bay, Iron Cove, NSW, 33°51.5'S 151 °09.5'E, on leaves of Avicennia marina, collected by D.T. Anderson, Aug 1984. Materials and Methods Additional material examined. Part specimens: DA All specimens used for anatomical studies were bleached plates (Fig. 2b,c,d), 212; body (Fig. 2f), 59; spe~. collected in August and September 1984, at the Dl, oral cone (Fig. 4a), 54A; labrum (Flg. 4b), 65; DB p~ms (Fig. 4g), 217. Slides: cirri I-VI, D22, .13 May 1~83, ng.ht height of the breeding season for H. pOI!eiana. a~d cirri, 85; spec. D5, right palp and nght mandlble (FIg. within the long period of maximum breedmg actIvIty 4c,d), 19; spec. D5, left maxillule and left maxilla (fig. 4e,f), of H. n.sp. All specimens were therefore sexually 17. Hexaminiusfoliorum was also collected from the lower mature. Observations on cirral activity and (immersible) leaves of Avicennia marina at Gosford and copulatory activity (see below) were also made at the Woy Woy, on Brisbane Waters, and at Hawks Nest, Port time. Larval cultures of H. popeiana (see below) were Step hens; all localites on the Central Coast of NSW reared in August and September 1984. between 33-34°S. Thirty individuals of each species, collected from Diagnosis. Hexaminius of low conical f~rm; b~sal Iron Cove NSW were investigated anatomically by margin ovoid with bluntly protrudmg nbs; the dissedtion of fresh specimens and specimens conspicuous reddish brown radial bands externally. preserved in 5% formalin in seawater (V/v). Limbs Wall plates and opercular plates thin, fragile. Wall and mouthparts were mounted in polyvinyl plates with thin basal margins; basis m~mbranous, lactophenol after staining in lignin pink. Wall plates thick, semiopaque. Tergoscutal flaps ":Ith shallow and opercular plates were disarticulated and cleared carinal fold. Scutal depressor muscles thm. Prosoma using sodium hypochlorite. Podomere counts were elongate, fusiform; cirri IV-VI with long rami and made for all limbs. long setae. Pedicel of penis not enlarged. During or subsequent to each dissection, measurements were taken as follows: basal length Description. The following description is based on and width; orifice length and width; scutum length 30 additional specimens from Iron Cove, NSW, (along occludent margin) and height (from occludent which were destroyed during the course of the study. EXTERNALS. Shell (Fig. 2a) to 7.0 mm m margin to basitergal ang~e); tergum length (from.apex to tip of spur) and width (from scutal margm to rostrocarinal basal length, low conical, even when carinobasal angle); ramal length for cirri Ill, IV-VI; crowded· basal length markedly greater than width, length (from second tooth to base) and width (across basal m~rgin ovoid. Colour when live, pale brown, cutting edge) of the mandible; length (from middle of with darker reddish brown radial bands between low ribs. Preserved, cream with brown radial ban~s.