Cnidaria, Hydrozoa, Solanderiidae)
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JOURNAL OF NATURAL HISTORY, 1988, 22, 1551-1563 Redescription and affinity of the large hydroid Chitîna ericopsis Carter, 1873 (Cnidaria, Hydrozoa, Solanderiidae) J. BOUILLON Laboratoire de Zoologie, Université Libre de Bruxelles, A venue F. D. Roosevelt 50,1050 Bruxelles, Belgium, and Station Biologique Léopold in, Laing Island, via Madang, Papua New Guinea P. F. S. CORNELIUS Department ofZoology, British Muséum ( Natural History), Cromwell Road, London SW7 5BD {Accepted 16 December 1987) Chitina ericopsis Carter, 1873, is a large, rarely-reported, arborescent hydroid with a thick internai chitinous skeleton. Ils afïinities have been unclear since its first description more than one hundred years ago. Study of new material which includes hydranths confirms that it should be referred to the family Solanderiidae which comprises just one other genus, Solanderia. The distinctions between the two included gênera are summarized. The species is redescribed, détails of hydranth morphology and cindocysts being given for the first time. Type material is identified and redescribed. KEYWORDS: Hydroid, Solanderiidae, Chitina, Solanderia, hydranths, cnidocysts, New Zealand, type material, systematics. Introduction The seldom-reported and unusually large arborescent hydroid Chitina ericopsis Carter, 1873, was originally described from colonies up to 40 cm in height collected in New Zealand waters by 'Dr Sinclair and Sir G. Grey' (Carter, 1873). Colonies of such size are unusual among hydroids and although longer colonies occur in some families in no other are the main trunks so massive. They resuit from the fusion of numerous typical hydrozoan chitinous perisarc tubes the walls of which are perforated by numerous fenestrae. This basic pattern is modified during development so that a robust, three-dimensional network results (Figs 4, 5). Since the skeletons of most athecate hydroids are rather uniform particular systematic interest attaches to the morphology of the feeding-polyps or hydranths. However, despite the conspicuousness of the colony, that of Chitina has remained undescribed until now. The genus Chitina was proposed by Carter when describing the species and has remained monotypic ever since, so that the validity of the genus has been questioned. Ail known material of the sole species has corne from New Zealand. The first-described spécimens of C. ericopsis were dried, and are preserved in the collections of the British Muséum (Natural History) [BMNH], London. Vervoort Contribution No. 119 from the Station Biologique Léopold IIL 1552 J. Bouillon and P. F. S. Cornélius (1962), studying thèse spécimens, regarded the genus Chitina as dubious since the material, being dried, did not provide sufficient information for taxonomic assessment. Other dried material, preserved in the Australian Muséum, Sydney, and evidently of the same nominal genus, was examined later by Wineera (1968) who considered it had affinity with the genus Solanderia. About seven species of this genus are recognized. He thought that the two gênera should not be separated on the basis of skeletal structure, and concluded also that C. ericopsis was almost certainly conspecific with the often- reported Solanderia misakinensis (Inaba, 1892). Recently Bouillon (1985) followed the opinion of Vervoort (1962) in regarding the nominal genus Chitina as dubious, but had no new évidence. Additional material in the BMNH stored in alcohol is sufficiently well preserved for a more detailed description of Chitina ericopsis to be made than previously possible. Consequently the validity of the genus and the affinity of the included species can be reassessed. Taxonomy Solanderiidae Marshall, 1892 The family is defined thus (after Bouillon, 1981): Colony erect, ramified, with chitinous internai anastomosing skeleton formed from coalescing and modification of adjacent hydrocauline tubes. Coenosarc covering entire colony and penetrating skeletal interstices. Hydranths borne over whole surface of colony, uniform in structure, cylindrical; with single circlet of capitate tentacles surrounding mouth and numerous similar tentacles dispersed over column. Gonophores, where known, eumedusoid sporosacs arising directly from coenosarc. Two recognized gênera are now included (Bouillon, 1974, 1985): Solanderia Duchassaing & MicheUn, 1846, and Chitina Carter, 1873. As noted by Wineera (1968) the similarity of the fine structure of the skeletons of their included species implies close affinity. However, the colony form of Chitina ericopsis differs markedly from that of virtually ail known species of Solanderia (see below). Second, the limited material of C. ericopsis available indicates that the distribution and method of insertion of the hydranths on the colonies also differs between the two (see définition). Hence it seems useful to main tain a generic séparation. The two gênera are distinguished and defined as follows : - colonies bushy, branching in ail planes; hydranths without hydrophoral structures, confined to hydrocladia Chitina - colonies fundamentally branching in one plane, roughly fan-shaped; hydranths generally accompanied by hydrophoral structures, distributed on both hydrocladia and hydrocauli Solanderia Chitina ericopsis Canei, 19^3 (Figs 1-5, 7) Chitina ericopsis Carter, 1873: 13-14; Carter, 1877: 46, 56, 60, 73; Carter, 1878: 308; Higgin, 1878: 45; Steinmann, 1878: 106; Brazier, 1887: 577; Spencer, 1892: 8, 21 ; Steinmann, 1893: 480, 482; Weltner, 1893: 15; Delage & Hérouard, 1901: 74; Hickson, 1903: 113-114; Hartlaub, 1905: 515 ; Stechow, 1909: 41 ; Bedot, 1912:271; Bedot,1916:73; Bedot, 1918: 95 ; Bedot, 1925:135; Vervoort, 1962: 510-512,536-537; Wineera, 1968:1-2; Bouillon, 1985:91. Material examined Type material. Dried, skeletal material only. Ail spécimens originally BMNH regd Redescription of Chitina ericopsis 1553 no. 1857.1.2.36 (re-registered 1962.4.15.1-3 and 1973.7.11.13); ail collected 'New Zealand' by Sir George Grey and 'Dr Sinclair' ; mentioned, Carter, 1873: 13 14.7 large erect ramified fragments, measuring approx. 38, 33, 31, 30, 23, 22, 19 cm in height; probably representing 4-5 colonies. Each colony bushy, comprising trunk-like main stem up to c. 2 0 cm diameter, polysiphonic, consisting of up to c. 40 hydrocauli running nearly straight and approximately parallel, anastomosing by short cross- connections at fréquent but greatly varied intervais. Main trunk forking successively but irregularly; resulting branches tortuous but ail continuing generally upwards, flexuose but roughly parallel. Finer branches 3-lOmm in diameter, not broadly divisible into component tubes, comprising fine interlaced three-dimensional mesh- work of chitinous threads with overall honeycombed appearance; in finest branches, positions of original coenosarc tubes marked by parallel tubular cavities running through meshwork; latter evidently laid down as network surrounding each tube, but networks of contiguous tubes shared ; no évidence of former positions of hydranths on this dried skeletal material. Other dried material. One spécimen, BMNH 1877.5.21.210, carried a printed label dating from around the first quarter of the présent century with the wording ' Australia/ Bowerbank Coll.'. We are indebted to Professer W. Vervoort, Rijksmuseum van Natuurlijke Historié, Leiden, for pointing out to us that if the locality were confirmed this spécimen would represent the first AustraUan record of the genus Chitina. However, the locality is in doubt since a further spécimen, 1877.5.21.124, is labelled in manuscript 'New Zealand/Bowerbank Coll.'. The printed label with the first spécimen was probably prepared for its public exhibition, and transcription error cannot be eliminated. No original label survives with this spécimen. A dried spécimen labelled 'Chitina ericopsis, Oko-Siri, Japan, 41° 59'N, 138° 30'E, 25-30 fm, coll. HMS Actaeon, près. J. F. R. Ayler (Master)', BMNH regd no. 1862.7.16.9, was re-identified as Solanderia cf misakinensis (Inaba) by our colleague S. J. Moore in 1986. The former positions of the hydranths are évident on both hydrocladia and hydrocauli, and we concur with his generic re-assignment. Spirit-preserved material. Several colonies collected by Dr Patricia M. Ralph, Wellington Harbour, New Zealand, 18 m (' 10 fathoms'), BMNH regd nos 1957.11.3.1, 1958.1.2.1,1961.4.8.1; described below. Description The description is based jointly on the type material and the more recently collected spirit-preserved spécimens. General morphology. (Figs 1-2). Colony erect, recorded up to c. 38 cm, branched in ail planes, compactly bushy even in tall spécimens; comprising one to several principal, much branched, hydrocauline trunks up to 2-5 cm in diameter from which hydrocladia anse at irregular intervais and in ail planes; trunks and stems yellowish-fawn. Trunks themselves bifurcating several times in succession, resulting in gradually finer and more délicate branches and branchlets. When several main trunks, thèse anastomosing through short latéral struts especially in basai région. In contrast hydrocladia anastomosing only exceptionally. Main trunks issuing from well developed meshwork of interlaced and frequently anastomosing hydrorhizal fibres which anchor colony (R in Figs 1-2). Thickest main trunks having latéral tubular expansions probably representing positions of future branches. Skeleton. (Figs 2-5, 7). Skeleton internai, permeated by and covered with communal coenosarc of colony. Skeleton of main trunks and principal branches a dense network of interlaced and frequently anastomosed