Kotatea aurantiaca Quoy & Gaimard, 1833

Kessel, Gustav M., Alderslade, Philip, Bilewitch, Jaret P., Schnabel, Kareen E., Norman, Jerry, Potts, Romana Tekaharoa & Gardner, Jonathan P. A., 2022, Dead man’s fingers point to new taxa: two new genera of New Zealand soft corals (Anthozoa, Octocorallia) and a revision of Alcyonium aurantiacum Quoy & Gaimard, 1833, European Journal of Taxonomy 837, pp. 1-85 : 17-24

Kotatea aurantiaca Quoy & Gaimard, 1833 gen. et comb. nov.

Figs 1B–C , 2B, L , 3A–C , 7B , 8–10 , 11A–C, E

Alcyonum aurantiacum Quoy & Gaimard, 1833 [spelling of Alcyonium – lapsus calami]: 277, pl. 22 figs 16–18.

Alcyonium aurantiacum – Benham 1928 in part: 71–75, figs 6–11.

? Alcyonium aurantiacum – Powell 1947: 8, fig. 14. — Doak 1971: 44–46, pl. 20. — Westerskov & Probert 1981 in part: 111, pl. 28. — Grange et al. 2010 in part: 148.

non Alcyonium aurantiacum – McFadden et al. 2006b (= Ushanaia fervens gen. et sp. nov.): 517, 521, 523, figs 1, 3.

Diagnosis

Colonies with branching lobes, colour varying from pale to dark orange, with white polyps. Collaret and points colourless and composed of warty spindles and some irregular and branched forms, clubs rare and small. Tentacles contain irregular, warty, scale-like sclerites. Polyp neck contains spiny spindles and warty rod-like sclerites. Polyp mounds contain slender, spiny clubs and warty rod- and spindle-like sclerites. Lobe and base surface contains radiates, clubs, and occasionally leafy spheroids. Lobe and base interior contains slender spindles and radiates.

Etymology

The species name is the feminine form of aurantiacum , the original species epithet ( Quoy & Gaimard 1833).

Material examined

NEW ZEALAND – Northland • 1 specimen; ~ 14 km N of Piwhane / Spirits Bay ; 34.3163° S, 172.7925° E; depth 68 m; 28 Jan. 1999; NIWA exped.; stn Z9702 ( KAH9901 /73); NIWA 3972 • 1 specimen; ~ 14 km NW of Piwhane / Spirits Bay ; 34.3510° S, 172.7088° E; depth 57 m; 28 Jan. 1999; NIWA exped.; stn Z9697 ( KAH 9901 /64); NIWA 3978 • 2 specimens; ~ 12 km NW of Piwhane / Spirits Bay ; 34.3533° S, 172.7487° E; depth 55 m; 22 Apr. 1999; NIWA exped.; stn Z9753; NIWA 12656 • 1 specimen (cut into 2 fragments); ~ 3 km N of North Cape ; 34.3668° S, 173.0003° E; depth 89 m; 27 Jan. 1999; NIWA exped.; stn Z9695 ( KAH 9901 /59); NIWA 3979 • 1 specimen; ~ 8 km N of Piwhane / Spirits Bay ; 34.3690° S, 172.8250° E; depth 55 m; 25 Jan. 1999; NIWA exped.; stn Z9677 ( KAH 9901 /25); NIWA 3980 • 2 specimens; ~ 5.5 km NNE of Cape Reinga ; 34.3745° S, 172.7013 ° E; depth 53 m; 27 Jan. 1999; NIWA exped.; stn Z9688 ( KAH 9901 /47); NIWA 3977 • 1 specimen; ~ 7 km E of Takou Bay ; 35.0750° S, 174.0183° E; depth 67–72 m; 19 Jul. 2009; Oceans Survey 2020 exped.; stn TAN0906/245; NIWA 57596 • 1 specimen; Bay of Islands , ~ 1.5 km NE of Harakeke Island; 35.1457° S, 174.1517° E; depth 55 m; 7 Jul. 2009; Oceans Survey 2020 exped.; stn TAN0906/57; NIWA 55142 • 2 specimens; same collection data as for preceding; NIWA 55164 • 1 specimen; Bay of Islands , ~ 500 m NW of Motutara Island; 35.2075° S, 174.1940° E; depth 23–27 m; 3Sep. 2009; Oceans Survey 2020 exped.; stn KAH 0907 /195; NIWA 58551 • 1 specimen; ~ 4 km NE of Whananaki ; 35.4858° S, 174.5012° E; depth 59–63 m; 5 Jul. 2009; Oceans Survey 2020 exped.; stn TAN0906/21; NIWA 54641 • 1 specimen; same collection data as for preceding; NIWA 54700 • 2 specimens; ~ 6 km ENE of Whananaki ; 35.5002° S, 174.5415° E; depth 64–66 m; 4 Jul. 2009; Oceans Survey 2020 exped.; stn TAN0906/3; NIWA 54535 • 4 specimens; Matapouri , ~ 4 km NE of Matapouri Bay; 35.5525° S, 174.5525° E; depth 57 m; 5 Jul. 2009; Oceans Survey 2020 exped.; stn TAN0906/25; NIWA 54766 . – Bay of Plenty • 1 specimen; ~ 5 km NW of Whakaari / White Island ; 37.4785° S, 177.1280° E; depth 12–17 m; 30 Apr. 1999; Coral Reef Research Foundation exped.; stn Z15884 ; MAGNT C013957 • 67 specimens; same collection data as for preceding; NIWA 101181 • 1 specimen; Whakaari / White Island, Volkner Rocks ; 37.5167° S, 177.1833° E; depth 12–20 m; 30 Apr. 1999; J. Starmer leg.; MAGNT C015225 • 1 specimen; same collection data as for preceding; MAGNT C015231 • 1 specimen; same collection data as for preceding; MAGNT C015232 • 2 specimens; same collection data as for preceding; MAGNT C015233 • 1 specimen; same collection data as for preceding; MAGNT C015234 • 1 specimen; same collection data as for preceding; MAGNT C015235 . – East Cape • 5 specimens; Ranfurly Bank ; 37.5472° S, 178.8925° E; depth 68–70 m; 30 May 2011; Oceans Survey 2020 exped.; stn TAN1108/213; NIWA 75330 • 4 specimens; Ranfurly Bank; 37.5823° S, 178.8975° E; depth 42–48 m; 31 May 2011; Oceans Survey 2020 exped.; stn TAN1108/217; NIWA 75393 . – Unknown location • 6 specimens; older than 1995; stn B5/96, no other data available; NIWA 142997 .

Type locality

Firth of Thames, North Island, NZ, depth ~ 14– 18 m.

Preliminary remarks

Quoy & Gaimard’s (1833) description alone lacks the detail needed to distinguish which of several species of Kotatea gen. nov. could be A. aurantiacum . However, when their original colour plate ( Fig. 11A ) and the photograph of the syntype specimens of A. aurantiacum ( Fig. 11E ) are considered in conjunction with the morphology and distributional range of all available specimens, the material here ascribed to K. aurantiaca gen. et comb. nov. is almost certainly conspecific with Quoy & Gaimard’s (1833) species.

Apart from sclerites, Kotatea aurantiaca gen. et comb. nov. specifically differs from all of its congeners as follows: Kotatea amicispongia gen. et sp. nov. has only been collected from much greater depths than Quoy & Gaimard’s (1833) material; K. kapotaiora gen. et sp. nov. and K. teorowai gen. et sp. nov. do not match the colour described for the original material as they are white rather than orange; K. raekura gen. et sp. nov. is known only from Manawatāwhi/Three Kings Islands and not near the type locality; and K. kurakootingotingo gen. et sp. nov., K. lobata gen. et sp. nov., and K. niwa gen. et sp. nov. all tend to differ in colony growth form. Ushanaia gen. nov. also differs in growth form.

Description

Colony form

Kotatea aurantiaca gen. et comb. nov. produces irregularly branched, lobate colonies. Lobes are usually finger-like but can appear more robust when contracted ( Figs 3A–C , 7B , 8 ). Preserved specimens vary in colour from very pale to dark orange and measure up to 7 cm in height and 7 cm in width ( Figs 7B , 8 ), but the species may attain larger sizes (see remarks below). Finger-like lobes emerge, often profusely, from a broad base that is usually lighter in colour than the rest of the colony, and which may be very short ( Fig. 8 ), or from a stalk ( Fig. 7B ). Polyps are most densely concentrated at lobe tips and tend to become sparser towards the base of the colonies, from which they are usually absent. Polyps are white in preserved specimens, are 0.5–1 mm tall when expanded and have colourless collaret and points ( Fig. 2B, L ).

Sclerites

Points are composed of warty spindles (~ 0.15–0.4 mm long) and a few small clubs distally (~ 0.1 mm long) ( Fig. 9A–B ). Proximally, the spindles become larger and more crescentic (~ 0.3–0.5 mm long), transitioning into a transverse orientation and merging with the collaret, which is five to seven rows deep ( Figs 9A , 10C ). Among the spindles, both the collaret and the points also contain some similarly sized, irregular, sometimes branched sclerites. The tentacles contain irregular, warty, scale-like forms that are often slightly crescentic (~ 0.06–0.2 mm long) ( Fig. 9C ), the polyp neck contains spiny spindles and warty rod-like forms (~ 0.06–0.15 mm long) ( Fig. 9D ), and the polyp mounds contain slender, spiny clubs and a few warty rod- and spindle-like forms (~ 0.06–0.15 mm long, clubs ~ 0.03 mm wide) ( Fig. 9E ). The surface of both the lobes and the base contains radiates and clubs (~ 0.05–0.12 mm long), with clubs being more common in the lobe surface ( Fig. 9F, H ). Surface sections may also occasionally include leafy spheroids ( Fig. 9F ). The interior of both the lobes and the base contains long, slender spindles with branches and/or complex tubercles, as well as radiates, with radiates being more common and spindles tending to be more branched in the interior of the base ( Figs 9G , 10A ). Interior sclerites are ~ 0.06–0.26 mm long.

Variability

Colonies of this species can expand and contract to a considerable degree. Consequently, the presence of a stalk may be difficult to discern, and although K. aurantiaca gen. et comb. nov. can resemble K. lobata gen. et sp. nov. when highly contracted (compare NIWA 101181 in Fig. 8 to Fig. 18 ; also see illustrations in Doak 1971), the material to hand indicates that the lobes of the latter are usually considerably longer and more robust.

Point clubs are overall more common in some specimens than in others and can be absent from some polyps. Additionally, leafy spheroids are present in low numbers in the surface sections of most colonies but may be absent. Beyond this, there is very little variability in the sclerites across all specimens, with size ranges falling within those described for the holotype in all cases ( Figs 9–10 ).

Comparisons

Kotatea aurantiaca gen. et comb. nov. is most similar to congeners which commonly exhibit branching of the lobes: K. amicispongia gen. et sp. nov., K. kapotaiora gen. et sp. nov., K. raekura gen. et sp. nov., and K. teorowai gen. et sp. nov. Differences from K. amicispongia are discussed under that species.

Other than colour (compare Figs 7B , 8 and 14C ), Kotatea aurantiaca gen. et comb. nov. specimens differ markedly from K. kapotaiora gen. et sp. nov. in lacking the latter’s large and robust clubs in the lobe surface (compare Figs 9F and 12F ), and in possessing interior sclerites composed largely of slender spindles, while those of K. kapotaiora specimens are distinct, irregular radiates with minimal branching processes (compare Figs 9G , 10A and 12G , 13B ).

Kotatea aurantiaca gen. et comb. nov. specimens can be easily differentiated from K. raekura gen. et sp. nov. by their colourless collaret and point sclerites, which are always coloured orange in K. raekura Sclerites of the collaret and point, polyp neck, polyp mound and surface regions also clearly differ between the two species, with those in K. aurantiaca gen. et comb. nov. specimens being much smaller and more slender than the overall more robust sclerites found in K. raekura (compare Figs 9A, D–F, H and 25A, D–F , 27B ). Additionally, K. raekura specimens have shallower collarets (3–5 vs 5–7 rows) and may be restricted to Manawatāwhi/Three Kings Islands judging from the available material.

As for Kotatea kapotaiora gen. et sp. nov., K. aurantiaca gen. et comb. nov. differs in colour from K. teorowai gen. et sp. nov. (compare Figs 7B , 8 and 24B ). Notably, K. teorowai completely lacks the slender interior spindles that are present and abundant in all K. aurantiaca gen. et comb. nov. specimens, possessing only rare, irregularly branched radiates in its interior (compare Figs 9G , 10A and 27H ). Additionally, while leafy spheroids are not common in any K. aurantiaca gen. et comb. nov. specimen, these sclerites are well-developed and feature conspicuously in the lobe surface of K. teorowai (compare Figs 9F and 27F ).

Kotatea aurantiaca gen. et comb. nov. and K. lobata gen. et sp. nov. are probably the most commonly encountered species of the genus. Specimens of K. lobata are distinctive in possessing very large, highly branched, antler-like sclerites in their interiors, especially in the lobes. By contrast, K. aurantiaca gen. et comb. nov. specimens entirely lack these sclerites, and their interiors are instead composed predominantly of slender spindles (compare Figs 9G , 10A and 20A , 21A ). Equally characteristic are the very large spindle-like sclerites found in the surface sections (particularly of the base) of K. lobata specimens, which are again absent in K. aurantiaca gen. et comb. nov. (compare Figs 9F, H and 19F , 20B ). Additionally, point clubs are more abundant in K. lobata specimens (compare Figs 9B and 19B ).

While their growth forms are, in general, sufficiently distinct to allow for differentiation, in many cases colony-scale morphological overlap between Kotatea aurantiaca gen. et comb. nov. and K. lobata gen. et sp. nov. may prevent species identification by eye, especially in small or very contracted colonies and in areas where both species may be present (e.g., compare K. lobata specimen NIWA 108960 in Fig. 18 with some of the K. aurantiaca gen. et comb. nov. colonies of NIWA 101181 and NIWA 54535 in Fig. 8 ).

Habitat and distribution

Specimens were collected from northern New Zealand, between Piwhane/Spirits Bay and East Cape at depths of ~ 10–90 m ( Fig. 1B–C ). Many of the specimens were collected from rocky, gravelly and shelly substrates alongside seaweed, hydrozoans, ascidians, bryozoans and large numbers of various species of sponge.

Remarks

Quoy & Gaimard (1833) most likely did not observe the tentacles of their specimens in an expanded state, as they describe the tentacles as short and rounded and their plate ( Fig. 11B ) also shows these to be contracted.

Since Quoy & Gaimard’s work, the only other taxonomic treatment of Alcyonium aurantiacum is that by Benham (1928), who described the morphology of three specimens and pointed out that Quoy & Gaimard’s description omits the ‘i’ in Alcyonium .Again, Benham’s descriptions have limited usefulness in distinguishing among the closely related species described here. However, judging from one of Benham’s sketches, it is possible that one of his specimens collected from the Mahia Peninsula (reproduced here in Fig. 11C ) may have been K. aurantiaca gen. et comb. nov., while an encrusting specimen collected in Dusky Sound and growing around a black coral fragment ( Fig. 11D ) almost certainly represents Ushanaia fervens gen. et sp. nov.

The identity of Benham’s third specimen, collected at Tasman Bay/Te Tai-o-Aorere, is unclear. This is described as stalked and lobed in growth form, with noticeably orange collaret sclerites, and so is likely a member of Kotatea gen. nov., but most likely not K. aurantiaca gen. et comb. nov., which possesses colourless collaret sclerites. Benham (1928) also believed a specimen from the Auckland Islands was A. aurantiacum , and while the exact identity of this specimen cannot be ascertained from his descriptions, it points to the possible presence of Kotatea or Ushanaia gen. nov. in New Zealand’s subantarctic islands, but at present no samples are known from this far south.

Similarly, Grange et al. (2010) illustrate what appears to be a Kotatea gen. nov. colony, possibly K. aurantiaca gen. et comb. nov., from Fiordland, and Powell (1947) mentions having commonly dredged what is likely K. aurantiaca gen. et comb. nov. from depths of ~ 10–15 m between Motuihe Island/Te Motu-a-Ihenga and Waiheke Island in the Hauraki Gulf/Tīkapa Moana. Additionally, observations recorded on iNaturalist (https://www.inaturalist.org, accessed Jan. 2021) of what appears to be K. aurantiaca gen. et comb. nov. indicate that it may reach at least as far south as Kaikōura. Therefore, this species and the genus in general may be (or have been) considerably more widely distributed around coastal New Zealand than available specimens would suggest.

Alcyonium aurantiacum occurring in southern Australia, as noted by Grange et al. (2010), is probably in reference to A. etheridgei Thomson & Mackinnon, 1911 . This species is Alcyonium -like and superficially similar in appearance as it is red when alive, but not related to Kotatea gen. nov. ( Verseveldt &Alderslade 1982; Alderslade pers. comm.), which should for now be considered endemic to New Zealand.

Alcyonium aurantiacum has previously been reported to grow intertidally ( Morton & Miller 1973; Morton 2004; Grange et al. 2010), particularly among ascidians and sponges on moderately exposed shores ( Westerskov & Probert 1981). Here, it has also been reported that the native nudibranch, Tritonia incerta Bergh, 1904 , grazes on A. aurantiacum ( Morton & Miller 1973; Westerskov & Probert 1981). However, intertidal observations are probably of K. lobata gen. et sp. nov., and not K. aurantiaca gen. et comb. nov.

Kotatea aurantiaca gen. et comb. nov. likely reaches a height of at least 30 cm when fully expanded in vivo, as noted by Grange et al. (2010) for A. aurantiacum . Present preserved material does not exceed ~ 7 cm in height.

Figures

Fig. 1. Collection sites of specimens of Kotatea gen. nov. and Ushanaia gen. nov. Holotype collection sites are indicated by white symbols, paratypes and other material by black symbols. In the case of K. aurantiaca gen. et comb. nov., the type collection site refers to that described by Quoy & Gaimard (1833). A. Manawatāwhi/Three Kings Islands. B. Far northern NZ, from Piwhane/Spirits Bay to the Poor Knights Islands. C. Northern NZ, from the Mokohinau Islands to East Cape. D. Central NZ, Cook Strait to Banks Peninsula. E. South-East NZ, Fiordland. Note that the holotypes of K. kapotaiora gen. et sp. nov. and K. teorowai gen. et sp. nov. share the same collection site.

Fig. 2. Polyps. A. Kotatea amicispongia gen. et sp. nov., holotype (NIWA 156312). B. K. aurantiaca gen. et comb. nov. (NIWA 75330). C. K. kapotaiora gen. et sp. nov., holotype (NIWA 3974). D. K. kurakootingotingo gen. et sp. nov., holotype (NIWA 101538). E. K. kurakootingotingo gen. et sp. nov., paratype, small polyp with coloured collaret and points (MAGNT C015221). F. K. lobata gen. et sp. nov., holotype (NIWA 101313). G. K. niwa gen. et sp. nov., holotype (MAGNT C015226). H. K. raekura gen. et sp. nov., holotype (NIWA 101537). I. Ushanaia ferruginea gen. et sp. nov., holotype (NIWA 156313). J. U. fervens gen. et sp. nov., holotype (NIWA 156311). K. U. solida gen. et sp. nov., holotype (NIWA 102133). L. K. aurantiaca gen. et comb. nov., in situ (uncollected specimen), Poor Knights Islands, photo by Ian Skipworth (ianskipworth.com). M. U. fervens gen. et sp. nov., in situ (uncollected specimen), Fiordland, photo by Shane Geange. Scale bars = ~1 mm.

Fig. 3. In situ photographs of species of Kotatea gen. nov. usually possessing finger-like lobes. A–B. K. aurantiaca gen. et comb. nov. (uncollected specimens), expanded colonies with a contracted colony inupper right background of B, Poor KnightsIslands, photos by Ian Skipworth (ianskipworth.com). C. K. aurantiaca gen. et comb. nov. (MAGNT C013957/NIWA 101181), large colony, Bay of Plenty, photo by Coral Reef Research Foundation. D. K. raekura gen. et sp. nov., holotype (NIWA 101537), Manawatāwhi/Three Kings Islands, photo by NIWA. Scale bars = ~3 cm.

Fig. 7. Preserved specimens. A. Kotatea amicispongia gen. et sp. nov. B. K. aurantiaca gen. et comb. nov., expanded and stalked specimens. Note NIWA 154046 and NIWA 142997 contain additional fragments that are not depicted. * = holotype.

Fig. 8. Selected preserved specimens of Kotatea aurantiaca gen. et comb. nov. Note that most specimen lots include small additional fragments that are not depicted and NIWA 101181 comprises a total of 67 similar colonies, all of which were examined.

Fig. 9. Kotatea aurantiaca gen. et comb. nov. (NIWA 54766), SEMs of sclerites. A. Collaret and points. B. Distal points. C. Tentacles. D. Polyp neck. E. Polyp mound. F. Lobe surface. G. Lobe interior. H. Base surface.

Fig. 10. Kotatea aurantiaca gen. et comb. nov. (NIWA 54766), SEMs of sclerites. A. Base interior. B. Polyps (in situ).

Fig. 11. A–B. Copies of Quoy & Gaimard’s (1833) illustrations of “A. aurantiacum”. C–D. Copies of the sketches of upright and encrusting specimens by Benham (1928), identified as A. aurantiacum. E. A. aurantiacum syntype, specimen MNHN-IK-2000-128 (photo by Marie Hennion).

Fig. 18. Selected preserved specimens of Kotatea lobata gen. et sp. nov. Note that NIWA 142995, NIWA 101740 and NIWA 108960 contain additional fragments that are not depicted. * = holotype.

Fig. 14. Preserved specimens. A. Kotatea kurakootingotingo gen. et sp. nov. B. K. niwa gen. et sp. nov. C. K. kapotaiora gen. et sp. nov. Note that MAGNT C015224 and NIWA 58543 contain additional fragments that are not depicted. * = holotype.

Fig. 12. Kotatea kapotaiora gen. et sp. nov., holotype (NIWA 3974), SEMs of sclerites. A. Collaret and points. B. Distal points. C. Tentacles. D. Polyp neck. E. Polyp mound. F. Lobe surface. G. Lobe interior.

Fig. 13. Kotatea kapotaiora gen. et sp. nov., holotype (NIWA 3974), SEMs of sclerites. A. Base surface. B. Base interior. C. Polyps (in situ).

Fig. 25. Kotatea raekura gen. et sp. nov., holotype (NIWA 101537), SEMs of sclerites. A. Collaret and points. B. Distal points. C. Tentacles. D. Polyp neck. E. Polyp mound. F. Lobe surface.

Fig. 27. Kotatea teorowai gen. et sp. nov., holotype (NIWA 27358), SEMs of sclerites. A. Collaret and points. B. Distal points. C. Tentacles. D. Polyp neck. E. Polyp mound. F. Lobe surface. G. Base surface. H. Lobe and base interior.

Fig. 24. Preserved specimens. A. Kotatea raekura gen. et sp. nov. B. K. teorowai gen. et sp. nov. Note that lot AK 656516 includes additional fragments that are not shown. * = holotype.

Fig. 20. Kotatea lobata gen. et sp. nov., holotype (NIWA 101313), SEMs of sclerites. A. Lobe interior. B. Base surface.

Fig. 21. Kotatea lobata gen. et sp. nov., holotype (NIWA 101313), SEMs of sclerites. A. Base interior. B. Polyps (in situ).

Fig. 19. Kotatea lobata gen. et sp. nov., holotype (NIWA 101313), SEMs of sclerites. A. Collaret and points. B. Distal points. C. Tentacles. D. Polyp neck. E. Polyp mound. F. Lobe surface.