Sciurella indivisa Allman, 1883

Galea, Horia R., Maggioni, Davide & Di Camillo, Cristina G., 2021, Towards a better understanding of the genus Sciurella Allman, 1883 (Cnidaria: Hydrozoa: Plumulariidae): evidence from an integrative study, Zootaxa 5040 (1), pp. 1-32 : 5-14

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Sciurella indivisa Allman, 1883


Sciurella indivisa Allman, 1883

Figs 1–7 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE 5 View FIGURE 6 View FIGURE 7 , 19 View FIGURE 19 , 20 View FIGURE 20 ; Table 2

Sciurella indivisa Allman, 1883: 9 , 15, 23, 26, pl. 5 figs 1–4.— Allman, 1888: lxv, lxix.— Kirkpatrick, 1890: 609.— von Lendenfeld, 1885a: 479.— von Lendenfeld, 1885b: 626.— von Lendenfeld, 1885c: 644.— von Lendenfeld, 1887: 28.— Nutting,

1900: 15, 31.— Billard, 1908: 759.— Bedot, 1916: 196.— Bedot, 1917: 17, 18.— Bedot, 1918: 231.— Van Praët, 1979:

933.— Di Camillo et al., 2008: 1592. Nemertesia indivisa — Billard, 1910: 38.— Billard, 1913: 60, fig. 50.— Bedot, 1917: 18.— Bouillon et al., 2006: 369.— Ramil &

Vervoort, 2006: 123. Nemertesia sp. 1 — Schuchert, 2015: 348, fig. 20. non Sciurella indivisa —Kirckpatrick, 1890: 604, 609 [= Sciurella cylindrica ( Kirchenpauer, 1876) ]. non Nemertesia indivisa — Schuchert, 2003: 215, fig. 63 [= Sciurella cylindrica ( Kirchenpauer, 1876) ].

Material examined. MNHN H.L. 1373, Australia, QLD, Cape York, Somerset Island: a slide mounted by A. Billard containing a 2.4 cm long, unstained stem fragment bearing distally a gonotheca [this is part of the type material housed in the Natural History Museum, London, UK; the slide is labelled in Billard’s handwriting « Sciurella indivisa Allm. / type du Challenger / Nemertesia indivisa (Allm.) / 9.X.1908].— MNHN H.L. 1374: a slide (with the same label as the preceding one) mounted by A. Billard, containing three unstained cladia, of which one was detached from the stem together with its apophysis and a pair of gonothecae.—MHNG-INVE-0137405, Indonesia, North Sulawesi, North Minahasa Regency, Bunaken Marine Park, Manado Tua I., 1.623017°, 124.714263°, 25 m, 15 Feb. 2005: several stems and fragments up to 8.5 cm, detached from substrate above insertion on hydrorhiza; a 5 cm high stem bears distally many female gonothecae, the others are sterile.—MHNG-INVE-0137406, Indonesia, North Sulawesi, North Minahasa Regency, Bunaken Marine Park, Siladen I., 1.626414°, 124.802058°, 20 m, 20 Feb. 2005: several stems and fragments up to 6.4 cm high, all sterile.—MHNG-INVE-0091092, Japan, Okinawa I., 26.713720°, 127.878600°, 29 m, 19 Jun. 2008: DNA extracted from a portion of colony identified by Schuchert (2015) as Nemertesia sp. 1 , GenBank: MZ 099712 View Materials (16S), MZ 348925 View Materials (18S), MZ 099659 View Materials (28S).

Description. Colonies forming clumps of several stems growing on dead corals, with no evident hydrorhizae ( Fig. 1A View FIGURE 1 ); stems up to 8.5 cm high, simple or forked, monosiphonic, with canaliculate coenosarc ( Fig. 5H View FIGURE 5 ), divided into a regular succession of moderately long internodes by means of deep, transverse constrictions of the perisarc ( Figs 2B–D View FIGURE 2 ; 5A–C View FIGURE 5 ); each internode with several apophyses supporting the cladia, and a varied number of nematothecae on surface ( Figs 3A View FIGURE 3 ; 5B, C View FIGURE 5 ); there are generally 4 apophyses, occasionally as few as 2 or 3, or as many as 5, 6 or 8, their number being directly related to the length of the internode; apophyses in younger, slenderer stems given off irregularly ( Fig. 2B View FIGURE 2 ), but showing a distinct tendency to adopt a bipinnate arrangement in older, thicker stems (two opposite, or nearly so, cladia near to the proximal node, and two opposite, or nearly so, cladia near the distal node; Fig. 2D View FIGURE 2 ); apophyses relatively short, pointing upward, bearing a conspicuous, adcauline mamelon with broad, circular aperture, two pairs of axillar nematothecae inserted on each side at the origin from internode, as well as an unpaired, central nematotheca above ( Fig. 3B, C, E View FIGURE 3 ) (the latter only present when the apophysis is not situated just below the distal stem internode, Fig. 3D View FIGURE 3 ); stem internodes with a belt of several nematothecae near the proximal node, as well as a number of nematothecae irregularly scattered on the surface of the internode, each nematotheca in relation to a given coenosarcal tube ( Figs 3A View FIGURE 3 ; 5B, C View FIGURE 5 ). Cladia given off at ca. 45° with the stem, up to 11 mm long, delimited or not from the corresponding stem apophysis by a proximal, oblique node; the presence of a first, short, quadrangular, ahydrothecate internode provided with an adaxial nematotheca (exceptionally two) is nearly constant in one stem ( Fig. 3A, B View FIGURE 3 ), but decidedly absent ( Figs 3C View FIGURE 3 ; 5D View FIGURE 5 ) in others, including the fertile stem; remainder of cladium homomerously-divided, composed of a regular succession of up to 18 hydrothecate internodes separated by slightly oblique, distinct nodes; each internode moderately long, with a hydrotheca in middle, spanning about half its length, and its complement of three nematothecae; two internal perisarcal ridges, one proximally and one distally ( Fig. 4A, B View FIGURE 4 ) (the former absent in proximal most cormidium, Fig. 3A, C View FIGURE 3 ). Hydrotheca long, tubular, fully adnate ( Figs 4B View FIGURE 4 ; 5E View FIGURE 5 ), abaxial wall distinctly thick, straight for most of its length, imperceptibly concave proximally and slightly flared distally; adaxial wall straight, with perisarc thickened laterally at junction with the internode behind ( Figs 3C View FIGURE 3 ; 4B View FIGURE 4 ); base concave, hydropore set next to the origin of abaxial wall; aperture transverse, circular in apical view, rim even, except for two lateral sinuations at fusion site with the internode behind; three nematothecae, of which one mesial, on a conspicuous projection, far below the hydrothecal base ( Figs 4K View FIGURE 4 ; 5E View FIGURE 5 ), and a pair of laterals ( Figs 4A, I, J View FIGURE 4 ; 5E–G View FIGURE 5 ); hydranths contracted, column elongated (filling most part of the hydrothecal lumen), distally provided with a reduced number of filiform tentacles. All nematothecae, including those of the stem, relatively short, movable and bithalamic; rim of upper chamber deeply scooped adaxially ( Fig. 4J, K View FIGURE 4 ). Gonothecae occurring in pairs, given off from each side of the stem apophyses, just below the mamelon ( Figs 6A View FIGURE 6 ; 7B–F View FIGURE 7 ); female in present material; body broadly ovoid in frontal view, provided with a number of prominent, conical spines, generally arranged in three lateral pairs, one unpaired apically, and 1–2 abaxial ones ( Figs 6B View FIGURE 6 ; 7E View FIGURE 7 ); each spine with 1–2 nematothecae, of which one is apical, and the second commonly basal, although it could be borne in the middle of the spine ( Fig. 7G View FIGURE 7 ); each spine is served by an independent coenosarcal tube derived from the central spadix, each branch ending up in a nematophore ( Fig. 6A View FIGURE 6 ); occasionally, some spines could be bifid, and some nematothecae unrelated to a spine, but simply borne on the surface of the gonotheca; in lateral view, the gonotheca appears distinctly flattened dorsoventrally, and is concave on its adaxial side and convex on the abaxial side; it is widest in middle, and thins out toward its margins; a circular aperture is borne on a small projection of the adaxial side ( Fig. 6C View FIGURE 6 , arrowhead); two large, spherical oocytes are contained within each gonotheca, and subsequently released and confined to a «brooding chamber» formed by the adaxial wall of the gonotheca with the corresponding stem internode ( Fig. 7F View FIGURE 7 ). Color in life: cauli pale-brown, cladia white, female gonothecae translucent, oocytes white, spadix brown.

Remarks. In one stem from Siladen, nearly all cladia begin with a short, quadrangular, nematothecate internode ( Fig. 3A, B View FIGURE 3 ), while in some others it is either occasionally present or totally absent ( Fig. 3C View FIGURE 3 ). As no signs of damage followed by subsequent regeneration could be noted, it is assumed that its occurrence is variable in this species. Its presence is also illustrated by Allman (1883: pl. 5, figs 2–3) in his original account, but does not seem to occur in all stems, as demonstrated here through the reinspection of the type (e.g. MNHN H.L. 1373, Fig. 3D View FIGURE 3 ), nor in the material assigned by Schuchert (2015: 348, fig. 20B) to Nemertesia sp. 1 (see below a discussion on this material).

In some stems, the cauline apophyses give rise occasionally to two cladia, one in the normal position, while the other is given off laterally from below one side of the mamelon, through a short athecate apophysis, followed by a quadrangular, nematothecate segment.

Stems of this species reportedly reach as much as 25 cm in height ( Allman 1883: 26, «ten inches»). According to the original account, the cladia are arranged «in four longitudinal alternating series» ( Allman 1883: 26), but verticils of three were found in the part of the type material (MNHN H.L. 1373) examined here ( Fig. 2A View FIGURE 2 ). In the quite rich material of the Siboga , the cladia are reportedly said either scattered or verticillate 16. In some of the youngest stems from Bunaken examined here, the cladia are either alternately-arranged ( Fig. 2C View FIGURE 2 ) or decidedly scattered to spirally-arranged ( Fig. 2B View FIGURE 2 ), while in the oldest ones, they adopt a rather bipinnate arrangement ( Fig. 2D View FIGURE 2 ).

The adequate understanding of the morphology of the fully-formed gonotheca was a gradual process. Allman (1883: 26) described it as «urn-shaped in front view, with two symmetrically placed hollow lateral processes near the distal end», and «compressed laterally, and when viewed in profile is seen to have its axis curved backwards nearly in a semicircle». Later on, Billard (1908), upon the reexamination of the type, described the gonothecae as concave,

16 “They are either scattered or verticillate” ( Billard 1913: 60) [translated from French].

with an irregular outline, their distal part being provided with three lobes 17. Even later, Billard (1910), evidently having been in possession of the hydroid collection of the Siboga , stressed again the lobate condition of the gonothecae, while specifying that they were both inaccurately drawn by Allman and imprecisely described earlier by himself 18. Finally, he reiterated ( Billard 1913) the statements from his 1910 work in nearly the same words, and provided an illustration of two gonothecae, one seen frontally and the other almost in profile view ( Billard 1913: fig. L). Unlike the gonothecae met with in our specimens from Bunaken ( Figs 6B View FIGURE 6 ; 7A–G View FIGURE 7 ), they were provided with much broader, bifid or even trifid marginal lobes, only the proximal most still adopting an elongated, conical shape. The two gonothecae present on slide MNHN H.L. 1374 are certainly not those depicted by Billard, as they only bear simple, conical projections; one of them is illustrated in Fig. 6D View FIGURE 6 herein (the other is even more crushed). Consequently, it is difficult to tell whether Billard’s illustrations are either diagrammatic representations or real gonothecae with much more structurally complex shapes than those examined in the course of the present study.

In light of the data provided by Billard (1913) 19 upon the examination of numerous specimens, it seems that the structure of the stem apophyses supporting the cladia is the same as that met with in our material, i.e. two pairs of nematothecae flank an apophysis provided adaxially with a conical mamelon ( Fig. 3E View FIGURE 3 ), while an uneven nematotheca is found a short distance above on the stem internode ( Fig. 3B, C View FIGURE 3 ). The same also results from the examination of specimen MNHN H.L. 1373 belonging to the type series ( Fig. 3D View FIGURE 3 – note in this example the absence of the unpaired, superior nematotheca due to the proximity of a distal node). A similar situation occurs in the Japanese material assigned to Nemertesia sp. 1 by Schuchert (2015: fig. 20B). In addition, the bipinnate arrangement of their cladia along the stems, and the relatively long cormidia provided both proximally and distally with an internal, annular ridge towards the nodes (compare Fig. 4F and 4B View FIGURE 4 ), suggests that his material is, with little doubt, S. indivisa . Cladial internodes in S. indivisa are highly variable in length: those of the type series (specimens MNHN H.L. 1373 17 “[…] in the preparations of the type species that I have examined, the distal part of the gonothecae is lobed (three lobes), instead of being entire. These gonothecae are thus irregular in shape and, additionally, are concave-convex, instead of being planar” ( Billard 1908: 759) [translated from French].

18 “ The gonothecae are not accurately illustrated by Allman either; their distal part is irregularly lobed, instead of being entire, like in Allman’s drawing, or provided with three lobes, as I stated earlier […]. They are, additionally, concave-convex, instead of being planar; their concavity faces the stem, so as to form a sort of brooding chamber, as I was able to document it in the specimens gathered by the « Siboga » Expedition” ( Billard 1910: 39) [translated from French].

19 “The apophysis supporting the hydrocladium is provided with two pairs of axillar nematothecae and a basal mamelon bearing an aperture; there is a stem nematotheca above the insertion of hydrocladia” ( Billard 1913: 60) [translated from French].

and 1374) are quite short ( Figs 4E and 4D View FIGURE 4 , respectively), while those in material from Bunaken are comparatively longer ( Fig. 4B, C View FIGURE 4 ), especially above the hydrotheca. Such size variations were also noted by Billard (1913) in the abundant material of the Siboga . The hydrothecae, too, similarly display varied depths (compare Figs 4B, C, F and 4D, E View FIGURE 4 ; see also Table 2).

Distribution. Australia: Cape York, QLD ( Allman 1883). Indonesia: Sape Strait, between Sumba and Komodo islands ( Siboga Stn. 49); W of Jampea I., Selayar Islands Regency ( Siboga Stn. 65); Borneo bank, off SE Kalimantan ( Siboga Stn. 77); between Misool I. and West Papua, Raja Ampat Regency ( Siboga Stn. 164); off the W coast of Kur I., Moluccas ( Siboga Stn. 250); NE of Aru Islands Regency ( Siboga Stn. 274); Sailus Besar I., Pangkajene Dan Kepulauan Regency ( Siboga Stn. 315) ( Billard 1913, as Nemertesia indivisa ); Bunaken Marine Park, North Minahasa Regency (present study); Komodo National Park, West Manggarai Regency (present study; N.B.: the specimen illustrated in Fig. 1B View FIGURE 1 was not sampled and therefore not included in the specimens examined in the formal identification). Japan: Okinawa I. ( Schuchert 2015, as Nemertesia sp. 1 ) ( Fig. 19 View FIGURE 19 ). Depth range: 9–29 m (compiled from the references given in the synonymy and our data).


Museum National d'Histoire Naturelle


Museum of the Earth, Polish Academy of Sciences














Sciurella indivisa Allman, 1883

Galea, Horia R., Maggioni, Davide & Di Camillo, Cristina G. 2021

Sciurella indivisa

Kirkpatrick, R. 1890: 609
von Lendenfeld, R. 1887: 28
von Lendenfeld, R. 1885: 479
von Lendenfeld, R. 1885: 626
von Lendenfeld, R. 1885: 644
Allman, G. J. 1883: 9