An illustrated key to the species of the genus Narella (Cnidaria, Octocorallia, Primnoidae) Author Cairns, Stephen D. Author Taylor, Michelle L. text ZooKeys 2019 822 1 15 http://dx.doi.org/10.3897/zookeys.822.29922 journal article http://dx.doi.org/10.3897/zookeys.822.29922 1313-2970-822-1 1589695A15C044719F8A5BB3912EF356 Genus Narella Gray, 1870 Narella Gray, 1870: 49; Cairns and Bayer 2009 : 43. Stachyodes Wright & Studer in Studer 1887 : 49. Calypterinus Wright & Studer in Studer 1887 : 49-50. Diagnosis. Colonies branched dichotomously (laterally or equal), pinnately, in a lyrate fashion, or unbranched. Polyps arranged in whorls, all polyps facing downward in contracted condition. Each polyp covered with three (rarely four) pairs of abaxial body wall scales (i.e., one pair of basals, one or rarely two pairs of medials, and one pair of buccals) and a variable number of pairs of smaller adaxial scales, nonetheless leaving the adaxial face largely naked. Articular ridge not present on basal scales. Paired infrabasal scales often present. Opercular scales keeled on inner surface. Coenenchymal scales thin and imbricate or thick and mosaic in placement, and sometimes prominently ridged. Type species. Primnoa regularis Duchassaing & Michelotti, 1860, by monotypy. Discussion. Currently there are 50 valid species in the genus Narella , the most speciose in the family Primnoidae ( Taylor and Rogers 2015 ). The species in both keys (tabular (Table 1) and polychotomous, below) are roughly presented in an order that follows the major characters as outlined below, these characters we purport to be valuable in the distinction of species of Narella . Table 1. Tabular Key to the species of the genus Narella .

Species	Dorsolateral edge of basal scale	Pairs of body wall scales	Polychaete commensalism	Branching mode	Body wall scale thickness	Coenenchymal scales: imbricate, thickness; ridged	Polyps/whorl; whorl diameter (mm)	Polyp length (mm)	Distal edge of basal scales	Other characters	Geographic and depth range
N. macrocalyx
N. gilchristi
N. ferula
N. hawaiinensis
N. muzikae
N. merga
N. fordi
N. cristata
N. alvinae
N. bayeri
N. alaskensis
N. arbuscula
N. pauciflora
N. bowersi
N. gaussi
N. parva
N. regularis
N. valentine
N. virgosa
N. bellissima
N. ornata
N. spectablis
N. abyssalis
N. laxa
N. horrida
N. hypsocalyx
N. clavata
N. ambigua
N. aurantiaca
N. leilae
N. alata
N. vermifera
N. allmani
N. obscura
N. dampieri
N. mosaica
N. vulgaris
N. orientalis
N. calamus
N. versluysi
N. speighti
N. grandiflora
N. studeri
N. biannulata
N. candidae
N. japonensis
N. gigas
N. dichotoma
N. megalepis
N. compressa

Dorsolateral edge of basal scale ridged or not ridged: The dorsolateral edge (the point of inflexion of the scale from the dorsal region to the lateral region) of the basal scale is consistently ridged or not ridged (Fig. 1E) in each species, with the only exception of N. macrocalyx , which is inconspicuously ridged, and sometimes (rarely) lacks the ridge. This external ridging may help give strength to basal sclerites. The ridge may be single and extend from the base to the tip of the scale (Fig. 1A, B), or partial, occurring only at the base of the scale (Fig. 1C). Or, there may be multiple short ridges occurring in this region of the scale (Fig. 1D). The ridges may be tall or low. This character is relatively easy to observe, but usually requires the removal of a polyp from a whorl, drying the specimen, and then applying a dye to help see the characteristic ridging structure. Number of pairs of body wall scales: Most species of Narella have three pairs of abaxial body wall scales (basal, medial, and buccal, Fig. 1F), but in three species there is consistently an extra pair of medial scales (Fig. 1G). Also, specimens of some species that have otherwise three pairs of body wall scales will have occasional polyps with four pairs of body wall scales. This is a fairly easily observed character when using a dissecting microscope. Worm commensalism: The commensal association with a polychaete worm, usually a polynoid ( Cairns and Bayer 2008 , Cairns 2012 , Britayev et al. 2014 , Serpetti et al. 2017 ), is considered to be characteristic of the species, and is easily observed even without a microscope. The facing basal scales of two adjacent polyps are greatly enlarged and modified (reflexed) in order to make an elongate cylindrical tube for the worm (Fig. 1H, I). Branching mode: The mode of branching, and thus colony shape, is considered to be characteristic of the species. Modes include: unbranched (Fig. 1J), branching from a common basal coenenchyme or bolus (Fig. 1K, L), sparse equal dichotomous branching (Fig. 1M), equal dichotomous branching (Fig. 1N), and lyrate (Fig. 1O), which is often followed by dichotomous branching. Lyrate branching might be considered as a special case of dichotomous branching in which the outer component of each bifurcation maintains a straight line while the inner branches remain roughly parallel to one another. Figure 1. A, B lateral view of a polyp showing dorsolateral ridge for entire height of basal scale (A N. parva from Versluys (1906) B N. bayeri ) C basal scale of N. hawaiinensis showing dorsolateral ridge only on lower half of scale D whorl of polyps of N. pauciflora showing multiple dorsolateral ridges on the basal scales E basal scale of N. vulgaris showing the lack of a dorsolateral ridge, and a lobate distal edge F polyp whorl of N. bellissima showing the three pairs of body wall scales G polyp of N. laxa having four pairs of body wall scales H polyp whorl of N. hypsocalyx showing highly modified basal scales forming a cross section view of a cylindrical worm tube I polyp whorl of N. vulgaris showing highly modified basal scales forming a lateral view of a cylindrical worm tube J unbranched colony of N. versluysi K, L branching from a basal bolus of N. hypsocalyx M sparse, dichotomous branching of N. macrocalyx N equal, dichotomous branching of N. vulgaris O lyrate branching of N. bellissima P massive basal scales of N. clavata . Body wall scale thickness: In some species the body wall scales are quite thick, or massive (Figs 1P, 2A). This trait is often correlated with having thick coenenchymals as well (see next character). This character is best seen using scanning electron microscopy of individual sclerites. Coenenchymal scales arrangement and ornamentation: The coenenchymal scales of most species are relatively thin, having the same thickness as a body wall scale, and have edges that slightly overlap those of other adjacent coenenchymal scales (Fig. 2B). But some species have quite thick scales (Fig. 2 C-D ) that are so massive that they cannot overlap adjacent scales and thus produce a mosaic, polygonal, or tessellate pattern, also called "cobblestone" ( Williams 1992 ). The term mosaic is used herein. Coenenchymal scales usually have a finely granular outer surface (Fig. 2D), but many species have scales that bear a single longitudinal (Fig. 2F) or multiple complexly arranged ridges (Fig. 2E). If these ridges are quite tall they have been termed sail scales ( Cairns 2016 )(Fig. 2G). Mosaic coenenchymals are not usually ridged (Fig. 2D). This character is best seen using SEM. Polyps/whorl; whorl diameter: Although every specimen and species has a range of polyps/whorl and whorl diameter, sometimes these numbers help to differentiate species. This character is easily determined using a dissecting microscope. Polyp length: As above, this character has a range for every specimen and species, but can sometimes differentiate among species. The polyp length is essentially the horizontal length of the polyp, which consist of the length of the buccal scale and whatever part of the operculars protrude from the buccal scale. This character is easily determined using a dissecting microscope. Shape of the distal edge of basal scales: The distal edge of the basal scales are usually slightly lobate and smooth (Fig. 1E), but in some species are serrate (Fig. 2H, K) or even spinose (e.g., N. horrida , Fig. 2I). It may extend far beyond its junction with the proximal edge of the medial scales as a cowl (Fig. 2K) or be quite short (Fig. 1D). This character is also easily determined using a dissecting microscope. Other characters: Other characters that are used to describe and differentiate species but are not consistently addressed in the keys include: shape and number of adaxial body wall scales (Fig. 2J), external ridging of the medial and buccal scales, closure of the body wall rings, aspects of the opercular scales, body wall formula (i.e., ratio of length of basal: medial: buccal scales), and number of polyps per cm. Geographic and depth range. All ocean basins, 128-4594 m ( Cairns 2012 ). Figure 2. A massive basal scales of N. clavata B thin, imbricate coenenchymal scales of N. fordi C thick, mosaic arranged coenenchymal scales of N. mosaica D individual thick coenenchymal scale of N. mosaica with a finely granular outer surface E complexly ridged coenenchymal scale of N. muzikae F single medial coenenchymal ridge of N. pauciflora G sail scale of N. spectabilis H serrate distal margin of body wall scales of N. bowersi I spinose body wall scales of N. horrida J adaxial body wall scales of N. dampieri K polyp pair of N. leilae showing extensive cowl and serrate distal edges of body wall scales (from Bayer, 1951).