Aglaopheniid hydroids (Cnidaria: Hydrozoa: Aglaopheniidae) from off New Caledonia collected during KANACONO and KANADEEP expeditions of the French Tropical Deep-Sea Benthos Program Author Galea, Horia R. text European Journal of Taxonomy 2020 2020-03-13 615 1 47 journal article 10.5852/ejt.2020.615 b2465730-6237-4a14-a98d-7fd2d1096cb0 3711000 637FC87F-13B5-4B32-BC52-11A9B30ECF1D Macrorhynchia spiralis sp. nov. urn:lsid:zoobank.org:act: DA1E42B9-C682-4639-B3B1-4A9A7E2C7143 Figs 16 E–G, 19; Table 3 Diagnosis Species of Macrorhynchia with a distinctly spirally-twisted stem, giving rise at each bend to a side branch that branches soon again, forming a bifid ramification. Stem and side branches strongly fascicled. Internodes short, with a proximal frontal nematotheca, a cladial apophysis above with its simple, conical nematotheca and a fronto-axillar nematotheca; cauline nematothecae saccate, with deeply scooped adaxial walls. Cladia alternate, fairly close to one another; cormidia moderately long, hydrotheca cupshaped, rather deep, aperture slightly sloping frontally, rim with 9 triangular cusps with rounded tips, of which the median, abaxial one is the most prominent; a slightly sigmoid, adaxial, intrathecal septum given off softly obliquely, reaching almost to abaxial wall; mesial nematotheca with short, spoutlike aperture, not reaching middle level of hydrotheca; lateral nematothecae barrel-shaped, with rim scooped out adaxially. Pseudophylactocarps, composed of chains of atrophied hydrothecae, borne on proximal parts of branches. Phylactocarp borne on a modified cladium, after 1–2 normal cormidia; first (occasionally second) segment, an atrophied hydrotheca, with its mesial nematotheca displaced more distally, and a short, lateral projection (on one side) bearing a couple of nematothecae similar to the laterals; gonotheca lenticular, flimsy, on top of atrophied hydrotheca, between the lateral nematothecae; gonophore, a medusoid. Etymology From the Latin ‘ spīra ’, meaning ‘spiral’, with reference to the distinctive shape of the stem. Material examined Holotype PACIFIC OCEAN • upper part of 1 colony, ca 6.5 cm high, with female gonophores; off New Caledonia , stn DW4954; 24°12ʹ S , 159°41ʹ E ; 300 m ; 5 Sep. 2017 ; KANADEEP leg.; MNHN-IK-2015-519. Paratypes PACIFIC OCEAN • upper part of 1 colony , ca 8 cm high, with gonophores; off New Caledonia , stn DW5007 ; 22°12ʹ S , 159°02ʹ E ; 290–750 m ; 19 Sep. 2017 ; KANADEEP leg.; MNHN-IK-2015-520 • 1 entire sterile colony, ca 10.5 cm high; off New Caledonia , stn DW5008; 22°12ʹ S , 159°02ʹ E ; 300–750 m ; 19 Sep. 2017 ; KANADEEP leg.; MNHN-IK-2015-522 . Description Colonies erect, sympodially built, with fascicled stems and side branches, grading to monosiphonic distally. Stem straight basally for a varied length, then bends at irregular intervals adopting a spiral arrangement. At each bend, a side branch arises as a rectilinear prolongation of the preceding portion of the stem, and usually ramifies once a short distance from its origin, both branches bearing hydrocladia for most of their length; proximally, a linear succession of saccate nematothecae with large apical foramina. Division into internodes indistinct wherever the perisarc is thick, but becomes evident towards the distal parts of the branches; internodes relatively short ( 400–420 µm long, 245–250 µm wide at node), with a large, proximal nematotheca, a cladial apophysis above, a fronto-axillar nematotheca, and a simple, conical nematotheca (with small, rounded, apical aperture) on the basal part of apophysis; nematothecae saccate, adnate for most of their length, having apically a large aperture whose rim is lowered adaxially. Cladia alternate, up to 14 mm long, composed of short cormidia ( 475–490 µm long, 95–100 µm wide at node), each accommodating a hydrotheca and its three nematothecae: one mesial and a pair of laterals. Hydrotheca cup-shaped, rather deep ( 390–400 µm ), abaxial wall 215–240 µm long, straight for most of its length, gently expanding towards aperture; the latter 205–215 µm wide, slightly sloping frontally; rim with 9 rounded cusps, the uneven, abaxial one being the most prominent, and slightly inwardlycurved, the 4 pairs on the edges being comparatively broader and of varied development; an internal septum, of a slightly sigmoid appearance, is given off softly obliquely from the lower part of the adaxial thecal wall, penetrating deep into the lumen, almost reaching the abaxial wall; mesial nematotheca fused to the abaxial wall of hydrotheca for most of its length, leaving distally a short ( 45–60 µm long), spout-shaped aperture, not reaching the middle of the hydrotheca; lateral nematothecae broadly barrelshaped, 110–120 µm long and ca 50 µm wide at aperture, rim scooped out on adaxial side. Fertile colonies produce two distinct structures: one not associated to the gonosome, the other being part of it. The former are pseudophylactocarps, composed of a chain of modified hydrothecae devoid of lumina, but retaining their complement of nematothecae (the latter are all tubular, with a rounded distal aperture and an ovoid, proximal, adaxial one); they are abundantly borne on small, alternate apophyses arising regularly and slightly laterally to the nematothecae of the proximal (acladiate) portion of branches, and encircle their long axis; no gonothecae have been observed associated with the pseudophylactocarps. The gonosome proper is borne on the distal parts of distally-modified cladia that retain proximally 1 (occasionally up to 4) normal cormidia, followed by 1–2 elongated internodes representing atrophied hydrothecae (devoid of lumina) that retain their normal complement of nematothecae, in addition to which a pair of nematothecae (similar to the laterals) is mounted on a conspicuous lateral, subterminal projection. There is some variation in the structure of the proximal part of the fertile cladium: the proximalmost hydrotheca (when only one is present) may be suppressed and replaced by one or two short internodes, the proximalmost bearing one nematotheca, the second three (one mesial and a pair of laterals). Gonothecae lenticular, 1150–1380 µm long and 750–795 µm wide, with flimsy perisarc, inserted between the pair of lateral nematothecae of the distalmost, atrophied hydrotheca; the gonophore is a medusoid, composed of a central spadix encircled by a single layer of large oocytes (the holotype is a female colony), and bearing apically a belt of refringent corpuscles. Fig. 19. Macrorhynchia spiralis sp. nov. , from sample MNHN-IK-2015-519. A– B . Portion of stem (A) and detail of the nematothecae from the vicinity of a cladial apophysis (B). C . Pseudophylactocarp. D–E . Hydrotheca(D) and one of its lateral nematothecae (E), both in lateral view. F–H . Same phylactocarp with empty gonotheca in two different views (F–G), and phylactocarp with two gonothecae, one of which bears a female medusoid gonophore (H). Scale bars: A, D = 200 µm; B, E = 100 µm, C, F–H = 500 µm. Table 3. Nominal species of Macrorhynchia Kirchenpauer, 1872 with saccate hydrothecae and adaxial but no abaxial, intrathecal septa, similar to those of Macrorhynchia spiralis sp. nov. , and their morphological differences to the new species. Nominal species Brief description, with emphasis on the distinctive features, and geographical distribution

M. allmani ( Nutting, 1900 )	Mesial nematotheca adnate for most of its length to the abaxial wall of hydrotheca, distal tip reaching the hydrothecal aperture. Gonosome described by Galea (2013) . Distribution: tropical western Atlantic ( Calder 1997 ).
M. clarkei ( Nutting, 1900 )	Mesial nematotheca adnate for most of its length to the abaxial wall of hydrotheca, distal tip reaching the hydrothecal aperture. Gonosome described by Ansín Agís et al. (2001) . Distribution: tropical western Atlantic, Cape Verde Islands ( Calder 2013 ).
M. curta ( Nutting, 1900 )	Poorly known species, with mesial nematotheca reaching middle part of the abaxial wall of hydrotheca. Gonosome unknown. Distribution: Bahamas ( Nutting 1900 ).
M. furcata ( Nutting, 1900 )	Poorly known species, with mesial nematotheca reaching middle part of the abaxial wall of hydrotheca. Gonosome unknown. Distribution: Bahamas ( Nutting 1900 ).
M. grandis (Clarke, 1879)	Poorly known species, with mesial nematotheca adnate for most of its length to the abaxial wall of hydrotheca, distal tip reaching or surpassing the hydrothecal aperture. Gonosome described by Nutting (1900) Distribution: tropical western Atlantic ( Calder 1997 ).
M. hawaiensis ( Nutting, 1905 )	Poorly known species, with mesial nematotheca becoming free in middle of hydrotheca, tip not reaching aperture; one median abaxial cusp and 2 pairs of 2 small, uneven lateral cusps . Gonophore described by Nutting (1905) . Distribution: Hawai’i ( Nutting 1905 ).
M. longiramosa ( Fraser, 1945 )	Poorly known species, with mesial nematothecae almost as long as the abaxial side of the hydrotheca, becoming free just below the thecal aperture. Gonosome unknown. Distribution: Dorsey Canyon, Gulf of Mexico ( Fraser 1945 ).
Poorly known species, with mesial nematotheca adnate for most of its length to the abaxial M. mercatoris wall of hydrotheca, tip almost reaching the rim; first cormidium with asymmetrical lateral ( Leloup, 1937 ) nematothecae. Distribution: Florida, USA ( Leloup 1937 ). Small hydroid. Hydrotheca constricted above the site where the mesial nematotheca becomes free; with abaxial carina ending distally in rounded crest ; mesial nematotheca partly adnate to lower abaxial wall of hydrotheca; lateral nematothecae reaching the rim; margin
M. mulderi (Bartlett, 1907)	with 1 median, abaxial cusp narrow and incurved, as well as 4 pairs of laterals: 1st rounded triangular, slightly projecting outwards , 2nd long, roughly rectangular , strongly incurved over hydrothecal aperture, 3rd broad, rounded at apex and projecting outwards , 4th long, narrow , slightly incurved over hydrothecal aperture ( Mulder & Trebilcock 1916 ). Gonosome described by Watson (1973) . Distribution: southern Australia ( Mulder & Trebilcock 1916 , Watson 1973 ).
M. racemifera ( Allman, 1883 )	Poorly known species, with mesial nematotheca adnate for most of its length to the abaxial wall of hydrotheca, distal tip overtopping the hydrothecal aperture. Gonosome described in the original account. Distribution: off Bahia, Brazil ( Allman 1883 ).
M. ramosa ( Fewkes, 1881 )	Poorly known species, with mesial nematotheca adnate for most of its length to the abaxial wall of hydrotheca, distal tip overtopping the hydrothecal aperture. Distribution: tropical western Atlantic ( Calder 1997 ).
M. robusta ( Fewkes, 1881 )	Poorly known species, with mesial nematotheca adnate for almost the whole length of the abaxial wall of hydrotheca. Gonosome unknown. Distribution: Montserrat ( Fewkes 1881 ). Poorly known species, with rather deep hydrothecae, with nearly horizontal aperture; rim with median, abaxial cusps and 3 pairs of laterals; mesial nematotheca becoming free in
M. similis ( Nutting, 1905 )	middle of hydrotheca, laterals tubular and narrow , distinctly overtopping the hydrothecal rim; phylactocarps with normal cormidium proximally, followed by a few short internodes carrying abortive (closed) hydrothecae that retain only their normal complement of nematothecae; no pseudophylactocarps . Distribution: Hawai’i ( Nutting 1905 ).

Remarks Hydrothecae with similar shape as those of M. spiralis sp. nov. , i.e., cup-shaped, with intrathecal adaxial but not abaxial ridge, are found in a number of congeners. Their distinguishing features with respect to the new species are emphasized in Table 3 . As noted above, this species produces a medusoid gonophore. At least three other nominal species reproduce in a similar way, namely M. filamentosa (De Lamarck, 1816) , M. philippina Kirchenpauer, 1872 and M. racemifera ( Allman, 1883 ) ( Galea 2018 ) , suggesting that this mode of dispersal could be the rule within the genus, as it probably is in the plumulariid genus Dentitheca Stechow, 1920 ( Galea et al. 2012 ). Distribution Known only from off New Caledonia (present study).