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Gulf Research Reports Volume 8 Issue 3 January 1991 Six New Genera in the Chaetognath Family Sagittidae Robert Bieri Antioch College DOI: 10.18785/grr.0803.01 Follow this and additional works at: http://aquila.usm.edu/gcr Part of the Marine Biology Commons Recommended Citation Bieri, R. 1991. Six New Genera in the Chaetognath Family Sagittidae. Gulf Research Reports 8 (3): 221-225. Retrieved from http://aquila.usm.edu/gcr/vol8/iss3/1 This Article is brought to you for free and open access by The Aquila Digital Community. It has been accepted for inclusion in Gulf and Caribbean Research by an authorized editor of The Aquila Digital Community. For more information, please contact Joshua.Cromwell@usm.edu.

Gulf Research Reports, Vol. 8, No. 3, 221-225, 1991 SIX NEW GENERA IN THE CHAETOGNATH FAMILY SAGITTIDAE ROBERT BIER1 Biology Department, Antioch College, Yellow Springs, Ohio 45387 ABSTRACT The following six new genera of Sagittidae, with type species listed in parentheses, are proposed: Adhesisagitta (Sagitta hispidu Conant, 1985), Demisagitta (Sagitta demipenna Tokioka and Pathansali, 1963), Decipisugitta (Sugittu decipiens Fowler, 1905). Tenuisugittu (Sagitta tenuis Conant, 1986), Abacisagitta (Sagitta pulchra Doncaster, 1903), Oculosagitta (Sagitta megalophthalma Dallot and Ducret, 1969). INTRODUCTION While reviewing recently (Bieri, In Press) the genera of the chaetognath family Sagittidae, I noticed that some genera are heterogeneous and therefore neither parallel nor symmetrical with the other genera of the family. Among the asymmetries are differences in fin ray distribution, body proportions, body texture, and the shape and structure of the seminal vesicles. To correct these heterogeneous assemblages I propose the following new genera: Sagittidae Tokioka, 1965 Adhesisagitta, new genus Figure 1 Type species - Sagitta hispida Conant, 1895, by monotypy and present designation. Name - From the Latin adhaesus, clinging to, referring to the habit of this quasi-planktonic species of clinging to the substrate. Definition - Corona Ciliata long, extending from just anterior to eyes to well posterior on trunk but not reaching ventral ganglion; intestinal diverticula present; lateral fins completely rayed; body rigid; seminal vesicles slightly swollen at anterior end when mature, but not forming distinct knob ovaries moderately long and narrow with ova irregularly spaced rather than in a single row; mouth appearing sutured or buckled by 4 rodlike structures or bars about 1/3 as long as distance between them; species can cling to vertical substrate. Species included - One, Sagitta hispidu. Discussion - Published drawings and descriptions of this species are not consistent with one another in several critical features. Figure 1 is a composite of the drawings of Pierce (1951), Tokioka (1955), Michel Manuscript rcccived February 26, 1990; acccptcd March 9, 1990. (1984), McLelland (1989), and my own observations. In distinguishing Adhesisagitta from Sagitta and Ferrosagitta the structure of the seminal vesicle and variations in its appearance at different stages of maturity are important and are best illustrated by Tokioka (1955). Although perhaps not of generic significance, the space between the seminal vesicle and the tail fin is about 1/2 the length of the vesicle. Pierce shows the vesicle in contact with the tail fin, and Michel, although stating in her text that it is widely separated from the tail fin, illustrates it as rather close, as does McLelland. The irregular arrangement of the ova is distinctive. Although the number of chaetognath species examined with SEM now exceeds 17 (Thuesen et al 1988), A. hispida is the only species found to have well developed bars or buckles across the mouth. They are shown exceptionally clearly in the SEM photograph of Cosper and Reeve (1970). It is also the only sagittid known to cling habitually to the walls of glass aquaria (H. Michel, pers. comm.) and to eel grass (McLelland, pers. comm.), a behavior that may be related to its hyper-neritic distribution (Pierce, 1951). The conjectured buckles may function as adhesive organs. Adhesisagitta is distinguished from the possibly closely related genera Sagitta (sensu Tokioka) and Ferrosagitta not only by the mouth buckles but also by its clinging ability, shape of the seminal vesicles, and the irregular arrangement of the ova. Dcmisagitta, new genus Figure 2 Type species - Sagitta demipenna Tokioka and Pathansali, 1963, by montypy and present designation. Tokioka (1965) placed S. demipenna in Aidanosagitta Tokioka and Pathansali, 1963 Name - From the trivial name of the type species, referring to the short or half posterior fin, which is limited to the tail. 22 1

222 BIERI Heres 14. Composite and schematic habltns drawings of Chaetognatlta, In dorsal dew. (1) Adk&u@ilta hhpidu, based on Pierce 1951, Tokloka 1955, Mkhel 1984, McLeUand 1989, and the author s observations. (2) Dcmisa$h~ demiptnnu, based on Toktoka and Pathansell 1963, Pathansaw 1974, and Alvariiio 1967. (3) Decipkagi#a dtclpisns, based on Werrot- Bulta 1979, McLelland 1989, and the author s observetlons. (4) Tcnukugitta tcnuis based an Pierce 1951, Mclelland 1980, and the author s observations, (5) Abnciasagitta pufchro, bawd on Tokloka 1966, Alvsrliio 1967, and the author s observatlons. (6) Qculusagilta megdophthlrlma, based on DaIlot end Qucret 1969, and Mlchel 1984.

SIX NEW GENERA IN THE CHAETOGNATH FAMILY SAG 223 Definition - Very similar to Aidanosugittu as defined by Tokioka (1965). Corona ciliata beginning posterior to eyes; intestinal diverticula present; lateral fins completely rayed, rays at anterior ends of fins perpendicular to body wall; body very stiff and rigid, wider than in Aidunosugittu, more like Spudellu and Pterosugittu; tail relatively long; seminal vesicle situated just behind posterior fin and separated from tail fin, no extemal differentiation of anterior glandular part; mature ovaries reaching ventral ganglion, ova in single row, so large that they crowd and distort gut, maturing asynchronously; posterior fin extending anteriorly only to level of transverse septum. Species included - One, Demisugittu demipennu. Discussion - No other species of Sagittidae has the posterior fin limited to the tail. This remarkable feature clearly separates D. demipennu from all other known sagittids and requires modification of the usual textbook definition of Sagittidae. If D. demipennu were to lose the anterior fin and develop a pair of large hair fans, it would fit into the genus Pterosugittu, which demonstrates the close affinity of the Pterosagittidae to the Sagittidae. The occurrence of a demi-fin in the relatively short, widebodied, and heavily muscled genera Pterosugittu and Spudellu as in Demisugittu argues for a specialized form of locomotion associated with the short, powerful muscles and the short, stiff demi-fin. Discussion - Erection of this genus leaves Mesosugittu with a single species, M. minima (Grassi, 1881). Of the four species listed above, only two were recognized as valid by Pierrot-Bulb in her 1979 review of the group. Decipisugittu butuvu may have been described from deformed specimens of Sagitta setosu. It has not been reported since the original description by Bierstecker and van der Spoel (1966), who said that small intestinal diverticula, show in their figure 2, are usually present. No other chaetognath has been reported with diverticula usually present ; they are either present or absent. Although possibly due to poor preservation, this type of preservational artifact has not been reported previously (Bieri, 1989). The other three species of Decipisugittu are mesoplanktonic, whereas D. batuvu is epiplanktonic, which is also anomalous. However, the original description is so well done that the species should be retained pending further study. Although Pierrot-Bulb synonymized Sagitta neodecipiens Tokioka (1959) with S. decipiens Fowler (1905), there are very significant differences in the arrangement of the fin rays as shown by Tokioka (1959) in his original description of S. neodecipiens and in S. decipiens as drawn by Pierrot-Bults from the lectotype and pardectotype. In view of these striking differences and the differences in the eye pigments, S. neodecipiens should be retained as a valid species. Decipisagitta, new genus Figure 3 Type species - Sagitta decipiens Fowler, 1905. Name - From the trivial name of the type species. Definition - Agreeing mostly with Tokioka s (1965) definition of Mesosugittu. Corona ciliata beginning on neck and extending posteriorly onto anterior part of trunk; intestinal diverticula present; fin rays diagonal to body wall rather than perpendicular as in Mesosugittu, rays completely appressed, not irregularly spaced, missing in some areas of both lateral fins; more of posterior fin on trunk than on tail; seminal vesicles separated from posterior fin; maximum body width rather spread out; between 1/3 and 1/2 of trunk length anterior to transverse septum, in contrast to Mesosugittu in which less than 1/4 of trunk is anterior to transverse septum; no marked constriction of tail at transverse septum as in Mesosugittu; body muscles weakly developed, stronger than in Fluccisugittu but weaker than in Serrutosugittu, stronger and more opaque than in Mesosugittu; mature ovaries approaching but usually not reaching ventral ganglion, relatively long and narrow with ova in single row. Included species - Four: Decipisugittu decipiens, D. sibogue, D. neodecipiens, and possibly the doubtful species, D. butuvu. Tenuisagitta, new genus Figure 4 Type species - Sagitta tenuis Conant, 1896. Name - From the trivial name of the type species. Definition - Corona ciliata very elongate, extending from just anterior to eyes to well onto trunk; intestinal diverticula absent; lateral fins completely rayed, rays completely appressed; body rather transparent with longitudinal muscles of moderate strength, weaker than in Sagitta but stronger than in Mesosugittu; seminal vesicles simple but with circular process at anterior end that becomes ovoid when fully mature; ovary variable, in small species with ova arranged in single row and maturing synchronously, in large species with ova arranged in double row and sometimes maturing asynchronously. Species included - As presently understood, six species: Tenuisugittu tenuis, T. setosu, T. fnderici, T. euneriticu, T. peruviunu, T. popvicci. Discussion - Although Tenuisugittu bears a strong superficial resemblance to Purusugittu, the two genera differ in several significant features, including the presence in Purusugittu of intestinal diverticula, and large vacuolated gut cells containing NH4+ rather than Na+ resulting in a lower body density in Purusugittu than in Tenuisugittu (Bone et al., 1987). All included species of

224 BIBRI T~nuisogitra am epiplanktonic and more or less neritic. Abdmagitta, new genus Figure 5 Type species - Sagitto pufchra Doncaster, 1903, by monotypy and present designation. Name - From the Latin lab, off, away from, plus acia, thread, referring to lthe partial lack of rays in the lateral fins. Definition - Corona ciliata very long, extending fmm just anterior to eyes to well onto mk, intestinal diverticula absent: lateral fins with conspicuous rayless zones, especially large in posterior fins, much larger than in Zunosagitta; head not as wide as maximum body width which is a[ about midength; body muscles weaker and more mslucent than io Zonusagitra; anterior fins flatsided and broadest near posterior end, seminal vesicle in two parts rather than forming bulb as in Zunosugitta; ovary very Iong, reaching from midbody to venw ganglion when mature, ova viewed dorsally usually in a single row; tail about 1/5 as long as head and trunk coni bined. Species included - One, Sagitto pulchra. Discussion - Tokioka (1965) included S. pulchra in his new genus Zorrosagitta. Alvarino (1967), on the other hand, excluded it from her M~ti group C = &nosagim), stating that This species is not included in any group. In view of the fin differences as well as differences in the body muscles and the seminal vesicles, this species merits its own genus. Qculosagitte, new genus Flgnre 6 Type.s@es - Sagitro megduphrhoho Dallot and Ducret, 1969, by monotypy and present designation. Name - F m the Latin oculus, eye, referring to the importance the authors of S. tneggaloplitkalma placed on the size of the eye pigment. Definition - Corona ciliata ve y long, extending from just anterior to eyes to well back onto trunk but not reactiing vend ganglion: intestinal diverticula absent; lateral fms completely raye4 all rays appressed one to another, M y moderately firm; head barely wider thau mmimum lmdy with which is at about 1/3 of mnk length anterior to transversed septum; M y width tapering gradually towards head and tail, without marked consb-iction at msverse septum; seminal vesicle simple, ovoid, touching tail fin but widely separated from posterior fin; ova reaching only to anterior end of posterior fin (mature specimens may not yet be known), ova in double row or imgularly spaced when viewed dorsally; gut with large vacuolated cells in middle two-thilds easily visible in dorsal view, these cells may line the entire gut in juveniles. Species included - One: Sagitta megalophthabna. Discussion - In their original description Dallot and Dumt discussed Tokioka s genera and Alv~uiRo s groups and decided that S. megalophthalma did not fit into any of the extant genera or groups. Although somewhat similar to Sagitfa bipunctum and vaguely reminiscent QE Mesosagitta minim, the species does not fit in any sagittid genus and therefore merits its own genus. The large, heavily pigmented eyes, large head, slender body, small cellarette, and long series of vacoolated gut cells set it off from all other known species of Sagittidae. The figures am composites of figures published by several different authors as noted with mdifications based on the author s expeience. The separation of the fin rays, which are actually closely appressed, is much exaggerated. ParticuIar attention has been given tu body pmpod~n~, relative head width, location of seminal vesicles, extent of ovary and arrangement of ova, and disposition of the fins in relation to the ventral ganglion and msverse septum. A less than successfue attempt has been made to indicate the stiffness of the body musculature by the degree of shading. The author is personally well acquainted with all the type species from well preserved formalin specimens with the exception of Dem isagittu demipenna and Oculosagitta megalapihaha. Alvatifio, A. 1967. The Chattopatha of the Nag8 Expedition (1959-1961) in &t S ~ t h china S a and the Gulf of Thailand. Nagu Rep. 4(2):1--197. Bieri, R. 1989. Krohnittllidae and BathybeLdae, new fdes in the phylum Chaetopatha; the rejection of the f d y Tokiokai spadellidae and the genera Tokioka ispudella, Zehonya, and Aberrospudello. Pro E, Bioi. SOC. Wash, 102(4):973-976. In Press. Systematics of the Chattognatha. In! Q. Bone, A.C. Pierrot Bults, and H. Kapp, (eds), The Biology of?he Chactognotha. Uaivtrsity Press, Oxford. Bitrsteker, R. IT., & S. van der SpocL. 1966. Sagitta batova n. sp. from the Scheldt Estuary, the Nethedatds ( Chaet opah). Beu ufa rria 14( 167):6 1 4. Born, Q.q C. Bmwnlee, G. W. Bryan. G. R. Burt, P. R. Dando, M. E. Liddicoat. A. L. Pulrrfod, & K. P. Ryan 1987. On

~ and SIX NEW GENERA IN THE WOGNATH FAMILY SAGITITDM 225 the differences between the two indicator species of chaetognath, Sagitta setosa and S. elegans. J. Mar. Biol. ASSOC. U.K. 67545-560. Conant, F. S. 1985. Description of two new chaetognaths (Spadella schizoptera and Sagitta hispida). Ann. Mag. Nut. Hist. (6)16288-292.. 1896. Notes on the chaetognaths. Ann. Mag. Nut. Hist. (6)18:201-214.. Cosper, T. C., & M. R. Reeve. 1970. Structural details of the mouthparts of a chaetognath as revealed by scanning electron microscopy. Bull. Mar. Sci. Gulf. Caribb. 20:441-445. Dallot, S., & F. Ducret. 1969. Un chaetognathe m6soplanctonique nouveau: Sagitta megalophthalma sp. n. Beaufortia 17:13-20. Doncaster, L. 1903. Chaetognatha, with a note on the variation and distribution of the group. Fauna Geogr. Maldive- Laccadive Archipel. 1:209-218. Fowler, G. H. 1905. Biscayan plankton collected during a cruise of the H.M.S. Research, 1900. IU. The Chaetognatha. Trans. Linn. SOC. London (Zool.) 10(3):55-87. Grassi, B. 1881. Intomo ai Chaetognati. Rend. R. 1st. Lombard0 SC. Lett., (2) 14199-214. McLelland. J. A. 1980. Notes on the northem Gulf of Mexico occurrence of Sagitta friderici Ritter-Zahoney (Chaetognatha). Gulf Res. Rept. 6(4):343-348.. 1989. An illustrated key to the Chaetognatha of the northem Gulf of Mexico with notes on their distribution. Gulf Res. Rept. 8(2): 145-172. Michel, H. B. 1984. Chaetognatha of the Caribbean Sea and adjacent areas. NOAA Tech. Rep. NMFS 15, U.S. Dept. of Commerce. 33 pp. Pathansali, D. 1974. Chaetognatha in the coastal waters of peninsular Malaysia with descriptions of two new species. Fish. Bull. No. 2, Minist. Agric. Rural Develop. Malaysia, 29 pp. Pierce, E. L. 1951. The Chaetognatha of the west coast of Florida. Biol. Bull. 100.206-228. Pierrot-Bults, A. C. 1979. On the synonymy of Sagitta decipiens Fowler. 1905. and Sagitta neodecipiens Tokioka, 1959, and the validity of Sagitta sibogae Fowler, 1906. Bull. Zool. Mus. Univ. Amst. 6137-143. Ritter-Zbhony, R. yon. 1911. Revision der Chaetognathen. Dtsch. Siidpol. Exped. 1901-3. 13(1):1-71. Thuesen, E. V., S. Nagasawa, R. Bieri, & T. Nemoto. 1988. Transvestibular pores of chaetognaths with comments on the functions and nomenclature of the vestibular anatomy. Bull. Plankton SOC. Jpn. 35133-141. Tokioka, T. 1955. Notes on some chaetognaths from the Gulf of Mexico. Bull. Mar. Sei. Gulf Caribb. 552-65.. 1959. Observations on the taxonomy and distribution of chaetognaths of the North Pacific. Publ. Set0 Mar. Biol. Lab. 7:349-456.. 1965. The taxonomical outline of Chaetognath. Publ. Set0 Mar. Biol. Lab. 12335-357.. 1966. Phylum Chaetoptha. Pages 181-190 in: I. Yamazi, Illustrations of the Marine Plankton of Japan. Hoikusha hbl. Co., Osaka, Japan. & D. Pathansali. 1963. Another new chaetognath from Malay waters, with a proposal of grouping some species of Sagitta into subgenera. Publ. Seto Mar. Biol. Lab. 11~119-123. II Robert Bieri died on July 31,1990 after a lengthy bout with cancer. He was Professor Emeritus of Environmental Studies at Antioch College in Yellow Springs, Ohio, where he taught for 30 years. He previously held posts at the Scripps Institution of Oceanography and the Lamont-Doherty Geological Observatory of Columbia University. His numerous expeditions included excursions to the Arctic Ocean, the Pacific Coast of Mexico, the Caribbean Sea, the Mediterranean Sea, and the Boreal North Atlantic. He spent two sabbatical years in Japan and three months in Ecuador as a visiting scientist at the Instituto Oceanographico de la Armada. He made numerous contributions to the knowledge of chaetognath biology and systematics, will be greatly missed as a reviewer and contributor to Gurf Research Reports. II