From Southwest Pacific Ocean Три Новых Вида, Два Новых Рода И Новое Семейство Biphragmosagittidae (Сhaetognatha) Из Юго-Восточной Части Тихого Океана

Home , Aphragmophora, Eukrohniidae, Parasagitta, Phragmophora, Pterosagittidae, Sagittidae

ZOOSYSTEMATICA ROSSICA, 20(1): 161–173 28 JULY 2011

Three new species, two new genera, and new family Biphragmosagittidae (Сhaetognatha) from Southwest Pacific Ocean Три новых вида, два новых рода и новое семейство Biphragmosagittidae (Сhaetognatha) из юго-восточной части Тихого океана

A.P. KASSATKINA

А.П. КАСАТКИНА

A.P. Kassatkina, Pacific Institute of Oceanography, Far East Division, Russian Academy of Sciences, 43 Baltiyskaya St., Vladivostok 690041, Russia. E-mail: [email protected]

Resuming published and own data, a revision of classification of Chaetognatha is presented. The family Sagittidae Claus & Grobben, 1905 is given a rank of subclass, Sagittiones, charac- terised, in particular, by the presence of two pairs of sac-like gelatinous structures or two pairs of fins. Besides the order Aphragmophora Tokioka, 1965, it contains the new order Biphrag- mosagittiformes ord. nov., which is a unique group of Chaetognatha with an unusual combination of morphological characters: the transverse muscles present in both the trunk and the tail sections of the body; the seminal vesicles simple, without internal complex compartments; the presence of two pairs of lateral fins. The only family assigned to the new order, Biphragmo- sagittidae fam. nov., contains two genera. Diagnoses of the two new genera, Biphragmosagitta gen. nov. (type species B. tarasovi sp. nov. and B. angusticephala sp. nov.) and Biphragmofas- tigata gen. nov. (type species B. fastigata sp. nov.), detailed descriptions and pictures of the three new species are presented. На основе литературных и собственных данных проведена ревизия типа Chaetognatha. Ранг семейства Sagittidae повышен до подкласса, Sagittiones, который охарактеризо- ван, в частности, наличием двух пар мешковидных желеобразных структур или двух пар плавников. Помимо отряда Aphragmophora Tokioka, 1965, этот подкласс содержит новый отряд Biphragmosagittiformes ord. nov., который является уникальной группой Chaetognatha с необычным сочетанием морфологических признаков: 1) поперечная му- скулатура имеется в полости туловищного и хвостового отделов, простые семенные ме- шочки без внутренних сложных камер; 2) две пары боковых плавников. Единственное семейство нового отряда, Biphragmosagittidae fam. nov., содержит два новых рода. При- ведены их диагнозы, а также диагнозы включенных видов – Biphragmosagitta gen. nov. (типовой вид B. tarasovi sp. nov. и B. angusticephala sp. nov.) и Biphragmofastigata gen. nov. (типовой вид B. fastigata sp. nov.) с детальными описаниями и изображениями. Key words: classification, Сhaetognatha, Eukrohniones, Sagittiones, Biphragmosagittiformes, Biphragmosagittidae, new subclasses, new order, new family, new genera, new species Ключевые слова: классификация, Сhaetognatha, Eukrohniones, Sagittiones, Biphragmo- sagittiformes, Biphragmosagittidae, новые подклассы, новый отряд, новое семейство, но- вые роды, новые виды

INTRODUCTION tive organs. Based on these descriptions he subdivided the phylum Chaetognatha, the While the phylogenetic position of class Sagittoidea, into two subclasses – Syn- Chaetognatha has became central to the gonata (with the ducts connecting the ova- question of early bilaterian evolution, the ries in the trunk section of the body and the internal systematics of the phylum are still testes in the tail section) and Chorismogo- not clear (Papillon et al., 2006). The history nata (with no ducts connecting the genitals of taxonomy of Chaetognatha was reviewed present). The Syngonata thus included only in details in some in early publications (e.g. a single order Biphragmophora Casanova, á Ritter-Z hony, 1911; Kuhl, 1938; Hyman, 1985 with a single family Heterokrohniidae 1959; Beauchamp, 1960). Tokioka (1965a, Casanova, 1985, and the Chorismogonata – 1965b) revised the classification based two orders, Monophragmophora Casanova, on the data on presence/absence of the 1985 (with families Spadellidae Tokioka, transverse coelomic muscles (the so-called 1965 and Eukrohniidae Tokioka, 1965) phragmes, or phragma). He considered that, and Aphragmophora Tokioka, 1965 (with from a morphological point of view, the most families Sagittidae Claus & Grobben, 1905, fundamental character was the presence or Pterosagittidae Tokioka, 1965, Krohnittidae absence of the phragmes, and thus defined Tokioka, 1965). However, the morphological the established orders, Phragmophora and data presented by Casanova (1985) were not Aphragmophora. Since then, Chaetognatha well supported by illustrative materials, and, has been typically divided into two orders, in some case, contradict to earlier known Phragmorphora and Aphragmorphora, each data on gonadal structure (e.g. Elpat’evskiy, comprising three families (Pierrot-Bults, 1910, 1913, 1914) and some new discoveries 2004). However Tokioka (1965a, 1965b) Kassatkina & Stolyarova, 2008). did not take into consideration morpho- Molecular data showed (Papillon et al., logical features of Krohnittella Germain & 2004, 2006) that there are three main mono- Joubin, 1912, a rare genus that does not phyletic groups: Sagittidae/Krohnittidae, have a transverse musculature but, by most Spadellidae/Pterosagittidae, and Eukroh- other characters, displays a clear similarity niidae/Heterokrohniidae. In this scheme, to Heterokrohnia Ritter-Záhony, 1911. This the group of Aphragmophora without is why, later, Bieri (1974) did not accept the Pterosagittidae (Sagittidae/Krohnittidae) Tokioka’s scheme (Tokioka, 1965a, 1965b) is monophyletic, the Spadellidae/Pterosag- with the two orders, Phragmorphora and ittidae and Eukrohniidae/Heterokrohni- Aphragmorphora. Dallot and Ibanez (1972) idae families are very likely clustered, the analysed 32 characters and received dif- Krohnittidae and Pterosagittidae groups ferent conclusions on taxonomic relation- should no longer be considered as families ships within the phylum: the order Aphrag- as they are included in other groups desig- mophora Tokioka, 1965 contained a single nated as families, and the Syngonata/Cho- family, Sagittidae, with two pairs of fins rismogonata and the Monophragmophora/ while genera with one pair of lateral fins, Biphragmophora hypotheses were thus Pterosagitta Costa, 1869 and Krohnitta Rit- rejected. More recent data (Jennings et al. ter-Záhony, 1910, that had been given rank 2010) also support the main divergence be- of families of their own by Tokioka (1965a) tween the Eukrohniidae+Heterokrohniidae were excluded. and Spadellidae s.l. further supporting the Casanova (1985, 1986) published on main scheme of Tokioka (1965a, 1965b) findings of new genera and species Archet- based on the transverse muscles presence/ erokrohnia rubra Casanova, 1986 and a new absence data. species Heterokrohnia murina Casanova, Salvini-Plawen (1986) gave reasons to 1986 with a description of their reproduc- consider Spadellidae as quite specialised

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 161–173 A.P. KASSATKINA. NEW CHAETOGNATH FAMILY, GENERA AND SPECIES FROM PACIFIC OCEAN 163 coastal chaetognaths rather than a group Yunnan, China (Eognathacantha ercainella ancestral for all other Chaetognatha. Tel- Chen & Huang, 2002 and Protosagitta spi- ford and Holland (1997) confirmed a dis- nosa Hu, 2005) and the Middle Cambrian tinct position of Spadella from all groups Burgess Shale of British Columbia (Oe- of Chaetognatha based on molecular data. sia disjuncta Walcott, 1911) (Szaniawski, Salvini-Plawen (1986) established three 2005; Vannier et al., 2007). A more recent species groups and proposed two new gen- chaetognath, Paucijaculum samamithion era based on the number of pairs of fins and Schram, 1973 has been described from the the locations and presence/absence of the Mazon Creek biota from the Pennsylva- digitate adhesive organs. While Bowman & nian of Illinois (Schram, 1973). As it can be Bieri (1989) agreed that the presence of ad- seen from the published pictures (e.g. that hesive organs is taxonomically important, of Protosagitta spinosa in Vannier et al., they questioned the importance of the num- 2007: fig. 1a), the chaetognaths from the ber of lateral fin pairs. They point out that Cambrian possessed the transverse muscles immature individuals of Paraspadella have (phragmes). only a single pair of fins, but, with sexual de- In general, the chaetognath locomotory velopment, the fins become separated into muscles exhibit astonishing variations. As anterior and posterior parts by the lateral described by Casanova and Duvert (2002), growth of the vagina. the transverse muscle, present in the less The Von Salvini-Plawen’s (1986) opin- evolved genera living near or on the bottom, ion on the ancestral state of the absence of has two forms: either classical cross-striated the phragmes was not confirmed by palae- or, in more or less benthic species, supercon- ontological data. There was published a de- tracting. Supercontraction is supposed to scription of the genus Parmia Gnilovskaya, be a derived character. The peculiarities of 1998 from southern Timan, northeast of the primary, secondary and transverse mus- the Russian Platform, – worm-like crea- cles of the truly benthic chaetognaths are tures inhabiting the ocean during the Late different from those of all the planktonic or Riphean of an approximate age of about 1 benthoplanktonic chaetognaths. These data Ga (Gnilovskaya et al., 2000, cited by Fe- confirm well the importance of the trans- donkin, 2003). Though Parmia is interpret- verse muscles structure for the taxonomy ed as a probable predecessor of the annelid in the Chaetognatha revealed by Tokioka worms (Fedonkin, 2003) this may be the (1965a). Schemes by Casanova and Duvert earliest chaetognath. I base this conclusion (2002) show the degree of reduction of the on the fact that the two well-preserved im- transverse musculature and its evolution in prints of the animal (Fedonkin, 2003: figs one-pair-of-fin Chaetognatha (Casanova & 6, 7) clearly show a regular homonomous Duvert, 2002: fig. 5, 2B). segmentation (7–11 segments per milli- Besides the transverse muscles structure meter of body length) that may represent and its presence/absence, the number of fins transverse muscles; any fins or sac-like ge- is also very important for understanding latinous structures (SGS) are absent. An- phylogeny and evolution of Chaetognatha; cient Chaetognatha were obviously able this character may even have a greater sig- to bend the body as it can be seen from the nificance because the fin development and, imprints but, having no fins or SGS, were respectively, the buoyancy, made it possible probably bottom dwellers. Several other for Chaetognatha to colonise open waters of fossil chaetognath species have been de- the ocean. scribed that appear to have originated in The most stable, during the develop- the Cambrian period (Vannier et al., 2007). ment and the maturation, morphological Complete body fossils are known from the character is, to my opinion, the number of Lower Cambrian Maotianshan shales of pairs of lateral fins. It was noticed as early

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 161–173 164 A.P. KASSATKINA. NEW CHAETOGNATH FAMILY, GENERA AND SPECIES FROM PACIFIC OCEAN as in first publications devoted to arrow As it was already mentioned above, the worms (Krohn, 1853; Langerhans, 1880; earlier opinion on the origin of Sagittidae Beraneck, 1895; Fowler, 1906). with two pairs of fins from Chaetognatha Ancient Cambrian Chaetognatha did with one pair of fins were caused by the in- not possess fins but already had SGS as it sufficient knowledge at that time. Findings can be seen, for example, in P. spinosa hav- of finless both Eukrohniidae and Sagittidae ing specific flaps on sides of the body and at (Kassatkina, 1998, 2001, 2002, 2003, 2006; the terminal body end (Vannier et al., 2007: Kassatkina & Stolyarova, 2006) provided fig. 1). Due to the presence of SGS, Cam- additional arguments against this opinion. brian Chaetognatha might have been able SGS could appear in different regions of the to dwell in pelagic waters. body; the presence of SGS instead of fins Sac-like gelatinous structures, which appears to be a plesiomorphic character re- serve for the establishing neutral buoyan- tained in a few recent species (Kassatkina, cy, are not fins in a proper sense. A fin fold 2006). Species with one pair and two pairs never has gelatinous filling as it occurs in of fins could evolve independently in differ- SGS. In contrast to SGS, the fins are rud- ent chaetognath lineages. I suppose that the ders adapted to strong surface flows in the chaetognaths with one pair of fins and the ocean (Ghirardelli, 1968). Sac-like gelati- chaetognaths with two pairs of fins repre- nous structures may be located in different sent independent clades originated from places of the body (on dorsal, ventral and Protochaetognatha, which possesses nei- lateral surfaces of the body) being separated ther fins nor SGS. Genetic data (Telford, by unequal gaps as it can be seen in Abosa- Holland, 1993, 1997; Thuesen et al., 1993) gitta, Accedosagittaminuta, Pseudosagitta, support a considerable distinctiveness of the chaetognaths with one pair of fins from Pseudeukrohnia, Protoeukrohnia, Praeu- the chaetognaths with two pairs of fins. krohnia and Paraeukrohnia (Kassatkina, In plankton samples from the south- 1998, 2001, 2002, 2003, 2006; Kassatkina & western part of the Pacific Ocean there Stolyarova, 2006, 2010). However, lateral were found unique Chaetognatha with SGS only have a triangular-convex shape. an unusual combination of morphological Flattened margins of the SGS can serve characters: 1) the transverse muscles pres- as fins, for example, in Paraeukrohnia. It ent in both the trunk and the tail sections of seems that the fins evolved from some SGS the body, the seminal vesicles simple, with- through their flattening and the appearance out internal complex compartments (that of rigid supporting elements, transversal is characteristic of Heterokrohniidae with and longitudinal, inside a SGS as can be one pair of fins); 2) presence of two pairs of seen in Praeukrohnia fidus (Kassatkina & lateral fins (earlier considered to be typical Stolyarovа, 2006) and A. minuta (Kassat- only for Sagittidae). This made it necessary kina, 1982). True fin rays are not similar to revise the whole system of Chaetognatha to these transversal and longitudinal ele- and find a position of the new taxa in it. The ments for they consist of elastoidin fibres as high-level classification of Chaetognatha in agnathans and fish (Reisinger, 1968). In has been revised as given below with de- Pennsylvanian Paucijaculum samamithion, scriptions of two new genera (three new that is undoubtedly a chaetognath, large species) belonging in a new family assigned rays can be seen; some of them are located in a respective new monotypic order. on the body aside of the fins while others, in the fin region (Schram, 1973). Most proba- MATERIAL AND METHODS bly, the fins with rays present in extant species are the result of an intergation of the fin Plankton samples were collected by fold and the rays. a hydrobiologist V.G. Tarasov from the

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 161–173 A.P. KASSATKINA. NEW CHAETOGNATH FAMILY, GENERA AND SPECIES FROM PACIFIC OCEAN 165 south-west Pacific Ocean during the cruis- two pairs of rows. Neck glands absent. Eggs es of Research Vessel (RV) “Academic Nes- released into water, non-adhesive to sub- meyanov”, using a Jeddy net, in April 1990. strate, marsupial sacs absent. Holotypes and paratypes are deposited in Two orders. the collection of the Pacific Institute of Oceanography, Far East Division, Russian Aphragmophora Tokioka, 1965 Academy of Sciences, Vladivostok. Transverse muscles absent; lateral body TAXONOMIC PART bending lacking. Epi-, meso-, and bathypelagic forms. Epipelagic forms closely ap- EUKROHNIONES new subclass proaching coast for spawning.

One pair of SGS on lateral fields of body Biphragmosagittiformes new order or one pair of fins. Internal ring of corona ciliata (sometimes absent in bathypelagic Transverse muscles present; body bend- species) without secretory cells. Eyes of ing in lateral directions. Small benthic everted type. Alveolar tissue commonly forms, commonly on shelf. single-layer, containing of flat rhomboid One family. cells. Teeth in two or one pairs of rows. Neck glands present. Eggs released into wa- BIPHRAGMOSAGITTIDAE fam. nov. ter, not adhesive to substrate, commonly in marsupial sacs hanging from opening of ovi- Type genus: Biphragmosagitta gen. nov. ducts and carrying by individual. Diagnosis. Trunk and tail sections with transverse muscle (phragmes; apparently, SPADELLIONES new subclass supercontraction). Two pairs of lateral fins. Sac-like gelatinous structures at bases of One pair of SGS on lateral fields of body lateral fins or elsewhere absent. Two pairs of or one pair of fins. Pair of soft tentacles on rows of teeth on head. sides of head in some species. Internal ring Two genera. of corona ciliata without secretory cells in most species, and special adhesive organs present at tail section. Corona ciliate and Biphragmosagitta gen. nov. eye reduced in some bathypelagic species. Type species: Biphragmosagitta taraso- Eyes of inverted type. Alveolar tissue multi- vi sp. nov. layer, containing of large vesicular cells. Diagnosis. Corona ciliata located below Teeth in two pairs of rows. Neck glands eye level, sometimes on neck. Most part of absent. In Spadellidae Tokioka 1965, sper- posterior fins located on trunk in adult ma- matophore released behind corona ciliate. ture animals. Thick rays in fins present. No Eggs released into water, commonly adhesive to substrate. gut diverticula. Eyes present. Inner nuclear eye zone smaller than peripheric one. Hooks not serrated. SAGITTIONES new subclass Two species. Two pairs of SGS on lateral fields of body Comparative remarks. The new genus or two pairs of fins. Internal ring of corona differs from the other genera, Biphragmo- ciliata (sometimes absent in bathypelagic fastigata gen. nov., of the family Biphrag- species) without secretory cells. Eyes of mosagittidae in having the corona ciliata inverted type. Alveolar tissue multi-layer, located below the eye level, sometimes on containing of large vesicular cells. Teeth in the neck.

Biphragmosagitta tarasovi sp. nov. (Figs 1a–b, 2a–e, 4)

Holotype. Inventory No. PHRAG-1/1, south-west Pacific, 16°39’S 168°28’E. RV “Aca- demic Nesmeyanov”, station 1, sample 1 horizon 100–0 m over depth 1020 m, plankton sample using Jeddy net, 4 April 1990; Pacific Institute of Oceanography, Far East Division, Russian Acad- emy of Sciences, Vladivostok. Paratypes. Ten specimens from the same sample as holotype in the same collection. Description (holotype). Transverse muscle (phragmes; apparently, supercontraction) in trunk being 13% of entire trunk length and, in tail, 4.8% of entire tail section length. Body rigid and muscular. Head 3 times as broad as neck. Trunk narrow; no considerable narrowing in region of seminal receptacles. Trunk region non-transparent, dark brown. Terminal end of body sharp. Body length 6.5 mm. Head region long, 14% of entire body length. Tail region 22.5%, ventral ganglion 6.6% of entire body length. Protuberances around mouth without papillae. Gut without diverticula. One pair of rows of hooks and two pairs of rows of teeth on head. Hooks (7 or 8 on each side) without unguiculi and not serrated. Front teeth (7 or 8 on each side) apically located. Posterior teeth (10 or 11 on each side) located on protuberances behind front teeth. All teeth of conical form as in Sagitta. Anterior end of anterior lateral fins located in front of ventral ganglion posterior level. Part of fin situated at level of ventral ganglion 0.7 times as long as ganglion, 4.7% of body length. Length of anterior fins 27% of body length, about 1.2 times as long as Fig. 1. Biphragmosagitta tarasovi sp. nov. (holo- posterior fins. Rays of anterior lateral fins type: specimen at 2nd maturity stage); habitus short. Borders, anterior and posterior ends (a), lateral view (b). Scale bar: 1 mm. of anterior lateral fins lacking rays. Dis- tance between anterior end of posterior fins and posterior end of anterior fins short, end of tail fin concave (lyre-shaped) as in being 2.3% of body length. Length of pos- Pseudosagitta lyra (Krohn, 1853). Some of terior fins 22.9% of body length; trunk part short rays present on tail part of posterior of posterior fins is 1.3 times larger than tail fins. Remaining part of posterior fins with- part. Some short rays located on tail part out rays. Ovaries short, about 8% of body of posterior fins. Tail fin about 5% of body length, containing numerous immature length and 22.4% of tail length. Posterior eggs set in two rows.

Fig. 2. Biphragmosagitta tarasovi sp. nov. (a–e): tail region in paratype at 4th maturity stage, ventral view (a), tail region in paratype at 1st maturity stage, ventral view (b), head region in holotype, ventral view (c), eye in holotype (d); hook in holotype (e); Biphragmosagitta angusticephala sp. nov. holotype (f–h): eye (f), head region, ventral view (i); Biphragmofastigata fastigata sp. nov. holotype: eye (g), head region, ventral view (h). Scale bar: 0.7 mm (a), 0.25 mm (b), 0.3 mm (c), 0.04 mm (d), 1 mm (e), 0.03 mm (f), 0.02 (g), 0.45 (h), 0.14 (i).

The specimen is at the 2nd maturity stage. It has a rounded corona ciliata located below the eye level, with its most part on the head, and a lesser part on the neck. The alveolar tissue is fringing neck, the ventral and dorsal body nearly to the middle tail region. The eyes are rounded, with a dark pigment zone in the center of the eye. There is a black crosslike structure inside the dark pigment zone. Remarks. The paratype at the 1st maturity stage are without defects. Specimens at the 4th maturity stage have traces of the lysis of the muscles. This defect shows some resemblance to the abnomaly of the chaetognathen species which was described from a volcanic bay (Kraternaya Bay, Ushishir Is- land) (Kassatkina, 1995). There are variations of the transverse muscle extension into the cavity of the tail section: a specimen at the 2nd maturity stage (holotype) has the smallest extension (13% of tail section length); a specimen at the 1st maturity stage has a longer extension (32.7% of tail section length); a specimen at the 4th maturity stage has the longest extension (45.3% of tail section length) (paratypes). This fact can probably be interpreted by some influ- ence of geophysical activity of this region in the south-west Pacific Ocean where the samples of this plankton chaetognath were collected. A specimen at the 4th maturity stage has elongate seminal vesicles contacting both pairs of lateral fins, but far from the tail fins. The seminal vesicles are very simple and differ from those in Sagittidae by the absence of any inner structure. Etymology. This species is named after V.G. Tarasov, a hydrobiologist, who organ- Fig. 3. Habitus: a, Biphragmofastigata fastigata ised the cruises of RV “Academic Nesmeya- sp. nov. (specimen at 2rd maturity stage); b, Bi- nov” to the south-west Pacific Ocean. phragmosagitta angusticephala sp. nov. (specimen at 4rd maturity stage). Scale bar: 1 mm. Biphragmosagitta angusticephala sp. nov. demic Nesmeyanov”, station 2, sample 2, hori- (Figs 2f–i, 3b) zon 300–200 m over depth of 390 m, plankton sample using Jeddy net, 5 April 1990; Pacific In- Holotype. Inventory No. PHRAG-2/1, stitute of Oceanography, Far East Division, Rus- south-west Pacific, 16°28’S 168°18’E. RV “Aca- sian Academy of Sciences, Vladivostok.

Paratypes. Five specimens from the same corona ciliata (elongated, without gyri) is sample as holotype in the same collection. located below the eye level, with its most Description (holotype). Transverse mus- part on the trunk, and the lesser part on the cle (phragma, apparently supercontraction) neck and head. Alveolar tissue fringes the in trunk 29.6% of entire trunk length, and terminal end of the body which is not sharp. transverse muscle in tail 16% of entire tail The eyes are rounded, with a dark pigment region length. Body rigid and muscular. zone in the center of the eye. Head narrow, as broad as trunk, but with Remarks. There are variations of the slightly broader neck. Trunk narrow; no transverse muscle extension into the cav- considerable narrowing in region of seminal ity of the tail section: a specimen at the 4th receptacles. Trunk region non-transparent, maturity stage (holotype) has the smallest dark brown. Terminal end of body not sharp. extension (16% of tail section length); a Body length 5 mm. Head section short, specimen at the 4th maturity stage (a para- 8% of entire body length. Tail section 24.8%, type) has the longest extension (30% of tail ventral ganglion 11% of entire body length. section length). Protuberances around mouth without pa- Comparison. This new species differs pillae. Gut without diverticula. One pair of from the other species, B. tarasovi sp. nov., rows of hooks, and two pairs of rows of teeth in particular, in the position of the anterior on head. Hooks (7 or 8 on each side) with- fins relative to the ventral ganglion. out unguiculi and not serrated. Front teeth Etymology. From Latin: angustus (nar- (3 on left side and 2 on right side) apically row) and cephalo (from Greek kephal, keph- located. Posterior teeth (16 or 17 on each alo) (head). side) located on protuberances behind front teeth. Anterior lateral fins without rays. Biphragmofastigata gen. nov. Anterior end of anterior lateral fins located behind posterior level of ventral ganglion: Type species: Biphragmofastigata fasti- distance 18 times shorter than length of gata sp. nov. ganglion (0.7% of body length). Diagnosis. Corona ciliata located at eye Length of anterior fins 22% of body level or higher. Larger part of posterior fins length, 1.2 times shorter than length of pos- located on trunk section in adult mature anterior fins. Distance between anterior end imals. Thick rays in fins absent. Gut diver- of posterior fins and posterior end of ante- ticula absent. Inner nuclear eye zone small- rior fins short, 4% of body length. Length of er than peripheric one. Hooks not serrated. posterior fins 26.7% of body length; trunk One species. part of posterior fins nearly 1.5 times larger Comparative remarks. The new genus than tail part. Rays of posterior fins short. differs from the other genus of the family Borders and anterior ends of posterior lat- Biphragmosagittidae, Biphragmosagitta eral fins lacking rays. Tail fin about 5% of gen. nov., in having the corona ciliata lo- body length and 20% of tail part. Posterior cated at the eye level or higher, and the ab- end of tail fin without concavity. Whole tail sence of thick rays in the fins. fin pierced with rays. Ovaries 15.8% of body length, containing numerous mature eggs Biphragmofastigata fastigata sp. nov. set in one row. (Figs 2g–h, 3a) The specimen is at the 4th maturity Holotype. Inventory No. PHRAG-3/1, 16°28´S stage. It has elongate seminal vesicles, con- 168°18´E, RV “Academic Nesmeyanov”, station 2, tacting both pairs of lateral fins, but far sample 2 horizon 300–200 m over depth of 390 m, from tail fins. The seminal vesicles are very plankton sample using Jeddy net, 5 April 1990; Pa- simple and differ from those in Sagittidae cific Institute of Oceanography, Far East Division, by the absence of any inner structure. The Russian Academy of Sciences, Vladivostok.

Fig. 4. Biphragmosagitta tarasovi sp. nov.: trunk region (ventral view), part with transverse muscles. Scale bar: 0.6 mm.

Paratypes. Four specimens from the same (8 on each side) located on protuberances sample as holotype, in the same collection. behind front teeth. All fins (anterior, poste- Description (holotype). Transverse rior, lateral, and tail ones) without rays. muscle in trunk section 13% of entire trunk Length of anterior fins 12.7% of body length, and transverse muscle in tail 5% of length, 2 times shorter than length of pos- entire tail section length. Body transpar- terior fins. Anterior end of anterior lateral ent, rigid and muscular. Terminal end of fins located behind posterior level of ventral body sharp. Trunk narrow; no considerable ganglion. This distance 1.5 times shorter narrowing in region of seminal recepta- than length of ganglion, and 6% of body cles. Head trapezoidal, wide, broader than length. Distance between anterior end of trunk. posterior fins and posterior end of anterior Body length 4.9 mm. Head region short, fins short, 6.6% of body length. Length of 5% of entire body length. Tail section 27.9%, posterior fins 26.4% of body length; trunk ventral ganglion 9% of entire body length. part of posterior fins nearly 1.2 times larg- Protuberances around mouth without pap- er than tail part. Length of trunk parts of illas. Gut without diverticula. One pair of posterior fins longer than anterior lateral rows of hooks and two pairs of rows of teeth fins. Tail fin about 3.5% of body length and on head. Hooks (6 on each side) without un- 12.7% of tail part. Ovaries 10.6% of body guiculi and not serrated. Front teeth (5 on length, containing numerous immature each side) apically located. Posterior teeth eggs set in two rows.

The specimen is at the 2nd maturity Comptes Rendus de l’Académie des Sciences stage. The corona ciliata is short, without de Paris, Série 3, 301(8): 397–399. gyri and located almost entirely on the Casanova, J.-P. 1986. Archeterokrohnia rubra n. head, just behind the brain. Alveolar tissue gen., n. sp., noveau Chaetognathe abyssal de l’Atlantigue nord-africain: description syste- fringes the neck and the trunk nearly to the matique, hypothes phylogenetique. Bulletin anterior lateral fins. Alveolar tissue is also du Musee d’Histoire Naturelle de Paris, Série present on the tail region between the pos- 4, 8A(1): 185–194. terior lateral fins and the tail fin. The eyes Casanova, J.-P. & Duvert, M. 2002. Compara- are rounded, with a dark pigment zone in tive studies and evolution of muscles in chae- the center of the eye. tognaths. Marine Biology, 141: 925–938. Remarks. There are variations of the Chen, J.-Y & Huang, D.-Y. 2002. A pos- degree of the transverse muscle extension sible Lower Cambrian chaetognath (arrow into the cavity of the tail section. A speci- worm). Science, 298: 187. Dallot, S. & Ibanes, F. 1972. Etude, prelimi- men at the 2nd maturity stage (holotype) naire de la morphologie et de l‘evolucion has the smallest extension (13% of tail sec- chez les Chaetognathes comparaison de trios tion length); a specimen at the 1st maturity techniques multivariable: classification au- stage (a paratypes) has the longerst exten- tomatique, classification cladistique et anal- sion (16% of tail section length). yse des coordonnees principales. Investiga- Etymology. From Latin: fastigata means cion Pesquera, 36(1): 31–41. having a sharp-pointed end. Doncaster, L. 1902. On the development of Sa- gitta with notes on the anatomy of the adult. Quarterly Journal of Microscopical Science, ACKNOWLEDGEMENTS 46: 351–398. The work was supported by the Far East- Elpat’evskiy, V.S. [Elpatiewsky, W.S.] 1910. ern Branch of Russian Academy of Sciences no. Die Entwiklungsgeschichte der Genital- 06-III-А-07-253. produkte bei Sagitta. I. Die Entwiklung der Eier. Biologicheskiy Zhurnal, 1(4): 1–10. Elpat’evskiy, V.S. [Elpatiewsky, W.S.] 1913. REFERENCES Origin of the ovum and embryological development of Sagitta. Part 1: Origin of the Beauchamp, P., de. 1960. Classe des cheto- ovum. Izvestiya Obschestva Lubiteley Estest- gnathes (Chaetognatha). In: Grassé, P. voznanya, Antropology, Etnography, 126(1): (Ed.) Traité de Zoologie, 5(2): 1500–20. Mas- 1–69. (In Russian). son, Paris. Elpat’evskiy, V.S. [Elpatiewsky, W.S.] 1914. Beraneck, E. 1895. Les Chaetognathes de la Origin of the ovum and embryological de- Baie d’Amboine. Revue Suisse de Zoologie, velopment of Sagitta. Part 2: Embryological 3(1): 1–137. development. Izvestye Obschestva Lubiteley Bieri, R. 1974. First record of the chaetognath Estestvoznanya, Antropology, Etnography, genus Krohnittella in the Pacific and descrip- 126(2): 1–48. (In Russian). tion of a new species. Wasmann Journal of Bi- Fedonkin, M.A. 2003. The origin of the Metazoa ology. 32: 297–301. in the light of the Proterozoic fossil record. Bowman, T. E. & Bieri, R. 1989. Paraspadella Paleontological Research, 7(1): 9–41. anops, new species, from Sagittarius cave Filatova, Z.A. 1951. Class Chaetognatha. In: Grand Bahama Island, the second troglobitic Zenkevich, L.A. (Ed.) Rukovodstvo po zo- chaetognath. Proceedings of the Biological ology [Manual in Zoology], 3(2): 592–608. Society of Washington, 102: 586–589. Sovetskaya nauka, Moscow. (In Russian). Bush, W. 1851. Beobachtungen über Anato- Fowler, G.H. 1906. The Сhaetognatha of the Si- mieund Entwiklung einiger wirbellosen Seeth- boga expedition. Siboga Exp. Monogr. Leiden. iere. A. Hirschwald, Berlin. 143 p. 21: 1–86. Casanova, J.-P. 1985. Description de l”appareil Ghirardelli, E. 1968. Some aspects of the biol- genital primitive du genere Heterokrohnia et ogy of the Сhaetognaths. Advances in Marine nouvelle classification des Chaetognathes. Biology, 6: 271–375.

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