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Gulf Research Reports Volume Issue 2 January 99 An Illustrated Key to the Chaetognatha of the Northern Gulf of Mexico with Notes on their Distribution Jerry A. McLelland Gulf Coast Research Laboratory DOI: 0.75/grr.002.07 Follow this and additional works at: http://aquila.usm.edu/gcr Part of the Marine Biology Commons Recommended Citation McLelland, J. A. 99. An Illustrated Key to the Chaetognatha of the Northern Gulf of Mexico with Notes on their Distribution. Gulf Research Reports (2): 45-72. Retrieved from http://aquila.usm.edu/gcr/vol/iss2/7 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.

Guy Research Reports, Vol., No. 2, 45-72, 99 Manuscript received August 5, 9: accepted September 2, 9 AN ILLUSTRATD KY TO TH CHATOGNATHA OF TH NORTHRN GULF OF MXICO WITH NOTS ON THIR DISTRIBUTION JRRY A. McLLLAND Invertebrate Zoology Section, Gulf Coast Research Laboratory, P.O. Box 7000, Ocean Springs, Mississippi 954-7000 ABSTRACT A key is provided to facilitate the identification of 24 species in nine genera of Chaetognatha occurring in the northem Gulf of Mexico. Included are the deep-water species, ukrohnia proboscidea,. calliops, Mesosagitta sibogae, and Sagitta megalophthalma, all recent additions to the known fauna of the region. Meristic data, brief descriptions, ecological notes, Gulf of Mexico records, and illustrations are also presented. INTRODUCTION Chaetognaths (arrow worms) have long been recognized as a significant trophic link between the vast numbers of marine copepods and larger predators, including many commercially impoaant species of fish (Heydom 959; Reeve 9; Nagasawa and Marumo 9). Certain species are associated with discrete physio-chemical conditions and have, thus, become useful as hydrological indicators in areas of intermixing water masses. Some species have gained recognition as vectors in the life cycles of marine parasites (Pearre 979; Boltovskoy 9; Jarling and Kapp 95). The chaetognath fauna of the Gulf of Mexico is known from various ecological studies concemed with the population structure in continental shelf and openocean plankton communities, but few of these works have included detailed descriptions of the animals or dealt with the entire reported fauna of the Gulf. However, descriptions of all species known to occur in the Gulf of Mexico were included in a recent work by Michel (94) on chaetognaths of the Caribbean Sea and adjacent areas. The present wok, limited in scope to the northem Gulf of Mexico, attempts to facilitate the identification of common species of chaetognaths one might expect to encounter in plankton samples from coastal and offshore waters of the region. The key also includes some rare species recently obtained from deep-water collections. Prior to 9, 2 species representing genera were recorded from the Gulf of Mexico. Of those, one is excluded from this key, the monospecific Bathybelos typhlops Owre, a rare, extremely deepdwelling species known only from a single specimen collected in the central Gulf (Owre 97). Recently, the deep-water species Mesosagitta sibogae Fowler, 90, ukrohnia calliops McLelland, 99,. proboscidea Fumestin and Ducret, 95, and Sagitta megalophthlma Dallot and Ducret, 99, were added to the Gulf fauna (McLelland and Perry 99), increasing the number of species to 25. At this writing, the fairly common epibenthic family Spadellidae has yet to be reported in the Gulf of Mexico; thus, only the planktonic species of Chaetognatha are considered in this key. There are, worldwide, currently about 25 described species of the phylum Chaetognatha belonging to two orders and six families of which the Sagittidae is the most diverse. In his 95 publication, Tokioka introduced a detailed phylogeny based on morphological similarities in which he arranged the 5 then recognized species into classes, orders, sub-orders, and families and restructured the former genus Sagitta into nine genera. Acceptance for Tokioka's proposal, especially the latter point, has been slow to gain support among specialists even though numbers of newly described species continue to mount. Recent systematic works such as those of Kassatkina (97, 92), Casanova (99, and Salvini-Plawen (9), have expanded upon Tokioka's original proposal. Accordingly, this work conforms to the revised taxonomy of the phylum, including the restructuring of the former genus Sagitta, in order to better account for differences among the distinct "groups" within the Sagittidae. MATRIALS AND MTHODS Chaetognath specimens (from 5-0% buffered formalin samples) were examined unstained under stereoscopic dissecting and compound microscopes. Using an ocular micrometer, total length measurements (excluding the caudal fin) were made and percentages of tail segment to total length (T%) were calculated. Ranges of hooks and teeth recorded in Table 45

4 MCLLLAND TABL Meristic values of chaetognaths from the northern Gulf of Mexico Total 95 Number Number Length Tail Number Anterior Posterior Number Source of Species (mm) Segment Hooks Teeth Teeth Specimens Specimens ukrohnia bathyantarctica.2 25. 2 -- 2. bathypelagica 5.-5.9 7.2-. 9.0-0..2. 24.-25. 25.-27. 22.-2.5 24.5-29.2 9-0 9-9 - 0 --._ -- -5 2-4 2-7 - 2 2 4 B, BB. calliops. fowler;. proboscidea.4 0.4-.7 2.-5.0.2-.4 2.5 5.2-5. 7.0 0.7-..-4.4 2.-2.5 0. 2.7 22.-.0 2.7-29. 24.-27.0 24.9.2-24. 9.0 20.0-20.9 22.4-24.0 2.-2.0 24.2-2- 2-2 9-0 2 0-2 - 0 -- -- -- -- -- -- 9-0 7-5 2-5-9 2 4-5 0-2 2 20-27 4 4 2 B B Krohnitta paccjica.5-5.0 5.-5..4-7.0 2.7-.4 29.-2. 2.-0.0-0 9 9-4 4 2 A D D K. subtilis 7.-9. 9.-0.4.2-2..9-4.2.4-7. 4.7-5.0.4-40..O-40.4 7 7-9 7-0- 0 0-2 9-2 -- -- 5 2 A,B B. A,B, Pterosagitta draco.7-4.9 5.-5.9.2-7.2 7.-9.5 7.-4.4 7.-4. 7 9-0 4-7 - 2-4 - 5 B.D B,D B,D Caecosagitta macrocephala.5 5.-.5 7.5-0.5.2 4. 9.7 2.7-7.7 27.7-7. 2.7.9 0 0-2 0-2 9- -7-0 -22 7-24 29 0 B BB B, Ferosagitta hispida.9-9.4..9.4-.5 2.2. 2.5-27.0 2.-27. 7- - 7 7-9 5-9 -9 0-0-4 2 5 2 A DP C FIaccisagitta enfrata.5-7.9 0. 4.0-7.2.-9.9 7.9-.0 7.9 5.4-.5 4.9-5. 9-9 -9-9 7 7-9 7-0 0-2- 2 5 4 D F D,F D,F F. hexaptera 9. 4.- 7.0 20.-2. 0.-2..4-44.0 4.-5.0 59.2 22..4-2.4 4.-22.9.7-22.0.7-..-9..2 9-0 7-9 -9-7 -7 4- -4-2- -4 4 4 - -9 2- - 4-5 4 4 2 A A,D. A,DB A,D, A,

A KY TO TH CHATOGNATHA OF TH NORTHRN GULF OF MXICO 47 TABL (Continued) Meristic values of chaetognaths from the northern Gulf of Mexico Total % Number Number Length Tail Number Anterior Posterior Number Source of Species (mm) Segment Hooks Teeth Teeth Specimens Specimens F. lyra 2.5-4. 20. 22.7-25.7 2.0-29.0 0.-4.7 7.7-20.0 5.0 4.9-7..-.0 5.0-5.5 7-9 -9-7 - 5- -7-7- -9 0 2-0 2-0 - 0 Mesosagitta decipiens.2-0..0-2. 2.9-. 22.-2.2 9.4-2. 2.2-24.4-7 7-9 7-9 -9-7 4-20 9 M. minima. 5.0-5..-. 22.2.5-20.5 5.7-9. 7 7-7 2-5 -4 4-5 7-2 9- M. sibogae 0.-. 2.4-.7 4.0-4.2 4.-4.9. 9.4-27.4 7.5-22.9 20.0-2. 2.0-2.9 2. -9 7-9 7-9 -9 9 4-7 5- -7 7 9-20 7 Sagitta bipunctata 4..5-9. 0.0-. 2.-.0 27. 24.-2.4 24.-2.0 25.-2.4 0 9-9- 0 5 5-7 -7 5-9- 0-7 -5 0-5 5 4 2 S. helenae.4.-4. 5.5-5.9 7.2-9.2 2.5 2.4-27. 2.-25.4 2.2-24. 7-7 9-5-9 5-7 2-4 4-5 2 4 2 S. friderici.0-7.9.-. 2.4-.7 25.0-0.4 25.7-0.2 27.-29. 7-7- -9 5- - 7-9 -7 4-9 7-22 22 0 9 S. megalophtltalma. 7.5 0.0.9-2. 25.9 24.5 25. 22.5-2.9 7 7-7 5 7-7 2 9-2 2 S. tenuis Serratosagitta serratodentata Solidosagitta planctonis 4. 5.-5.9.0-. 5.-.4.9-7.9.-9.7 0.0-0..4.4-..0-.4. 24.-2.2 25.7-2. 22.9-2.4 2.9-2.0 2.0-24.5 25.-27.2 2.0 2.-29.9 2.5-24.4 7-7- - 7-5- -9 0-9 5-5-7 - -0 9-0 9-0 5-9-0 2-4 0-0-5-9 5-20 -20 0-4-5 7-20 4 4 2

4 MCLLLAND represent those observed in a single set from either side of the head. The meristic values in the table are based on observations of specimens ranging in size from juvenile to adult from the northem Gulf of Mexico collections listed below. This list is referred to by letter under the Source of Specimens column in the table. A. U.S. Bureau of Land Management (BLM- SUSIO), 974; northeastem Gulf of Mexico continental shelf and slope, MAFLA lease tracts (see McLelland 94); 2 stations, 0-25 m, 54 samples; collected by R.A. Woodmansee, Gulf Coast Research Laboratory (GCRL). B. U.S. Dept. of nergy (LBL-OTC), 97; Mobile OTC site (29ON, 0OW), 0-000 m, 0 samples; collected by J.P. Steen, Jr., GCRL. C. NOAA R/V Oregon, winter cruise, 975; Mississippi River delta region, surface samples; collected by J.P. Steen, Jr., GCRL. D. Louisiana Dept. of Wildlife and Fisheries (LWF), Louisiana Offshore Oil Port Study (LOOP), 9; over continental shelf south of Timbalier Bay, Louisiana; collected by R. Ganzcak, LWF.. MARFIN Geryon cruises - August and November, 97 and February, 9; northem Gulf of Mexico continental slope, 5 sites; 20077 m; collected by Harriet Peny, GCRL (See McLelland and Perry 99). F. Personal collections of J.A. McLelland from various locations around the Mississippi Sound barrier islands. Trunk Segment Anterior Teeth Rayed \ Posterior Teeth \ I \ Gut Diverticulae Posterior Fin Deep-water Collections. The OTC samples (see B above) were collected during four cruises made between June 5 and October, 97, under Subcontract LBL 47 402, Lawrence Berkeley Laboratory, for the U.S. Department of nergy Contract No. W- 7405-NG-4. Closing plankton nets (0.75 m, 202 pm mesh) equipped with Niskin double-trip devices were used. General Morphology The general features by which most chaetognaths are identified are depicted in Figure. Chaetognaths are bilaterally symmetrical enterocoelous coelomates characterized by three body segments - head, mnk, and tail - separated by transverse septae. AU chaetognaths display lateral and caudal stabilizing fins, a head armed with chitinous grasping spines (hooks), transparent to semi-opaque bodies, and a lack of excretory and circulatory systems. The nervous system features a primary cerebral ganglion in the head connected to two dorsal eyes, and one large trunk ganglion on the ventral surface. The mouth and anus, both ventral, are connected by a straight digestive tube. As protandric Tail Segment Figure. General diagram of a chaetognath showing diagnostic characteristics. hermaphrodites, chaetognaths feature two oblong ovaries in the trunk segment opening dorso-laterally at the tail-trunk septum and two testes in the tail segment which are connected by minute sperm ducts to exterior seminal vesicles. This key is primarily intended for the identification of well-preserved adult animals and is based on easily observed extemal features and some intemal features such as mature ovaries. Therefore, caution must be taken when juveniles or sub-adults are examined. It is advisable for one to first become familiar with the

A I(Y TO TH CHATIWNATHA OF TH NORTHRN GULP OF MXICO 49 adult forms in a given sample before attempts are made to identify immature specimens. Illustrations for this work were prepared using preserved specimens at hand. Because quality specimens of all species were not available, some whole-animal illustrations were omitted. The reader is referred to Michel (94) and Alvariao (99) for additional illustrations, especially for the genus ukrohniu. GULF OF MXICO CHATOGNATHA species list Class Sagittoidea Claus and Grobben, 905 Order Phragmophora Tokioka, 95 Family ukrohniidae Tokioka, 95 ukrohniu Ritter-Zahony, 909. buthyunturcticu David, 95. buthypelugicu Alvariiio, 92. culliops McLelland, 99. fowleri Ritter-Zahony, 909. humuta (Mobius, 75). proboscideu Fumestin and Ducret, 99 Order Aphragmophora Tokioka, 95 Family Krohnittidae Tokioka, 95 Krohnittu Ritter-Zahony, 90 K. pacifica (Aida, 97) K. subtilis (Grassi, ) Family Pterosagittidae Tokioka, 95 Pterosugittu Costa, 9 P. druco (Krohn, 5) Family Sagittidae Claus and Grobben, 905 Buthybelos Owre, 97 B. typhlops Owre, 97 Cuecosugitru Tokioka, %5 C. mucrocephalu (Fowler, 905) Ferosugittu Kassatkina, 97 F. hispido (Conant, 95) Flaccisugittu Tokioka, %5 F. enfutu (Grassi, ) F. hexupteru (d'orbigny, 4) F. lyra (Krohn, 5) Mesosugittu Tokioka, 95 M. decipiens (Fowler, 905) M. minima (Grassi, ) M. sibogae (Fowler, 90) Sagitta Quoy and Gaimard, 27 S. bipunctatu Quoy and Gaimard, 27 S. helenue Ritter-Zahony, 90 S. friden'ci Ritter-Zahony, 9 S. megulophthalmu Dallot and Ducret, %9 S. tenuis Conant, 9 Serrutosugitta Tokioka and Pathansali, 9 S. serrutodentatu (Krohn, 5) Solidosugittu Tokioka, 95 S. plunctonis (Steinhaus, %) KY TO TH CHATOGNATHA OF TH NORTHRN GULF OF MXICO. a. One pairoflateral fins..... 2 b. Two pairs of lateral fins... 0 2. a. Lateral fins extending onto trunk segment; one set of teeth... b. Lateral fins limited to tail segment (Fig. 4); two sets of teeth... genus Pterosugittu... P. druco. a. Lateral fins extending to ventral ganglion... genus ukrohniu... 4 b. Lateral fins not extending to ventral ganglion... genus Krohnittu... 9 4. a. yes with pigment... 5 b. yes without pigment... 7 5. a. Apical gland-cell complex bilobate and protruding, causing head to appear pointed (Figs. 2B,): hook tips bent inward at 45-90" angles (Figs. 2D,G); transverse musculature extending past posterior edge of ventralganglion... b. Apical gland not prominent, a single lobed mass (Fig. B); hook tips straight (Fig. ), transverse musculature even with posterior edge of ganghon.... fowlen'

so MCLLLAND. a. ye pigment small, elongate or U shaped, in posterior region of eye (Fig. 2 0.... proboscideu b. ye pigment large, lunate, encompassing most of median portion of eye (Fig. 2C).... culliops 7. a. Number of hooks greater than, hook tips straight.... bathyunturcticu b. Number of hooks less than, hook tips bent inward.... a. Hooks stout, nearly straight; tail less than 25% of body length.... hamata b. Hooks slender, gently curved; tail usually greater than 25% of body length.... buthypelogicu 9. a. Outer margin of hooks obtusely angled (Fig. 5G); mature ovaries enlongate, may extend past edge of lateral fins (Fig. S)... K. pacifica b. Outer margin of hooks evenly rounded (Fig. 5D); mature ovaries compact, not extending past edge of lateral fins (Fig. 5A)... K. subtilis 0. a. Body flaccid, highly transparent; trunk musculature inconspicuous... genus Fluccisugittu (Fig. 7)..... b. Body rigid, translucent; trunk musculature prominent.... a. Anterior fins long, inserted close to ventral ganglion, connected to posterior fins by raised portion of body cuticle; caudal fin bilobate (Fig. 7D)... F. lyra b. Anterior fin short, rounded, well separated from ventral ganglion and posterior fins; caudal fin not bilobate... 2 2. a. Two to four elongate, thin anterior teeth which sometimes protrude anteriorly (Fig. 7C)...... F.Iiexoptera b. Four to eight short, wide, overlapping anterior teeth... F. enj7uta. a. Hooks finely serrate on inner margins as seen under loox magnification (Fig. IB)... genus Serrutosugitta... S. serratodentutu b. Hooks not serrate... 4 4. a. Collarette absent or indistinct... 5 b. Collarette hstinct... 5. a. yes without pigment; anterior fins entirely rayed, emerging at a point well separated from ventral ganglion (Fig. B)... genus Cuecosugitta... C. mucrocephula b. yes with pigment; anterior fins with anterior rayless zone, emerging at or near ventral ganglion... genus Mesosugittu.... a. Tail segment less than 22% of total body length mature ovaries compact, with three to five large ova (Fig. D)... M. minima b. Tail segment usually greater than 22% of total body length; mature ovaries elongate with numerous small ova... 7 7. a. Seminal vesicles located approximately equal distance from posterior fins and caudal fin (Fig. A); maximum body length at maturity less than 5 mm... M. decipiens

A KY TO TH CHAT~GNATHA OF TH NORTHRN GULF OF MXICO 5 I b. Seminal vesicles adjacent to caudal fin and well separated from posterior fins (Fig. B); maximum body length at maturity may reach 20 mm... M. sibogae. a. Gut diverticulae present*... 9 b. Gut diverticulae absent... genus Sagitta... 20 9. a. Posterior fins triangular with prominent rayless zone; walls of gut tube lined with large vacuolar cells (Fig. IOD)... genus Solidosagitta... S. planctonis b. Posterior fins rounded, completely rayed; no large vacuolar cells associated with gut tube (Fig. ).. genus Ferosagitta... F. ltispida 20. a. Posterior fins well separated from seminal vesicles... 2 b. Posterior fins contacting the seminal vesicles... 22 2. a. Anterior fins emerging posterior to ventral ganglion by a distance of about half the length of the ganglion; large vacuolar cells lining middle third of gut tube (Fig. 0A)... S. megalophthalma b. Anterior fins emerging level with posterior edge of ventral ganglion; no large vacuolar cells associated with gut tube (Fig. 9A)... S. bipunctata 22. a. Anterior teeth numerous (-), elongate, protruding outward in an overlapping, fan-shaped arrangement (Fig. 9D)... S. helenae b. Anterior teeth less numerous (-9), short, lying flat against head... 2 2. a. Mature ovaries with large ova which are few in number and arranged in one row (Fig. 9)...... S.tenuis b. Mature ovaries with small, numerous ova arranged in double rows (Fig. 9H)... S. friderici * Gut diverticulae are often undeveloped and thus difficult to see in immature specimens. UKROHNIA BATHYANTARCTICA David, 95 Body length at maturity unknown but probably greater than 20 mm. Hooks -4. Teeth usually -. T% 9-25. No eye pigment. No collarette. No gut diverticulae. Fins extendmg from anterior end of ventral ganglion to anterior third of tail segment. Mature ovaries short with small ova. Seminal vesicles ovoid, adjacent to lateral tins. Hooks gently curved with straight tips. Oceanic, bathyplanktonic, probably cosmopolitan. The most deep-living species of the genus in the Gulf of Mexico, occurring at depths to 200 m (Fagetti 9). Collected from great depths in the Antarctic and Caribbean Seas (David 95; Fagetti 9; Owre 972, 97). Listed as meso- to bathyplanktonic (>500-00 m) in the Florida Straits off Miami (Stepien 90). Gulf of Mexico records Open Ocean - Fagetti (9), Owre (97), McLelland and Perry (99). One specimen (.2 mm), an immature individual, was obtained from the bottom net sample (77 m) at Area 5 of the February, 9 Geryon cruise (Table ). It was undoubtedly a stray from deeper waters upwelled in the region. It had no denticulate hooks, a feature noted by Owre (97) for 4 smaller (7.5-2.0 mm) Caribbean specimens. Fagetti (9) failed to find denticulate hooks on any of her I specimens (.5-20.0 mm) from the Gulf of Mexico and Caribbean Sea.

52 MCLLLAND B L I C F I G A Figure 2. A-D, ukrohnia calliopr. -G,. proboscides (A) whde animal, ventral view; (B) head, dorsal view; (C) eyes; (D) hook with tip variations; () head, dorsal view; (F) left eye; (G) hook tip variations. Scales: A =.0 mm; B, = 0.S mm; C, D, F, G = 0. mm.

A KY TO TH CHATOONATHA OF TH NORTHRN GULF OF b c o 5 B D Figure. urrohnia fowled. (A) immature spedmen, ventral view; (B) head, dorsal view; (C) eyes; (D) adult hook; () hook tip variations; (F) juvenile hook. Scales: A, B =.0 mm; D = 0.5 mm; C, F = 0. mm; = 0.05 mm.

54 MCLLLAND UKROHNL4 BATHYPLAGICA Alvariiio, 92 vesicles ovoid-elongate, separate from caudal fin, anterior third overlapped by posterior edge of lateral fins. Hook tips sharply deflected inward (Fig. 2D). Body length at maturity 20-2 mm. Hooks 7-0. Teeth 7-22 (mature). T% 229. No eye pigment. No collarette. No gut diverticulae. Lateral fins extending from middle of ventral ganglion to posterior third of tail segment. Ovary tips have coiled appearance before becoming fully mature. Seminal vesicles oval, separated from lateral and caudal fins. Hook tips bent inward at 45-90' angles. Oceanic, meso- to bathyplanktonic, probably cosmopolitan; inhabits cold, deep waters from 200 to 400 m. Recorded from the North Pacific (Alvarifio 92) and Atlantic oceans (Ducret 95; Owre 97). Gulf of Mexico rccords Open Ocean - Owre (97), McLelland and Perry (99). Numerous specimens were found in the OTC (200-00 m) and in the Geryon (200-77 m) samples, but none were mature. They probably were strays from a deeper population offshore. The smallest specimen (5. mm) from the OTC samples did not have denticulate hooks; however, denticulations were present on the two ventral-most pairs of hooks of three specimens (7.2, 7.2, and 7. mm) from the Geryon samples. Denticulate hooks have been observed on small specimens of this species by Owre (97) and Nagasawa and Marumo (979). All specimens had the characteristic hook tip deflection (an abrupt bend of approximately 45 degrees). UKROHNIA CALLIOPS McLelland, 99 (Figs. 2A-D) Body length at maturity 5-22 mm. Hooks -. Teeth up to 2. T% 2-. yes large with dark, lunate pigment cups (Fig. 2C). Paired apical gland at anterior tip of head. Distinct neck. No collarette. No gut diverticulae. Lateral fins broadly rounded at posterior ends, tapering anteriorly, extending from anterior third of ventral ganglion to posterior third of tail segment; fin rays present only in outer-most posterior edge. Caudal fin large, broadly triangular. Mature ovaries short; ova large, round, in two rows. Seminal Figure 4. Pterosugittu druco. Whole animal, ventral view. Scale =.0 mm.

A &Y TO TH CHATO~NATHA OF TH NORTHRN GULF OF MUCO 55 Oceanic, mesoplanktonic, probably hypoplanktonic along the continental slope in the northem Gulf of Mexico. Gulf of Mexico records McLelland (99), McLelland and Perry (99). Remanks One hundred and eighty-three () specimens were collected at four sites from 200 to 77 m over the Northeast Gulf continental slope in August, 97, and February, 9 (MARFTN Geryon cruises). Tokyo Bay (Japan) specimens of a closely related species,. kitoui Kuroda, 9, on loan from Kazunori Kuroda, were compared to the Gulf of Mexico specimens and found to be substantially different (McLelland 99). No denticulate hooks were found on any specimens from either species. UKROHNIA FOWLRI Ritter-Zahony, 909 (Figs. A-F) Body length at maturity 4-40 mm. Hooks 9-. Teeth up 0. Head wide with conspicuous neck. ye pigment small, in center of eye (Fig. C). In well preserved specimens, thick band of epidermal tissue encircles body at ventral ganglion. Lateral fins with rays only in posterior third. Ovaries short; ova large, few in number, in two rows. Seminal vesicles ovoid, separated from caudal fin. Hooks gently curved with straight tips (Figs. D,), appearing reddish in preserved specimens. Three to four ventral-most pairs of hooks strongly denticulate in small specimens (Fig. F). Oceanic, meso- to bathyplanktonic, cosmopolitan; abundant at depths greater than 00 m (Ducret 95; Alvariao 99). piplaaktonic in polar seas, but submerged to colder waters in tropics (Fraser 952; Boltovskoy 9). Gulf of Mexico records Open Ocean - very (%), Fagetti (%), McLelland (99), Mcklland and Perry (99). UKROHNIA HAMATA (Mobius, 75) Body length at maturity 54 mm. Hooks -9. Teeth up to 25. T% 9-24. No eye pigment. Head small with distinct neck. No collarette. No gut diverticulae. Fins narrow, extending from middle of ventral ganghon to seminal vesicles, with rays only in posterior third. Mature ovaries with small ova arranged in four rows. Seminal vesicles ovoid, adjacent to lateral fins and apart from caudal fin. Tips of hooks bent inward at sharp angle (0-90"). Oceanic, cosmopolitan, epiplanktonic in cold-water regions, submerged to deeper depths toward the tropics (Ducret 95; Boltovskoy 9). Gdf of Mexico records Open ocean - Pierce (954). No specimens were found in northem Gulf collections. Pierce's specimens, collected at 495-749 m, were possibly. bathypelagica or. bathyantarctica, species unknown at the time of his work. UKROHNIA PUOBOSCIDA Fumestin and Ducret, 95 (Figs. 2-G) Length at maturity -0 mm. Hooks 0-. Teeth -25 (5-25 when mature). T% 20-. Apical gland bilobate. ye pigment light brown in color, elongate, situated in posterior region of eye (Fig. 2F). No collarette. No gut diverticulae. Lateral fins extending from posterior edge of ventral ganglion to midpoint of tail segment; rays only in posterior half and outer edge. Mature ovaries short; ova up to 4, in two rows. Mature seminal vesicles ovoid-elongate with anterior third overlapped by lateral fins; well separated from caudal fin. Hook tips bent sharply inward (Fig. 2G). Oceanic, meso- to bathyplanktonic in tropical and subtropical Atlantic. Type material f" 000 and 00 m off southwest African coast (Fumestin and Ducret 95). Four specimens reported from 79 to 2072 m in Caribbean Sea (Owre 97; Michel et al. 97).

5 MCLLLAND C F r Figure 5. A-D, Krohnirta subtifis. G, K. pacifica (A) whole animal, ventral view; (B) head, ventral view; (C) right eye; (D) hooks; () whole animal, ventral view; (F) right eye; (G) hooks. Scales: A, =.0 mm; B = 0.5 mm; D, G = 0. mm; C, F = 0.05 mm.

Gulf of Mexico Records A KY TO TH CHATOGNATHA OF TH NORTHRN GULF OF MXICO 57 McLelland (99), McLelland and Perry (99). The single specimen collected at Area 5 during the 9 Geryon cruise constitutes a new record for the Gulf of Mexico (see McLelland 99). At 0. mm, it is the smallest yet recorded for the species. Its position at 29"N, 04"W and depth of 77 m extends the range of the species vertically, northward, and westward where it is considered a stray from a Caribbean population. KROHNITTA PACIFICA (Aida, 97) (Figs. 5-G) Body length seldom longer than 7 mm. Hooks -0. Teeth -4. T% 2-2. ye pigment small, round (Fig. 5F). No collarette. No gut diverticulae. Lateral fins extending from seminal vesicles to a point midway between ventral ganglion and tmnk-tail septum; gently curved, narrow, with rayless zone comprising about half of fin area. Caudal fin spatulate, often continuous with lateral fins. Mature ovaries long, may reach ventral ganglion; ova round or cuboidal, in one row. Seminal vesicles small, oval, usually overlapped by lateral and caudal fins. Hooks flat, highly transparent, light brown in color, outer edges obtusely angled (Fig. 5G). A Semi-neritic, epiplanktonic, cosmopolitan in tropical and subtropical seas (Fumestin 9; Boltovskoy 9). Often associated with outer coastal regions of mixed water (Pierce and Was 92; Almeida-Prado 9; McLelland 97, 94). Gulf of Mexico ncords Coastal - Cuba: Sukz-Caabro (955). Florida: Ritter-Zahony (90), Pierce (95 I), Smith (955). Northeast Gulf: Mulkana and McIlwain (97), McLelland (97, 94). Texas: Pierce (92), Adelmann (97). Mexico: Vega-Rodriguez (94, 95), Mostajo (97). Open ocean - Kolesnikov (9), Alvarifio (99 - as K. mutabbii), Michel et al. (97). According to Owre (90), fitter-zahony's (90) specimens of K. subtilis were probably K. pacifica. Owre (97) considers K. mutabbii Alvariiio, 99 to be a synonym of K. pacifica. Figure. Caecosagitta macrocephala. (A) head, ventral view; (B) immature specimen, ventral view. Scales: =.0 mm.

5 MCLLLAND KROHNITTA SUBTILIS (Grassi, ) (Figs. 5A-D) Body length at maturity 2- mm. Hooks 7-9. Teeth 9-2. T% -40. ye pigment small, threepointed (Fig. SC). No collarette. No gut diverticulae. Fins semi-circular, delicate, almost totally rayless, extending from seminal vesicles to 2/ distance from septum to ventral ganglion. Ovaries short, not extending past edge of lateral fins; ova round or oval, in two rows. Seminal vesicles small, rounded, touching lateral and caudal fins. Hooks clear, highly transparent, cutlass shaped (Fig. 5D). Oceanic, lower epiplanktonic to mesoplanktonic, cosmopolitan in tropical to temperate waters (Alvariiio 95; Boltovskoy 9). Usually reported as mesoplanktonic (200-400 m) in tropical and subtropical seas (Fumestin 957; Owre 90; Mattlin 974). Not common in near-shore coastal waters. Guy of Mexico rccords Coastal - Cuba: Suh-ez-Caabro (9%). Horida: Maidana and Mostajo (90). Northeast Gulf McLelland (97, 94). McLelland and Perry (99). Texas: Pierce (92). Mexico: Vega-Rodriguez (94, 95), Mostajo (97). Open Ocean - Pierce (954), Kolesnikov (%), Kolesnikov and Alfonso (9), very (9), Fagetti ( 9). This species reaches a greater length at maturity than K. pacifica and is narrower, more transparent, and inhabits deeper strata. PTROSAGITTA DRACO Krohn, 5 (Figs. 4, 2H) Mature body length 7-0 mm. Hooks 7-0. Anterior teeth up to. Posterior teeth up to. T% 742. ye pigment small, elongate (Fig. 2H). Voluminous collarette extending onto tail segment, becoming continuous wjth lateral fins, possessing lateral tufts or bristles (seen on well preserved specimens). Gut diverticulae present. Tail segment constituting almost half of body length. Lateral fins entirely rayed, confined to tail segment. Ova large, in two rows; mature ovaries may extend full length of trunk cavity. Seminal vesicles elongate, contacting lateral fins and lying close to caudal fin. Oceanic, epiplanktonic, cosmopolitan in tropical and subtropical areas (Owre 90 Alvariiio %5). Apparently tolerates mixed water along continental shelf regions (Pierce 92; Pierce and Wass 92; Saint-Bon 9; McLelland 94). Gulf of Mexico records Coastal - Cuba: Suhz-Caabro (955). Florida: Alvariiio (99), Maidana and Mostajo (90). Northeast Gulf Alvariiio (99), McLelland (97, 94), McLelland and Perry (99). Texas: Pierce (92). Mexico: Vega-Rodriguez (94, 95). Open ocean - Central Gulf Pierce (954), Kolesnikov (9), Kolesnikov and Alfonso (9), very (9), Alvariiio (99), Owre (97), Michel et al. (97), Mostajo (97). CACOSA GITTA MACR 0CPHAL.A (Fowler, 905) (Figs. A,B) Body firm, opaque, with conspicuously large head. Body length at maturity 20-22 mm. Hooks 0-, reddish in color. Anterior teeth -0. Posterior teeth up to 0. T% 2-40. No eye pigment. No collarette. No gut diverticulae. Anterior fins short, completely rayed, separated from ventral ganglion by about half their length. Posterior fins longer and wider than anterior fins, with small rayless zone at anterior end; length equally divided over trunk and tail segment. Seminal vesicles oval, separated from posterior and caudal fins. Ovaries may reach ventral ganglion when mature, containing small ova in multiple rows. Oceanic, cosmopolitan, meso- to bathyplanktonic in all oceans (Alvariiio 95; Boltovskoy 9). Gulf of Mexico records Coastal - Northeast Gulf McLelland and Perry (99). Open ocean (as Sagitta macrocephala) - Pierce (954), very (9), Fagetti (9), Owre (97), Michel et al. (97). Most specimens have dark granular material in the

TH NORTHRN GULF OF MXICO 59 wall of the gut which may be red or orange pigmented in fresh specimens (Alvariiio 97). Several specimens, all immature, were examined from the OTC (200-000 m) and Geryon (200-77 m) samples. FROSAGZTTA HZSPZDA (Conant, 95) (Figs. Il, 2G) Body length at maturity -2 mm. Hooks -. Anterior teeth 5-9. Posterior teeth 0-4. T% 2-2. Head large, conspicuous. ye pigment square to beanshaped (Fig. 2G). Collarette extending /2 to 2/ distance from neck to ventral ganglion. Gut diverticulae present. Lateral fins entirely rayed anterior fins emerging at posterior edge of ventral ganglion; posterior fins extending to seminal vesicles. Mature ovaries may extend past midpoint of anterior fm; ova round, in two rows. Ripe seminal vesicles large, oval to rectangular, separated from caudal fin by half their length. Neritic, epiplanktonic, in inshore tropical-subtropical Atlantic waters (Alvariiio 95; Reeve 9). Indicates presence of inshore water when collected offshore (Owre 90; Pierce and Wass 92; Ducret 9; Michel et al. 97). Guy of Mexico records (As Sagitta hispida) Coastal - Cuba: SuArez-Caabro ( 955). Florida: Pierce (95), Smith (955), Hopkins (9), Kelly and Dragovich (9), Maidana and Mostajo Northeast Gulf Mulkana and Mcnwain McLelland (97, 94). Texas: Pierce Adelmann (97). Mexico: Vega-Rodriguez 95), SuArez-Caabro and Gomez-Aguirre Mostajo (97). Open Ocean - Kolesnikov (9), very Alvariiio (99). The genus Ferosagitta Kassatkina, 97 was created to separate the similar species F. ferox, F. hispida, and F. robusta from the former genus Parasagitta Tokioka, 95 (See Kassatkina 92). FLACCISAGITTA NFLATA (Grassi, ) (Figs. 7A, 2B) Body length at maturity -25 mm. Hooks -. Anterior teeth -. Posterior teeth up to. TI 4-. ye pigment star-shaped (Fig. 2B). No collarette. No gut diverticulae. Anterior fins short, narrow, well separated from ventral ganglion, rayed only on outer posterior edges. Posterior fins large, rounded, 2/ length on mnk, rayed only on outer posterior edges. Mature ovaries may reach anterior fins; ova rounded, varying in size, in 2- rows. Seminal vesicles small, rounded, contacting caudal fin. Mature testes with conspicuously compacted V shaped cluster of material at posterior end of tail segment. Oceanic and semi-neritic, cosmopolitan, epiplanktonic in tropical and temperate regions (Pierce 95, 95; Alvariiio 95; Boltovskoy 9). Little tolerance for low salinities or persistently high temperature (Pierce 95). Gulf of Mexico recordr {As Sagitta enflnta) Coastal - Cuba: Suhrez-Caabro (955). Florida: Pierce (95), Smith (-955), Hopkins (9), Kelly and Dragovich (9), Alvariiio (99), Maidana and Mostajo (90). Northeast Gulf Alvarifio (99), Mdkana and McIlwain (97), Perry and Christmas (97), Show (97), McLelland (97, 94). Texas: Pierce (92), Adelmann (97). Mexico: Vega- Rodriguez (94, 95), Suirez-Caabro and Gomez- Aguirre (95), Mostajo (97). Open Ocean - Pierce (954), Kolesnikov (9), Kolesnikov and Alfonso (9), very (9), Alvariiio (%9) Michel et ai. (97). This species is easily recognized tropical waters. the most commonly reported and chaetognath in tropical and sub- FLACCISAGITTA HXAPTRA (d orbigny, 4) (Figs. 7B,C; 2A) Body length at maturity -40 mm. Hooks -0. Anterior teeth 2- (usually 2-4). Posterior teeth - (usually -5). T% 7-20. ye pigment elongate, T- shaped (Fig. 2A). No collarette. No gut diverticulae. Anterior fins short, rounded, well separated from ventral ganglion; rays only on outer edge. Posterior fins large, rounded, 2/ length on uunk, with internal, anterior rayless zone. Mature ovaries reach ventral ganglion; ova small, in three rows. Seminal vesicles small, rounded, close to caudal fin.

0 MCLLLAND I I Figure 7. A, Flaecisagirtu enf4t4. B, C, F. Ircxapteru. D,, F. lyra. (A,B,D) whole animals, ventral views; (C) head, dorsal view; () head, ventral view. Scales =.0 mm.

A KY TO TH CHA~OGNATHA OF 'IH NORTHRN GULF OF MXICO I Figure. A, Mesosagitta decipiens. B, C, M. sibogae. D, M. minima. (A) posterior end of body, ventral view; (B, D) whole animal, ventral view; (C) head, dorsal view. Scales =.0 mm.

2 MCLLLAND Oceanic, cosmopolitan, epiplanktonic in tropical and temperate regions (Alvariiio 95). Associated with deeper epipelagic zones (00-500 m) in warm seas (Owre 90 David 9; Mattlin 974). Seldom occurs in shallow coastal waters. Gulf of Mexico records (As Sagitta hexaptera) Coastal - Cuba: Suhrez-Caabro ( 955). Florida: Alvdo (99), Maidana and Mostajo (90). Northeast Gulf Alvariiio (99), Mulkana and McIlwain (97), McLelland (97, 94), McLelland and Perry (99 - as F. hexaptera). Mexico: Vega- Rodriguez (94, %5), Mostajo (97). Open Ocean - Pierce (954), Kolesnikov (%), Kolesnikov and Alfonso (9), very (9), Alvariiio (99), Owre (97), Michel et al. (97). This species resembles F. enj7uta but is usually associated with higher salinity and deeper water. One of the largest chaetognaths, it may each 70 mm in cold waters (Alvarifio 97). Because of its large size, it has been implicated as a predator of larval fish (Alvariiio 90, Thuesen and Bieri 97). FLACCISAGITTA LYRA (Krohn, 5) (Figs. 7D, 2C) Body length at maturity 0- mm. Hooks -9. Anterior teeth -. Posterior teeth up to 0. T% 5-20. ye pigment elongate, Y-shaped (Fig. 2C). No collarette. No gut diverticulae. Anterior fins emerge near the posterior edge of the ventral ganglion, anterior 2/ of length rayless. Posterior fins triangular, with large laminate rayless zone. Lateral fins connected by raised portion of body cuticle, or "fin bridge". Caudal fin bilobate on posterior edge. Ovaries long; ova small, in four rows. Seminal vesicles small, oval, situated close to posterior fins. Hooks robust, transparent, with conspicuous flange at base. Oceanic cosmopolitan in tropical and temperate regions (Alvariiio 95). piplanktonic to mesoplanktonic (00-700 m) in open Ocean waters (Pierce 954; Owre 90; David 9). Gulf of Mexico records (As Sagitta lyra) Coastal - Texas: Pierce (92). Northeast Gulf McLelland (97, 94), McLelland and Perry (99 - as F. lyra). Open Ocean - Pierce (954), Kolesnikov and Alfonso (9), very (9), Owre (97), Michel et al. (97). As they mature, these animals tend to lose hooks and teeth. Fully mature specimens usually possess hooks (Fig. 7), -5 anterior teeth, and 2-4 posterior teeth. MSOSAGITTA DCIPINS (Fowler, 905) (Figs. A, 2) Body length at maturity 0-4 mm. Hooks -7. Anterior teeth 7-9. T% 9-2. Posterior teeth -20. ye pigment elongate, T-shaped (Fig. 2). No collarette. Gut diverticulae prominent. Anterior fins tapered, emerging level with posterior edge of ventral ganglion, with small rayless zone at anterior end. Posterior fins rounded, equally situated over trunk and tail segments, widest near posterior end, with small rayless zone at anterior end. Mature ovaries with large ova in one row, extending to anterior fins; anterior end of ovaries produced into a sharp point which continues as a fine fiber connected to the body wall (Fig. A). Seminal vesicles long and narrow, equidistant from posterior and caudal fins. Oceanic, cosmopolitan in tropical to temperate regions, lower epiplanktonic to deep mesoplanktonic (David 9; Alvariho 95; Fagetti 972; Pierrot- Bults 979). Associated with low levels of illumination and temperature (Owre %0). Gulf of Mexico records (As Sagitta decipkns) Coastal - Northeast Gulf McLelland (97, 94), McLelland and Perry (99 - as M. decipiens). Mexico: Mostajo (97). Open ocean - Kolesnikov (9), Kolesnikov and Alfonso (9), very (9), Alvariiio (99). Some of the above-mentioned records and ecological notes may be inaccurate because of confusion with M. sibogae, a very similar species (see remarks for M. sibogae).

A KY TO TII CHATOGNATHA OF TH NORTHRN GULF OF MXICO MSOSAGITTA MINIMA (Grassi, ) (Figs. D, 2D) Mature body length -7 mm, widest at trunk-tail septum. Hooks 7-. Anterior teeth -5. Posterior teeth up to 2. T% 5-22. ye pigment small, slightly elongate, T-shaped (Fig. 2D). Small head with distinct neck. No collarette. Gut diverticulae small. Lateral fins delicate with sparse rays on outer margins. Anterior fins slightly separated from ventral ganglion. Posterior fins with /4 length on trunk segment. Mature ovaries short, with -4 large ova in one row. Seminal vesicles oval, widely separated from posterior fins, contacting caudal fin. Oceanic to semi-neritic, cosmopolitan in tropical and temperate regions where mixing of coastal and oceanic water occurs (Alvariiio 95; Boltovskoy 9). Not a good hydrological indicator species because of its association with mixed water (Pierce 95; Pierce and Wass 92). piplanktonic to mesoplanktonic, shows affinity for cooler temperature (Alvarifio 95; Ducret 9). Gulf of Mexico records (As Sagitta minima) Coastal - Florida: Maidana and Mostajo (90). Northeast Gulf Alvariiio (99), McLelland (97, 94), McLelland and Perry (99 - as M. minima). Texas: Pierce (92). Mexico: Mostajo (97). Open Ocean - very (9), Alvdo (99), Michel et al. (97). MSOSAGITTA SIBOGe (Fowler, 90) (Figs. B,C; 2F) Body length at maturity 4-20 mm. Hooks. Anterior teeth 7-9. Posterior teeth 4-20. T% 9-27. ye pigment elongate, T-shaped (Fig. 2F). No collarette. Prominent gut diverticulae. Anterior fins tapered, emerging level with posterior edge of ventral ganglion, with small rayless zone at anterior end. Posterior fins rounded, equally situated over trunk and tail segments, widest near posterior end, with small rayless zone at anterior end. Ovaries long, may extend past anterior fin, rounded at distal end; ova uniform in size, in one row. Seminal vesicles situated near caudal fin and widely separated from lateral fins, with a clear, bulbous inflation at anterior end. Oceanic, mesoplanktonic in mid and lower Atlantic latitudes. Probably distributed shallower in the mesopelagic realm than M. decipiens; apparently does not migrate diumally (Pierrot-Bults 979, 92). Gulf of Mexico records McLelland and Perry (99). Body length at maturity and seminal vesicle position are the major differences between this species and M. decipiens. Pierrot-Bults (979) resurrected Sagitru sibogae after examining the lectotypes of S. sibogae and S. decipiens. Her detailed redescriptions of both species corrected the inadequacies of Fowler's (905, 90) original descriptions. Northeast Gulf specimens were compared to the lectotype of S. sibogue (ZMA 525-C, Siboga xp., sta. 4) on loan from the Zoological Museum of Amsterdam. SAGITTA BIPUNCTATA Quoy and Gaimard, 27 (Figs. 9A, 2) Body length at maturity 0-5 mm. Hooks -. Anterior teeth 5-7. Posterior teeth up to 7. T% 24-27. ye pigment slightly elongate (Fig. 2). Collarette prominent to level of ventral ganglion, often extending entire length of body. No gut diverticulae. Lateral fins entirely rayed; anterior fin slightly separated from ventral ganglion. Mature ovaries long, may reach level of ventral ganglion; ova round, in one or two rows. Seminal vesicles well separated from posterior fins; anterior portion with clear, bulbous structure which sometimes contains a circular, saw-toothed inset. Oceanic, cosmopolitan in temperate to tropical water (Alvariiio %5). piplanktonic to upper mesoplanktonic (Owre 90; Legad and Zoppi 9; Mattlin 974). An indicator of high salinity, oceanic water (Pierce 95; Grant 9). Gulf of Mexico records Coastal - Cuba: Surez-Caabro (955). Florida: Ritter-Zahony (90), Maidana and Mostajo (90). Northeast Gulf: Alvariiio (%9), Mulkana and McIlwain (97), McLelland (97, 94), McLelland and Pew (99).

4 MCLLLAND, D F H i B G Figure 9. A, Sagitta bipunctata. B, C, D, S. heknae., F, S. tenuis. G, H, S. fridcrici. (A, B,, G) whole animals, ventral view; (C) head, dorsal view; (D) anterior head, ventral view; (F, I) lateral view of ovaries. Scales =.0 mm.

A KY TO TH CHATOGNATHA OF TH NORTHRN GULF OF MXICO 5! B C F Figure 0. A-C, Sagitta megalophthalma. D-F, Solidosagitta planctonis. (A, D) whole animals, ventral View: (B, F) heads, dorsal view; (C) lefl eye; () right eye. Scales: A, D =.0 mm; B, F = 0.5 mm; C, = 0. mm.

MCLLLAND Open ocean - Pierce (954), Kolesnikov (%), Kolesnikov and Alfonso (9), very (9), Alvariiio (99), Owre (97), Michel et al. (97). The anterior teeth are similar in appearance and arrangement to those of S. helenae except they are less numerous and shorter. SAGITTA HLNA Ritter-Zahony, 90 (Figs. 9B-D, 2L) Body length at maturity 9- mm. Hooks 7-. Anterior teeth up to 9. Posterior teeth up to 5. T% 2-2. ye pigment square (Fig. 2L). Cdlarette extending about half way to ventral ganglion. No gut diverticulae. Lateral fins entirely rayed; anterior fins emerge at posterior edge of ventral ganglion; posterior fins contact seminal vesicles. Mature ovaries may extend past ventral ganglion; ova cylindrical, in two rows. Seminal vesicles oval when ripe, contacting posterior and caudal fins. Neritic, epiplanktonic, in Atlantic tropical-subtropical regions (Alvarifio 95; Boltovskoy 9). Often associated with higher salinity continental shelf waters (Pierce 95, 95). Gulf of Mexico rccords Coastal - Florida: Ritter-Zahony (90), Pierce (95), Smith (955), Hopkins (9), Kelly and Dragovich (9), Alvarifio (99), Maidana and Mostajo (90). Northeast Gulf: Alvarifio (%9), Show (97), Mulkana and Mcnwain (97), Perry and Christmas (97), McLelland (97, 94). Texas: Pierce (92), Adelmann (97). Mexico: Mostajo (97). Open Ocean - very (9), Alvarifio (99). SAGITTA FRIDRICI Ritter-Zahony, 9 (Figs.!GH, 2J) Body length at maturity e mm. Hooks 7-9. Anterior teeth 5-9. Posterior teeth up to 22. T% 25-0. ye pigment square (Fig. 25). Collarette short at neck. No gut diverticulae. Lateral fins entirely rayed anterior fin tapered, emerging level with posterior edge of ventral ganglion. Mature ovaries may reach mid-point of anterior fins; ova small, round, in two rows (Fig. 9H). Mature seminal vesicles contacting posterior and caudal fins, possessing anterolateral circular processes. Neritic, epiplanktonic, widely distributed in coastal waters of Atlantic and adjacent seas (Alvariilo 95; Michel et al. 97). Usually associated with lower salinity, near-shore waters but able to tolerate oceanic salinities (Fumestin 957; Fraser 9; McLelland 90). Gulf of Mexico records Coastal - Northeast Gulf: McLelland (97, 90, 94). Mexico: Suirez-Caabro and Gomez-Aguirre (99, Laguarda-Figueras (97). Tokioka (974) considers S. friderici and S. tenuis to be ecological forms of the Same species, with S. tenuis being the typical form. He attributes the larger size (up to 9 mm) of S. friderici and other "forms" of S. tenuis to be characteristic of slower growth rates of "matured individuals" in colder water, whereas the smaller size of individuals in warmer water results from energy being transferred away from body growth toward ovary development to achieve rapid maturation rates. Mature specimens of the two species from the Northeast Gulf, although similar, are readily separated by the methods presented in this key. Sampling e+ dence seems to indicate overlapping habitats, given the occurrence of both species not only at the same locations, but often in the same sample (McLelland 97, 90, 94). SAGITTA MGALOPHTHALMA Dallot and Ducret, %9 (Figs. OA-C) Length at maturity up to 9.5 mm. Hooks 7-. Anterior teeth 5-. Posterior teeth -2. T% 22-29. ye pigment distinctively large, rectangular (Fig. OC). Collarette well developed at neck. Gut thickened at anterior end, resembling small diverticulae. Large vacuoles along exterior of gut walls for about 2/ length of gut tube. Lateral fins completely rayed; anterior fins emerging posterior to ventral ganglion by a distance equal to /2 the length of the ganglion. Mature ovaries extending to anterior end of posterior fins; ova round, in thrae rows. Seminal vesicles ovoid, well separated from posterior fins.

A KY TO TH CHATOGNATHA OF TH NORTHRN GULF OF MXICO 7 Oceanic, mesoplanktonic; reported from the westem Mediterranean Sea (holotype from 00-00 m), the Tyrrhenian Sea, and Gulf of Guinea by Dallot and Ducret (99). A few specimens have been collected in the Caribbean Sea where it is a suspected indicator of North Atlantic water (Michel et al. 97) and in the Florida Straits (Stepien 90). Gulf of Mexico Records McLelland and Perry (99). The five specimens from a 200-500 m. Geryon Cruise sample represented a new record for the Gulf of Mexico. They were probably transported in the Loop Current from the Caribbean Sea. SAGITTA TNUIS Conant, 9 (Figs. 9,F 2K) Body length at maturity less than 7 mm. Hooks 7-. Anterior teeth 5-7. Posterior teeth 0-. T% 24-2. ye pigment square (Fig. 2K). Collarette small at neck. No gut diverticulae. Lateral fins entirely rayed anterior fin tapered, emerging level with posterior edge of ventral ganglion. Mature ovaries usually not reaching anterior fin; ova large, round, few in number, in one row (Fig. 9F). Mature seminal vesicles contacting posterior and caudal fins, possessing anterolateral circular processes. cdog, uryhaline, associated with neritic waters in tropical, subtropical, and temperate Atlantic regions; epiplanktonic (Alvarifio 95). Gulf of Mexieo records Coastal - Cuba: Sukz-Caabro (955). Florida: Pierce (95), Smith (955), Hopkins (9), Kelly and Dragovich (9), Alvarifio (99), Maidana and Mostajo (90). Northeast Gulf: Alvarifio (%9), Mulkana and McIlwain (97), Perry and Christmas (97), Mcklland (97, 90, 94). Texas: Pierce (92), Adelmann (%7). Mexico: Vega-Rodriguez (94, %5), Mostajo (97). R e d s Recorded occurrences of S. tenuis from Gulf of Mexico'and coastal Atlantic regions may include S. friderici owing to Pierce's (95) placing the two species in synonymy (McLelland 90). Juveniles and subadults of the two species cannot be confidently separated. SRRATOSAGITTA SRRATODh'TATA (fiohql5) (Figs. IA-D) Body length at maturity 0- mm. Hooks 5-. Anterior teeth -0, Posterior teeth 5-20 (mature). T% 2-2. ye pigment elongate, T-shaped (Fig. ld). Collarette vestigial, usually not distinct at neck. No gut diverticulae. Anterior fins emerging at posterior end of ventral ganglion, with small intemal rayless zone. Posterior fins "bell" or "guitar" shaped, with intemal rayless zone. Mature ovaries extending to anterior fins; ova round to cuboidal, in one row. Seminal vesicles contacting lateral fins, separated from caudal fin, with two anterolateral papillae, or "horns", when mature. Hooks finely serrate on inner margins with serrata being low in profile (Fig. lb). Oceanic, epiplanktonic, inhabits tropical and subtropical waters of Atlantic and adjacent seas (Alvariiio 95; Boltovskoy 9). Usually associated with high salinity but has been known to tolerate low salinity coastal water, especially in temperate regions (Pierce and Wass 92). Gdf of Mexico records (As Sagitta sematodentata) Coastal - Cuba: Suhz-Caabro (955). Florida: Ritter-Zabony (90), AlvariAo (99), Maidana and Mostajo (90). Noaheast Gulf Alvariiio (99), Mulkana and McIlwain (97), McLelland (97, 94), McLelland and Perry (99 - as Serrutosugitta serrutodentatu). Texas: Pierce (%2), Adelmann (97). Mexico: Vega-Rodriguez (94, 95), Mostajo (97). Open ocean - Pierce (954), Kolesnikov (9), Kolesdkov and. Alfonso (9), very (9), Alvarifio (%9), Michel et al. (97). According to Pierrot-Bults' (974) classification, the Gulf popularion belongs to the Atlantic subspecies Sagitta serrutodentata serrutodentata Krohn, 5 as opposed to the subspecies S. serrutodentata utlunricu Thompson, 947, an isolated population in the South Pacific. A compound microscope is usually required to observe hook serrata in smaller specimens.

MCL~LAND C D I Figure. A-D, Sematosagitta serratodentata., Ferosagitta hispida (A, ) whole animals, ventral view; (B) hook detail; (C) hook; (D) right eye pigment. Scales: A, =.0 mm; C = 0. mm; D = 0.0 mm.