<I>Teuthowenia</I> (Oegopsida)

BULLETIN OF MARINE SCIENCE, 36(1): 1-85, 1985

SYSTEMATICS, BIOLOGY AND BIOGEOGRAPHY OF THE CRANCHIID CEPHALOPOD GENUS TEUTHOWENIA (OEGOPSIDA)

Nancy A. Voss

ABSTRACT Teuthowenia is comprised of three discrete, closely related, allopatric species. Synonymies, definitions, diagnoses and keys to all developmental stages are presented, along with a review of the complex history of the genus and detailed illustrations. The discrete, ecologically distinct, distributional patterns of the three species reflect the influence ofa number of biological and physical factors. T. megalops (Prosch) is confined to the highly productive Atlantic subarctic and the highly productive areas of the North Atlantic temperate region. T. maculata (Leach) is restricted to the area of year-round, high productivity in the eastern tropical Atlantic. T. pellucida (Chun) is distributed circumglobally in the mixed and fringing waters of the Southern Subtropical Convergence. The species display similar patterns of ontogenetic descent from near-surface waters to midwater depths of about 1,000 m to in excess of 2,500 m where the animals mature, mate and spawn. Teuthowenia species have differentiated physiologically and developmentally as well as morphologically. Variations in the maturity-related morphological features among the species suggest differences in behavioral patterns for courtship and copulation. The genus displays a high rate of evolution in male and female secondary sexual characters. The relationships of Teuthowenia with Egea and Megalocranchia, which together comprise the monophyletic Megalocranchia group, and with the other taoniin genera are discussed.

The taxonomic confusion surrounding the cranchiids was resolved in part by the familial revision of N. Voss (1980). The phylogenetic history of the family was reconstructed by N. Voss and R. Voss (1983). This first ofa proposed series of individual generic revisions elucidates the specific differentiation that has occurred in the various genera, the inter- and intraspecific variations in growth and development that have resulted in confusion within and among taxa, and the vertical and geographic distributions of the individual species. Teuthowenia historically has been the most confused of all the genera of the Cranchiidae. I found the distribution of the genus to extend from the subarctic region of the North Atlantic to the circumglobal waters in the region of the Southern Subtropical Convergence. North-south disjunctions in the distributional pattern were revealed by analyses of the collections from the Cape Town-Denmark transect of the 1928-1930 Round-the-World cruise of the DANAand the Cape Town-Madeira transect of the 1971 cruise of the WALTHERHERWIG,together with the results from a series of stations made by the DISCOVERYat about 10° intervals in the eastern North Atlantic between 60° and lION (Clarke and Lu, 1974; 1975; Lu and Clarke, 1975a; 1975b). The defined populations represent three discrete, closely related, allopatric species.

HISTORICALREVIEW The history of the genus Teuthowenia as now recognized is complex. The first mention of a nominal species was by Leach in 1817 in the Zoological Miscellany, when he briefly characterized Cranchia maculata on the basis of a mantle of a larva collected off West Africa by John Cranch during the Tuckey Expedition to the river Zaire. The fOllowing year, Leach (1818) listed the species when he 2 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985 expanded the diagnosis of his new genus Cranchia. In 1847, Prosch created a new subgenus of Cranchia, Owenia, to contain his newly described and figured species mega lops, the larval type of which was collected off the Faeroe Islands. In the same work, maculata was again listed. A number of inaccuracies in Prosch's original description and figures of mega lops were pointed out by Morch in 1850. Six years later in a footnote in his paper on the cephalopod hectocotylus, Steenstrup (1856) briefly characterized a new species Leachia hyperborea collected high up in Baffin Bay. In his 1861 survey of the cephalopods in the collections of the Copenhagen museum, Steenstrup showed that the dorsal, pseudoarticulation of the head and mantle used by Prosch to distinguish his subgenus Owenia from Cranchia, which displays a dorsal fusion between head and mantle, was not present in the primary specimen described and figured by Prosch (1847, figs. 4-6). Steen- strup found that the error was due to the misidentification of additional larvae, one of which Prosch showed in figure 7, considered along with the primary specimen. The supplementary specimens, which lacked the dorsal nuchal fusion, were not identical to megalops but appeared to be young gonatids. In the same work, Steenstrup expanded the characterization of the 210-mm ML subadult type of his species hyperboreus, placed it along with Loligo pavo Lesueur in his newly proposed genus Taonius, and designated the older species pavo as the type-species. Cranchia maculata was regarded as an incompletely known species and was not listed. Later Steenstrup (1881) further clarified the Cranchia-Owenia-Gonatus confusion. About the same time, Tryon (1879) listed maculata and megalops under Cranchia and assigned hyperborea to Lamark's genus Loligopsis along with pavo which had, prior to Steenstrup (1861), been placed in Loligopsis by Orbigny in 1839. Verrill (1881) further complicated the picture by naming a new genus Des- moteuthis, on the basis of a 330-mm ML specimen from off New England that he mistakenly considered to be identical to Taonius hyperboreus (Steenstrup). Later in the same paper, a second species tenera based on two juveniles the largest measuring 116 mm ML, from off New England was added to the new genus. Subsequent to Rochebrune's (1884) transfer of hyperborea back to Lamark's Lo- ligopsis, Hoyle (1885) replaced Streenstrup's species in Taonius, and clarified the Desmoteuthis problem. Fortunately, Verrill's specimens were well described and illustrated, permitting Hoyle to recognize Verrill's error in identifying the 330- mm ML animal with hyperboreus, and to correctly identify the specimen as Tao- nius pavo. Consequently, Hoyle placed Desmoteuthis hyperboreus, Verrill in the synonymy of Taonius pavo and synonymized Desmoteuthis with Taonius. In the course of the study, Hoyle reexamined the type of hyperboreus and gave the first detailed description of the species. In addition, he tentatively placed Verrill's second species Desmoteuthis tenera in the synonymy of hyperboreus, an assign- ment fully supported by my examination of the two syntypes of tenera. The following year in the Challenger Report, Hoyle (1886) elaborated on the Taonius- Desmoteuthis problem. For the first time, Steenstrup's type of hyperboreus was figured; and additionally, after reexamining the type, Hoyle gave the first description of Cranchia maculata, previously known only from Leach's brief characterization given first in Latin (1817)- "sacco laevi pulcherrime nigro maculato; maculisovatis distantibus"-and then in English (1818). Pfeffer (1900) accepted Hoyle's placement of Desmoteuthis hyperborea, Verrill in the synonymy of Taonius pavo, and D. tenera in the synonymy ofSteenstrup's hyperborea, but unfortunately retained the genus Desmoteuthis for hyperborea. The nomenclatural web was further tangled by the introduction of his 1884 genus Megalocranchia which Pfeffer synonymized with Desmoteuthis. Though the type voss: REVISION OF TEUTHOWEN1A 3

of Pfeffer's type-species Megalocranchia maxima, a 39-mm MLjuvenile from off the Cape of Good Hope, is not extant, having been destroyed in World War II, the originally described, and later illustrated (Pfeffer, 1912) feature of two rows of suckers on the tentacular stalk, alone precludes membership in the genus now recognized as Teuthowenia. At the same time, Pfeffer raised Prosch's discarded subgenus Owenia to the generic level to contain megalops. Chun (1906) supported Pfeffer's concept of Desmoteuthis and Owenia but, in his 1910 monograph he proposed Teuthowenia for the preoccupied name Owenia to contain megalops, and added a new species antarctica to the genus based on a 13-mm ML larva from the Antarctic. Chun expressed uncertainty, however, as to the significance of the difference that he noted between mega lops and antarctica as to the number of rows of suckers on the tentacular stalks-four rows in the former species and two rows in his new species. This difference plus the presence of the digestive duct appendages only on the digestive gland in antarctica as originally described and figured, and confirmed by my examination of the type, rather than on the gland and digestive ducts as found in the larva and adult of Teuthowenia, suffice to show that the two species are not congeneric. In addition, Chun added a new species pellucida to Desmoteuthis, describing and figuring in detail the 77-mm ML larval type collected in the Benguela Current; and, with reservations, mistakenly placed Helicocranchia pfefferi Massy, 1907 into the synonymy of the genus. The previous year, Hoyle (1909) had similarly accepted Pfeffer's retention of Desmoteuthis for hyperboreus, a move in conflict with his correct conclusions previously expressed in his works of 1885 and 1886, but Hoyle differed from both Pfeffer and Chun by retaining megalops in Cranchia. Hoyle (1910) continued to so regard both hyperboreus and megalops. Again, seeking to solve the Taonius-Desmoteuthis problem, Berry (1912) came to the same logical conclusions arrived at by Hoyle in his earlier works (1885; 1886), that is, "that Desmoteuthis must be regarded as absolutely synonymous with Taonius." Berry then proceeded to use Pfeffer's genus Megalocranchia, which Pfeffer himself had earlier (1900) synonymized with Desmoteuthis, to contain hyperborea as well as Chun's new species pellucida and the unlike forms of Pfeffer's maxima (type-species of Megalocranchia), Goodrich's abyssicola and Berry's own fisheri. Berry considered Chun's move to include Helicocranchia pfefferi in the group as doubtful. In the same year in his monographic review of the oegopsid cephalopods, Pfeffer (1912) likewise resurrected Megalocranchia to hold the same species as appeared in Berry's paper, with the exception of hyperborea, for which he persisted in retaining Desmoteuthis. Pfeffer expanded Teuthowenia to include four subgenera: Teuthowenia sensu stricto containing megalops and Chun's antarctica, Hensenio- teuthis, Sandalops, and Helicocranchia. In 1916 Berry described and figured a new species pardus from the Kermadec Islands. Considering the 50-mm ML larval type "exceedingly close" to pellucida, but differing in the degree of development of the stalked eyes, funnel and dentic- ulation of the sucker rings, Berry added pardus to Megalocranchia. In a footnote Berry proposed a new genus Verrilliteuthis for Desmoteuthis tenera Verrill which, for undisclosed reasons, he removed from the synonymy of hyperborea where it had reposed since Hoyle, 1885. Subsequently, Grimpe (1922) in his confused classification listed hyperborea under Verrilliteuthis. Misidentifications of Chun (1913), and Grieg (1920) led Grimpe (1922; 1925), and later Clarke (1966), to list pellucida for European waters. Robson (1924a) had difficulty in placing his new species impennis in Pfeffer's classification, so proposed a new genus Anomalocranchia. Later the same year, 4 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

Robson (1924b) figured the 57-mm ML type from off Cape Town and expanded his description to include internal features, The fact that past records were comprised of single or but a few specimens and the workers had little or no comparative material, coupled with the confusion that existed in the literature, suffice to explain why little or no attempt was made to connect any of the growth stages represented by the different nominal species. It was the relatively large collection of cephalopods made by the 1908-1910 Danish Oceanographical Expeditions to the Mediterranean and Adjacent Seas that afforded Degner the opportunity to do the first developmental work in the group. Studying 63 specimens from the northeastern Atlantic that he identified to Desmoteuthis hyperborea, Degner (1925) described some of the morphological changes that occur during growth from about 6 to 84 mm ML, and figured the developmental sequence of the fins and the posterior end of the gladius, and of the ocular photophores. Degner lacked, however, the intermediate size stages that might have permitted him to connect the young with the 315-mm ML (380-mm GL), maturing female that he, later in the same paper, described as a new species Desmoteuthis thori. His description of thori contained the first known reference to the brachial end-organ, which is now known to occur on the ends of all the arms in females of the genus Teuthowenia to which this animal belongs (N. Voss, 1980). Due to the lack of comparative male specimens, Degner did not recognize the modification as a female character. Additionally, the glandular processes present near the base of midarm suckers in maturing females of the species were described and figured for the first time. Teuthowenia was always shrouded in doubt, characterized only by larval features and confused from its conception by Chun's (1910) addition of the unlike form antarctica which introduced into the generic concept, among other things, the mistaken idea of two or four rows of suckers on the tentacular stalk. It is no wonder that new species based on misidentified larvae were subsequently added to the genus-T. (Ascoteuthis) corona Berry, 1920 from west of Bermuda, and T. tagoi Sasaki, 1929 and T. elongata Sasaki, 1929 from Japanese waters. In my examination of two of the three types, I found that Berry's corona belongs to Megalocranchia, and that Sasaki's elongata, by vitrue of its two rows of carpal suckers and the confinement of the digestive duct appendages to the digestive glands, is excluded from Teuthowenia. The exact identification of this latter type must await further study. The type of tagoi is not extant, and, finding the original description and figures insufficient for identification, I consider it to be a species dubia. Thiele (1934) incorporated Berry's new Ascoteuthis as a subgenus, along with Teuthowenia, Hensenioteuthis sensu stricto, and He/icocranchia into the reinstated genus Hensenioteuthis Pfeffer, 1900. At the same time, Thiele recognized Verrilliteuthis for hyperborea and Anomalocranchia for impennis. Over the years scattered, additional records of the various nominal species appeared in the literature including Verrill, 1884; Hoyle, 1886; 1906; Massy, 1909; 1913; Pfeffer, 1912; Chun, 1913; Joubin, 1920; 1924; 1933; Bouxin and Legendre, 1936; and G. Voss, 1955; 1960. Of these, I have examined Joubin's specimens and found his 1920 material, identified as Teuthowenia megalops, to be He/icocranchia sp., and two that I located of the three 1924 specimens, identified as Megalocranchia hyperborea, to be correctly identified to that nominal species. The specimen figured and described by Joubin (1933) as a male of Teu- thowenia mega lops was located in the Paris Museum. It is an adult female of Helicocranchia papillata, and was refigured by N. Voss (1980: fig. 4a). Though the two larval and juvenile specimens from off Cuba referred to Verrilliteuthis hyperborea by G. Voss (1955) are in poor condition, having dried out and been voss: REVISION OF TEUTHOWENIA 5 partially restored, they can be unhesitatingly reassigned to Megalocranchia on the basis of the photophores on the digestive gland, and other features, that I found on reexamination. The 15 young of Megalocranchia megalops reported by G. Voss (1960) from Bermudan waters proved on reexamination to be Helicocranchia papillata(Nos. 78317, 78319, 78320(2), 78321, 78323, 78324),? H.pfefferi(Nos. 78320 (1), 78322, 78325), Megalocranchia sp. (No. 78318), and? identification (No. 78338). Finally, in 1956 Muus, studying the cephalopod collections from the North Atlantic made by the DANAduring 1947-1953, linked the larval stage represented by megalops with the later growth stages represented by hyperborea and thori into a single species to which Muus assigned the oldest specific name megalops but, unfortunately, chose Desmoteuthis for the genus, despite the fact that it had previously (Hoyle, 1885; 1886; Berry, 1912) been clearly shown to be an absolute synonym of Taonius. Muus then proceeded, apparently without examining the types or specimens involved, to synonymize with mega lops such diverse forms as Helicocranchia pfefferi Massy, 1907, Teuthowenia antarctica Chun, 1910 and Desmoteuthis pellucida Chun, 1910, among others. His mistaken consideration of the latter two species led Muus to believe that megalops was bipolar or cos- mopolitan in distribution. G. Voss (1960) was rightly critical of Muus' synonymy, but subsequently, influenced by Berry's (1912) incorrect conclusion that Megalocranchia was the valid genus to contain hyperborea, used that genus for megalops. The recurrence of specimens of Helicocranchia in the literature of megalops (Joubin, 1920; 1933; Bouxin and Legendre, 1936; Muus, 1956; 1962; and Voss, 1960) dates back to Chun's (1910) error in assigning Helicocranchia pfefferi to the synonymy of Teu- thowenia. The controversy continued when Adam (1962), critical of the decisions of both Muus and G. Voss, recognized Berry's illy conceived Verrilliteuthis as the valid generic name for his well-described 68-mm ML specimen from off Angola specifically identified as hyperborea. This reported capture appears to be the first for a specimen to be subsequently identified (present paper) to maculata since that species was proposed by Leach in 1817. The same year Muus (1962), unaware of Adam's paper, addressed the criticisms and taxonomic decisions of G. Voss (1960) regarding megalops and reaffirmed his own decisions of 1956. In the meantime, Dell (1959) added a new species Megalocranchia richardsoni to the group on the basis of three juveniles and one subaduIt taken in New Zealand waters. Though recognizing the relationship to Berry's pardus, Dell could not reconcile the morphological differences between his specimens, which were in the sessile-eye stage and pardus, which was described and known only in the stalked- eye stage. Since its description in 1916, two additional single larval specimens of pardus from offN.S.W., Australia and Wellington, New Zealand had been reported by Allan (1945) and Dell (1952). Muus' choice of Desmoteuthis for the generic name of megalops was also rejected by Clarke (1962a) who attempted to reexpand Taonius to Steenstrup's original scope to include megalops along with Lesueur's generically distinct species pavo. In his 1966 work, Clarke recognized seven of the nine known nominal species and assigned them to four different genera. He left pardus in Megalocranchia, impennis in Anomalocranchia, and transferred pellucida, thori and richardsoni to Taonius to join megalops which he had transferred in his previous paper. Verrill's tenera appeared in the synonymy of megalops, while Steenstrup's hyperboreus appeared both as a junior synonym of megalops and as a species in Verrilliteuthis. The last taxon to be named to the group was Megalocranchia megalops australis, which G. Voss (1967) proposed for four larvae, 24-89 mm ML, taken off South 6 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. ], 1985

Africa, which differed from mega lops by the multi pointed tubercle on the funnel- mantle cartilages, first described here and now known to be characteristic of the southern ocean species. Additional differences were found in the structure of the beaks and radula. Comparisons between m. australis and the other southern ocean nominal species were not given. Because of the confusion concerning the various species in the group, Voss chose to identify his specimens as a subspecies of megalops rather than as a separate species. Monod (1970) made a major contribution to the meager history of Leach's maculata by publishing the principal portions of the heretofore unpublished journal of John Cranch from the 1816 expedition to the Congo during which maculata was collected. The text and expedition track were given, in addition to Cranch's original aquarelles, thus providing, for the first time, the time and place of capture as well as a figure of the poorly known type. In his effort to put taxonomic order into the cranchiids, Nesis (1974a) attempted to solve the Taonius-Desmoteuthis-Megalocranchia problem by rejecting generic names based on larvae. He subsequently used Berry's Verrilliteuthis to contain two species megalops and Dell's richardsoni; and considered megalops to consist of two subspecies m. megalops from the North Atlantic and G. Voss' m. australis from the South Atlantic and southwestern Indian Ocean. Then, working only with the literature and but a single 100-mm ML specimen that he misidentified to Chun's pellucida, Nesis proposed a new genus Vossoteuthis to hold pellucida and Berry's pardus. N. Voss (1980) examined Nesis' specimen and found it to connect into a growth series identified to Liguriella podophtalma IsseI, 1908. In addition, Nesis placed Robson's Anomalocranchia impennis in the synonymy of Sandalops Chun, 1906. In a study of the distribution of the oceanic cephalopods of the southwestern Atlantic, Nesis (1974b) employed some of the same specimens collected by Russian research ships used in the above systematic work to dem- onstrate the association of megalops australis with the Southern Subtropical Con- vergence. In a series of papers on squid larvae that appeared during this period, Okutani and McGowan (1969) and Okutani (1969; 1974) misidentified a number of specimens (R. Young, 1972) to Teuthowenia megalops or Teuthowenia sp., thereby introducing the misconception that the normal distribution of the group extends into the North Pacific. Following Clarke's (1962a) transfer of megalops to Taonius, subsequent English workers primarily used that nominal combination for the species. Specimens identified to Taonius megalops and Verrilliteuthis hyperborea were included in the studies of Denton et al. (1958; 1969) on buoyancy mechanisms in squids. Dilly (1972) recorded observations on the "balling-up" behavior exhibited by a megalops larva when disturbed. Dilly's figures of the live animal show the characteristic, relatively widely-spaced chromatophore pattern on the mantle. In a series of papers on the vertical distribution of oceanic cephalopods in the northeastern Atlantic using open-closing nets at six stations made at about 10°intervals from 60° to lION, between 20° and 25°W, Clarke and Lu (1974; 1975) and Lu and Clarke (1975a; 1975b) demonstrated the occurrence of ontogenetic descent in the group, and revealed a break in the north-south distribution of what they considered to be a single species mega lops. Additionally, they found mega lops to be the commonest and second commonest squid taken at 53° and 600N, and noted a difference in the ontogenetic distribution in the water column between the northern (600-53°N) and southern (lIoN) populations. I have examined the specimens from the southernmost station, and they are included in the present paper under maculata, The specimens from the two northern stations were used by voss: REVISION OF TEUTHOWENIA 7

Dilly and Nixon (1976) and Nixon (1983) in their studies of certain aspects of morphology and life history of megalops. The former work included the histology of the ocular photophores and scanning electron micrographs showing details of the suckers, funnel organ and radula. Specimens identified to megalops were used in several studies on the structure of the statoliths (Dilly, 1976), the statocysts and photosensitive vesicles (J. Young, 1977; 1984) and the suckers (Nixon and Dilly, 1977) of cephalopods. The number of captures subsequently to be identified (this paper) to the little known species maculata from the eastern tropical Atlantic was increased by the larvae reported and figured as Teuthowenia mega lops by Roeleveld (1977) and Morales and Guerra (1977). I have examined Roeleveld's specimens and included them in my present study. The specimens of Morales and Guerra were not examined but I consider them very probably to be maculata. Like several earlier authors, Roeleveld considered Teuthowenia as a "larval genus" to hold megalops until the confusion regarding the valid genus was clarified. Clarification of the taxonomic confusion was not to be found in Imber's attempt to revise the cranchiids in 1978, but rather the problem was further complicated by his reshuffling of the various nominal species, misidentified specimens and unlike forms that have appeared over the years in the literature of the group. The different nominal species were assigned to two genera: Teuthowenia, comprised of the two species, megalops and antarctica, which Chun originally assigned to the genus; and Fusocranchia Joubin, 1920, comprised of one species pellucida. Imber recognized two subspecies of megalops: m. megalops, correctly identified as a North Atlantic form; and m. impennis (Robson, 1924), a form, distributed in the southern oceans north of the Subtropical Convergence, under which Me- galocranchia richardsoni Dell, 1959 and M. megalops australis G. Voss, 1967 were placed in synonymy. Imber distinguished Chun's unlike species antarctica solely on the basis ofthe originally described two rows of suckers on the tentacular stalks and of beaks which Imber identified to the species. The distribution of antarica was given as the southern oceans south of the Subtropical Convergence to about 65°S. As stated above, I have examined Chun's type of antarctica and found it definitely not to be of this genus. Unfortunately, the ontogenetic series necessary to determine the exact identity of the larval type is not yet complete. Imber's errors in resurrecting Joubin's Fusocranchia, the larval type of which I have examined and found to belong to Liocranchia, and his mistaken concepts of pellucida, and Berry's pardus, which is placed in the synonymy of pellucida, were discussed by N. Voss (1980). For the numerous erroneus synonymic assign- ments, the reader is referred to the synonymies given in the present paper. The affinity of megalops for Slope Water was demonstrated by the findings of Lu and Roper (1979). Stephen (1982) found the species to be one of the commonest squids off Canada. I have examined and included specimens from both studies in the present paper. Considering the abundance of the genus in the temperate and subpolar waters of the North Atlantic and in the region ofthe Southern Subtropical Convergence (Lu and Clarke, 1975a; Stephen, 1982; present paper), it is no wonder that the various nominal species have appeared regularly in the recent growing literature on cephalopod beaks found in the stomachs of predators. They have been reported in the stomachs of sperm whales (Clarke and MacLeod, 1974; 1976; Clarke et al., 1976; Clarke, 1980), northern bottlenosed whales (Clarke and Kristensen, 1980), blue sharks (Clarke and Stevens, 1974) and various marine birds (Imber, 1973; 1976; 1978; Imber and Russ, 1975). The exact identifications of the beaks, however, must be viewed with skepticism given the systematic confusion in the 8 BULLETINOFMARINESCIENCE,VOL.36, NO. I, 1985

Table I. List of nominal species of Teuthowenia

Cranchia maculata Leach, 1817 Megalocranchia pardus Berry, 1916 Cranchia (Owenia) megalops Prosch, 1847 Anomalocranchia impennis Robson, 1924 Leachia hyperborea Steenstrup, 1856 Desmoteuthis thori Degner, 1925 Desmoteuthis tenera Verrill, 1881 Megalocranchia richardsoni Dell, 1959 Desmoteuthis pe/lucida Chun, 1910 Megalocranchia mega lops australis G. Voss, 1967

group and the situation that most of the beaks were unaccompanied by identifiable remains, and few or no beaks, extracted from accurately identified specimens were available for comparison. Hopefully, a handbook for the identification of cephalopod beaks (Clarke, in prep. I) that resulted from an international workshop held in Plymouth, England, in June 1981 during which beaks extracted from specimens identified by cephalopod systematists were described and figured, will promote a higher degree of confidence in future studies. Finally after studying large, world-wide collections containing complete series of intermediate growth stages linking larvae with adults and examining the type- specimens of the 10 nominal species (Table 1), N. Voss (1980) redefined the genus in her revision of the Cranchiidae, and determined Teuthowenia Chun, 1910 to be the valid name. Verrilliteuthis Berry, 1916 and Anomalocranchia Robson, 1924 were declared junior synonyms. Voss recognized Teuthowenia, together with Megalocranchia and Egea, as belonging to one of three generic groups comprising the subfamily Taoniinae. The monophyly of the group, termed the Megalocran- chia group, was well corroborated in a subsequent analysis of the phylogenetic relationships among the cranchiid genera (N. Voss and R. Voss, 1983). Evolu- tionary rate inequalities estimated to have occurred in the Megalocranchia group are discussed by R. Voss et al. (1983).

MATERIALS AND METHODS

The 549 specimens I examined include a continuity of ontogenetic stages from early larval through adult for two of the species, megalops and pellucida. and early larval through early subadult for the third species, maculata. The specimens varied considerably in condition and frequently lacked various bodily parts. Not all size groups were equally represented. The examined material came from over the entire, in the case of megalops, or a wide part, in the cases of maculata and pe/lucida, of the geographic distribution ofthe species. All the type-specimens of the 10 nominal species (Table I) were examined, with the exception of the holotype and one of the three paratypes of Megalocranchia richardsoni. Some pertinent specimens misidentified in the literature were also examined. Collections are generally listed in alphabetical order by name of the collecting vessel or program. Depths are expressed in meters (m). Fishing-depths for DANA, GALATHEAand THOR hauls, originally given in meters of wire, were estimated using the method of Bruun (1943). The examined material belong to: Australian Museum, Sydney (AM); British Museum (Natural History), London (BMNH); Field Mu- seum of Natural History, Chicago (CNHM); Dominion Museum, Wellington (DMNZ); Fisheries and Oceans Canada, Halifax (FOH); Institut fUr Seefischerei und Zoologisches Museum der Universitllt, Hamburg (ZMH); Musee Oceanographique, Monaco (MOM); Institute of Oceanographic Sciences, Wormley (lOS); Laboratory, Marine Biological Association, Plymouth (MBA); Museum of Com par- ative Zoology, Harvard University, Cambridge (MCZ); National Museum of Natural History, Smith- sonian Institution, Washington, D.C. (USNM); St. John's Biological Station, Fisheries and Oceans Canada, St. John's, Nfld. (SJBS); S. S. Berry, private collection, Redlands, Calif. (SSB); Scripps In- stitution of Oceanography, La Jolla (SIO); South African Museum, Capetown (SAM); University of Miami Rosenstiel School of Marine and Atmospheric Science, Miami (UMML); University Museum, University of Tokyo (UMT); Yale Peabody Museum, New Haven (YPM); Zoologisches Museum der Humboldt Universitllt, Berlin (ZMH); Zoologisk Museum, University of Copenhagen (ZMC).

I M. R. Clarke. A handbook for the identification of cephalopod beaks (in prep.). voss: REVISIONOF TEUTHOWENIA 9

The problems presented by the larval types of the three recognized species were discussed with F. M. Bayer, Smithsonian Institution, a member of the International Commission on Zoological No- menclature. Following his suggestions, I have for each species selected and designated as a hypotype an adult or subadult specimen in good condition, and each species is characterized on the basis of that specimen, supported by data from additional material. Because of the close similarity of the member species prior to maturity and in an attempt to avoid needless repetition, the genus is defined in detail and the species are diagnosed. The early ontogenetic stages are similarly treated, broadly described for the genus and characterized for the individual species. The description of the maturity stage, on the other hand, where known, is specifically detailed. Terms, abbreviations, counts, measurements (given in millimeters (mm» and indices follow the definitions of Clarke (l962b; 1980; in prep.') [beaks], G. Voss (1963) [general], N. Voss (1969) [spermatophores], Toll (1982) [gladius], and Roper and G. Voss (1983) [general]. Because ofthe high degree of contraction (usually 10-40%, often externally undetectable) that occurs with these soft- bodied squids on preservation, all pertinent indices are based on gladius length (GL) rather than on the commonly used dorsal mantle length. The gladius was dissected and measured from all specimens, with the exception of the smallest larvae, on which meristic data were taken. Because of the difficulty involved, the gladius was not dissected from larvae smaller than about 45 mm ML; instead only specimens in the best and least contracted condition were selected for detailed measurements and counts. The use of mantle length (ML) in these instances is noted in the text and tables. Certain selected measurements and counts were made, however, on almost all of the specimens. Abbreviations and indices that appear for the first time in this paper, or that have been recently proposed or modified (Toll, 1982; Roper and G. Voss, 1983; Clarke, in prep") and are not yet in general use are: AF- formula of arm order based on comparative lengths; ASI, elIl -diameter of largest sucker on arm III as a percentage of gladius length; CILI -length of club as a percentage of gladius length; EDI -diameter across bulbus of eye as a percentage ofgladius length; FrRaWI, ant-greatest width at anterior end of free rachis as a percentage of gladius length; FrRaWI, mid-width at midpoint of free rachis length as a percentage of glad ius length; GiLl-length of gill as a percentage ofgladius length; GiLaC-count of gill lamellae on outer demibranch; GTILI-Iength ofgladius tail measured along the midline from the level of maximum width of the vanes to the posterior end of the gladius as a percentage of gladius length; LRLI - rostral length oflower beak as a percentage of gladius length; CISI-diameter oflargest sucker on club as a percentage of gladius length; MaSTC-count of teeth on largest manus sucker; NiGILI-length ofnidamental gland as a percentage ofgladius length; TtLl-length of tentacle as a percentage of gladius length; VLI-Iength of vanes measured along the midline from the anterior insertions to the posterior end of the gladius as a percentage of gladius length. In addition to the rostral lengths of the upper and lower beaks (URL; LRL) of selected specimens, the rostral edge visible from profile, crest, hood and wing lengths of the lower beaks were also taken in order to calculate the ratios of wing length to rostral edge (WiULREL), and crest length to hood length (LCrULHoL) which have been found useful in identifying beaks (Clarke, in prep. I). Abbreviations used in the tables to denote stage of development are: M - mature; nM-near-mature; S-sessile eye; St-stalked eye. Abbreviations used for collecting vessels are: B-BRANDAL: ELT- ELTANIN;T-THOR; WH-WALTHER HERWIG.Ranges when given for indices in some of the tables are followed by the means in parentheses. Total egg counts in mature females were estimated by aliquot sample. Individual eggs selected for measurements were taken from the oviducts. Serial sections of the glandlike organs on the anterodorsum of the mantle of mature females of Teuthowenia pellucida were stained with Mayer's Mucicarmine, Gomori trichrome or Schiff stain. Species synonymies include only references of systematic, distributional or biological significance. Only those references to specimens that I have been able to confirm either from an adequate description and/or figure or from actual examination of material are given.

Teuthowenia ehun, 1910

Owenia Prosch, 1847: 71 (subgen.) (type, by monotypy: Cranchia (Owenia) megalops Prosch, 1847; type in University Zoological Museum, Copenhagen; examined) [invalid junior homonym of Owenia Chiaje, 1844 (Polychaeta)]. Teuthowenia Chun, 1910: 376 [substitute name for Owenia Prosch, 1847]. Teuthowenia (Teuthowenia), Pfeffer, 1912: 741. Verrilliteuthis Berry, 1916: 64 (type, by monotypy: Desmoteuthis tenera Verrill, 1881: 412; type in U.S. National Museum of Natural History, USNM 574850; examined). Anomalocranchia Robson, 1924a: 9 (type, by original designation: Anomalocranchia impennis Rob- son, 1924a; type in British Museum (Natural History), BMNH 1924.9.9.38; examined). Hensenioteuthis (Teuthowenia), Thiele, 1934: 981. 10 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.1, 1985

Diagnosis.-Fins long, narrow, terminal-lateral; funnel-mantle fusion cartilages roughly oval with single or multipoint tubercle; funnel valve absent; dorsal pad of funnel organ with 1 median papilla plus 2 long, elliptical, lateral papillae; eye with 1 large plus 2 small, non-contiguous photophores; brachial end-organ present on ends of all arms in mature females; arms and clubs without hooks; carpal suckers in 4 rows, arranged in zigzag pattern, on tentacular stalks; digestive duct appendages on digestive gland and ducts; gladius with conus lacking; medium length, narrow, but not needlelike, hollow pseudoconus present in juvenile to adult stages. Definition.-Medium-sized to moderately large cranchiids (ca. 140-380+ mm GL). Conical mantle elongate, tapering from widest point near anterior margin to narrow posterior point; mantle wall thin, muscular; muscle layers not continuous across gladius. Small, spindle-shaped to oval funnel-mantle fusion cartilages with single or multipoint tubercle at mantle margin; small nuchal fusion cartilage elongate, oval to spindle-shaped, without tubercles; cartilage does not extend beyond anterior margin of mantle. In mature and near-mature females, row of glandlike organs present (? or absent) on anterior portion of middorsal line of mantle. Long, narrow, terminal-lateral fins with growth extend progressively up lateral margins of mantle to approximately 40-60% of mantle length, with anterior fin insertions opposite point of origin of gladius vanes; very small, anterior lobes present; fins taper very gradually posteriorly terminating in small lobes whose posterior margins extend beyond tip of gladius for very short distance and fuse with median cleft; in cross section fins thin, but muscular. Large, broad-based funnel extends to mideye level or beyond; dorsal pad of funnel organ inverted V-shaped, with 3 long papillae, elliptical in cross section (spatulate one on each narrow lateral arm and lanceolate one on mid posterior margin of anterior midportion); medium-sized ventral pads roughly triangular to kidney-shaped; funnel valve absent. Head dominated by huge, protruding, anteriorly oriented eyes with 3 non-contiguous, complex photophores (large, broad, crescent-shaped posterior photophore with smaller, long, narrow, crescent-shaped anterior photophore within its concavity on ventral surface, and third (rarely fourth) small, oval photophore on inner, anterodorsal margin). Olfactory organ small, narrow-stalked. Attachment formula for buccal membrane usually DDVV, with middorsal support bifurcate and remaining 6 supports sin&e. Short to medium length arms about 1/4 to 112 length of mantle, muscular, with length formula usually III>II>I>IV or III>II>IV~I in adult females, III> II> I> IV or III~I~II> IV in adult males, III> II> I> IV or III> II> IV=I in subadults. Small to medium-sized suckers greatly enlarge on midportions of III and II exceeding width of arms, similar but progressively less enlargement on I and IV; rings of basal and midarm suckers usually smooth; rings of terminal suckers with small, flat, usually blunt teeth on distal margins, except on modified ends of arms I (except in mega lops) and II in males. Well-developed, trabeculate protective membrane slightly more expanded on ventral margins of all arms; membranes basally fused to form low, inner web between all arms. Aboral keel present on distal half of III, and on distal quarters of I and II; lateral keel present on IV. All arms of adult males show some modification, particularly arms I and II, with varying degrees of elongation and increased robustness, and changes in dentition and overall appearances of suckers. Arms I may elongate with marked increased robustness, particularly in midportion, or terminal portion may elongate and display modified suckers on enlarged pedestals; terminal portion with suckers in 2 or 3-4 rows. Terminal portions of II elongate, with modified suckers on enlarged pedestals arranged in 2 or 3-4 rows. Modified suckers on both arms voss: REVISION OF TEUTHOWENIA 11

coequal, abruptly small or with slightly swollen appearance; apertures reduced; rings with uniform, minute, blunt to pointed teeth on entire margins. Modified, terminal portions of I and II sometimes display naked, whiplike ends. Suckers on basal portions of all arms with reduced apertures and swollen, fleshy collars. In mature females, all arms with brachial end-organs; glandular processes sometimes present near base of midarm suckers. Tentacles short, muscular, with 4 rows of carpal suckers set in zigzag fashion on distal 112 to 2/3 of stalk; carpal cluster absent; club slightly expanded with suckers on long pedestals; suckers gradually enlarge to midmanus with slight increased enlargement from dorsal to ventral margins; sucker rings incised on entire margins with numerous short, sharp teeth, longest on distal portion; trabeculate protective membrane well developed on both margins; dorsal keel present on distal half. Digestive gland stout, spindle-shaped, with small, spindle-shaped ink sac partially embedded in ventral third of median, posterior surface; digestive duct appendages in form of2 usually unequal-sized (left larger than right) lobes offollicles on posterodorsal surface of digestive gland and clusters of follicles which extend length of long, fused digestive ducts to small caecum. Spermatophore small to moderately large with long sperm mass, medium to long cement body, and short to moderately short ejaculatory apparatus; cement body without collar complex at oral end; aboral third of ejaculatory filament straight, midportion closely spi- raled prior to forming single, short loop at oral end. Mature eggs (known only for pellucida) small, mean size 2.6 x 1.9 mm; total egg count about 6,000-8,000. Gills short, about 10-13% of mantle length, with about 18-26 lamellae on outer demibranch. Lower beak with rostrum somewhat more than 112 length of wing; rostral edge nearly straight with hooked tip; obtuse jaw angle obscured in profile view by moderately low to prominent, rounded wing fold; shoulder ridge low or developed into distinct, broad-based, triangular tooth; hood set low or moderately high on usually straight, or sometimes slightly curved crest; distal edge of hood with moderately deep or shallow median notch; hood length about 112 that of crest; lateral wall with broad, prominent fold that extends to posterior edge intersecting at point about midway between crest and free comer. Upper beak with much curved rostrum; jaw angle usually slightly obtuse, sometimes slightly acute or slightly recessed; false jaw angle present; hood long, about % length of crest, set high on crest. Radula with tricuspid rachidian, bicuspid first lateral, and unicuspid second and third laterals; small marginal plates distinct or poorly developed; outer cusps of rachidian and first laterals small. Gladius with long, narrow, flexible free rachis that lacks dorsal keel; thin, delicate vanes bordering posterior 1/3 to in excess of 112 of length form long, moderately wide, spindle-shaped lanceola; lanceola with strong to moderately developed dorsal keel; posterolateral margins of lanceola ventrally infold and extend, without subsequent fusion, to form narrow, but not needlelike, hollow pseudoconus. Thin, outer layer of skin of mantle, fins, funnel, neck, arms and tentacles with finely mottled, dark, reddish-brown coloration. Under layer of skin with fewer and larger, discrete, reddish-brown chromatophores arranged in distinctive patterns on different body parts. (Expanded and modified from N. Voss, 1980: 400) Larva.-Larval stage extended, persisting to mantle lengths of about 53-100 mm. Larva distinguished by 4 rows of suckers extending nearly full length of tentacles; small, oval eyes, lacking pronounced ventral rostrum, set on short, stout stalks; short, stout arm pillar; stout, saccular, thin-walled mantle which becomes increasingly conical with growth; and distinctive chromatophore pattern. Chromatophore pattern of early larva of about 4-11 mm ML comprised of 12 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985 single large, reddish-brown chromatophores on midhead, mideye stalks, midarm pillar and midarms I on dorsal surface; and on midhead at base of funnel and mideye stalks on ventral surface. Single large, elongate chromatophore present on each side of buccal mass; row of large chromatophores extending full length of tentacles on aboral surface gives banded appearance to tentacles. Mantle with numerous small to relatively few large, oval (when expanded; mere points when contracted), reddish-brown chromatophores densely to widely spaced. Somewhat larger number of chromatophores occur on dorsal surface than on ventral. Ar- rangement of chromatophores on mantle irregular except for middorsal row which usually forms over free rachis of gladius. With growth, new chromatophores appear on developing structures, especially on funnel, clubs and on arms to form median row. Additional features and their progressive changes during larval development as follows. Anterior margin of mantle typically with scalloped appearance, about equally produced between 3 points of fusion. Single or 2-5 tubercles may appear on small irregular oval or spindle-shaped, funnel-mantle cartilages between about 10-80 mm ML. Fins appear in early larva as mere buds inserted on posterolateral margins oflanceola, widely separated by broad, rounded posterior end of gladius. With growth, fins become paddle-shaped, then roughly oval in individual outline. Separate fins progressively draw closer together and elongate as they follow formation and elongation of pseudoconus of gladius (Fig. 15a'-d'). Simultaneously, in mid to late larval stage, fins begin to grow anteriorly on mantle. Larva frequently found with posterior end and oftentimes head retracted into mantle. Large, broad-based funnel extends beyond level of midpoint of eyestalks. In early larva, dorsal element of funnel organ small, stout, roughly triangular pad with single, small, central papilla. By about 10-15 mm ML, lateral margins of pad have elongated and indentations on posterior margins define lateral arms on which 2 lateral papillae appear as small mounds. By late larval stage, sculpture of dorsal funnel pad closely resembles that of adult (Fig. 16d-f). Head short, length about 10-18% of mantle length. Small, oval eyes covered with golden iridophores except on dark dorsal surface. Eyes proportionately larger during early larval stage of about 7-18 mm ML than during midlarval stage of development. Subsequently, as larva approaches metamorphosis, eyes progressively enlarge to maximum diameter index of 10-11 in late, stalked-eye stage. Simultaneously, eyes gradually change shape to become more hemispherical. In early larva, first photophore appears as patch on narrow, posteroventral end of oval, stalked eye. With growth, patch becomes more defined, and enlarges to cover most of ventral surface of eye. At about 15-20 mm ML, second photophore forms as thickening along concave anterior margin of initial patch. As eye enlarges and gradually assumes more hemispherical shape, small, third, subtriangular to oval photophore, pinched off from inner anterior end of first photophore, becomes closely associated with second organ. All 3 photophores remain contiguous until near-sessile eye stage (Fig. 16g-i). Arm crown short (dorsal 2 pairs of arms first to appear followed by lateral and ventral pairs), with early arm length formula II> I> III> IV. Growth of arms III accelerated so that by about 12-15 mm ML, arm formula changes to III> 11=IV> I. This formula remains typical for succeeding larval stages during which arms grow at gradually increasing differential rates. In mid and late larvae, distal and lateral or entire margins of arm suckers incised with low, blunt teeth. Midarm suckers proportionately larger in early larva of about 7-20 mm ML than in midlarva up to shortly before onset of disproportionate enlargement of suckers on distal third- quarters of arms III and II. In late stalked-eye stage latter suckers enlarge to about twice diameter of basal ones. voss: REVISION OF TEUTHOWEN1A 13

Strong, moderately long tentacles often contracted in preserved larva. Suckers, which extend full length of stalk, arranged in 2's and 3's in basal 3-5 rows and thereafter in 4's, set in zigzag pattern (frequently obscured on contracted tentacles) characteristic of genus. Suckers set on low to moderately long pedestals; ring margins toothed. First definitions of club seen at about 10-15 mm ML with appearance of low protective membranes, small aboral keel and detectable, uniform enlargement of suckers (Fig. 15t). Stout, spindle-shaped digestive gland, covered with golden sheath of iridophores, oriented vertically to longitudinal axis of body. Digestive duct appendages found, as in adult, as two large lobes of unequal size (left larger than right) on posterodorsal surface of digestive gland and in clusters offollicles extending length of digestive ducts. Gills in larvae of about 5-24 mm ML, relatively small to moderate in length; gradually become proportionately smaller during subsequent larval development. Adult number of pairs of gill lamellae on outer demibranch, about 18-26, found in mid to late larvae. Beaks of early larva delicate with dark pigmentation confined to rostral edge and small portion of hood, gradually spread- ing with growth. Radula of mid and late larvae tends to be more homodont than in older animal, with outer cusps absent to poorly developed on rachidian and first laterals. Gladius of larva lacks true conus. Posterior end initially broad, rounded and spoon-shaped. At early stage, posterolateral margins tum ventrally and infold to form short, broad pseudoconus (Fig. 15g)which progressively elongates and narrows with growth.

Juvenile.-Onset of juvenile stage marked by development of sessile eyes. De- pending in part on environmental factors, metamorphosis may occur as early as 53 to as late as about 95 mm GL. It marks major change in developmental history of animal, that includes noticeable changes in growth of various parts of body. Growth of near-hemispherical eyes, limited in late larval stage to maximum diameter index of 10-11, irrespective of mantle length, abrupt (Fig. 21). Con- currently, 3 photophores on posteroventral surface of eyes begin complete sepa- ration (Fig. 16j). Narrow, anterior second organ which sometimes partially separates from large, posterior first organ during late stalked-eye stage, but usually remains attached at inner end, separates completely. Small, third organ, simultaneously, splits off from inner end of second organ and subsequently migrates to more anterior position. Metamorphosis of larva to juvenile accompanied by acceleration in growth of arms and rate of disproportionate enlargement of suckers on distal, third quarters of arms III (Figs. 23, 24) and II. Growth accompanied by gradual progressive loss of dentition on rings of suckers on basal portions of arms. Secondary sexual modifications of ends of arms I (except in mega lops) and II in males may first appear at this time, or be retarded to slightly later period in juvenile development. Because of continuing growth of arms during development, number of pairs of normal suckers prior to modifications varies considerably, from about 11-27, depending on varying lengths of larval stage and on timing of event in juvenile stage. Small suckers on affected ends of arms first appear as modified suckers rather than as normal suckers subsequently modified. Modified suckers characterized by: small, coequal size, abruptly reduced from that of normal suckers on preceding portion of arm; sucker rings with reduced apertures uniformly incised on entire margins with minute, sharp teeth, in contrast to rings of immediately preceding normal suckers which bear larger, blunt to pointed teeth confined to distal and lateral margins; and arrangement in 2-4 rows. Other characters show little marked changes in rate of development distinctive for juvenile stage: smooth progressive lengthening of fins as they grow posteriorly 14 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985 accompanying elongation of posterior end of gladius and simultaneously, grow anteriorly on mantle (Fig. l5e'); and little change in proportional lengths of tentacles and clubs or of maximum size of manus suckers. Progressive general enlargement of suckers on basal portion of club part of ongoing development of manus. Small sex organs continue to develop but show little proportional enlargement. Nidamental gland length index remains at 1-2 throughout juvenile to well into subadult stage. Additional chromatophores appear on different growing structures but overall pattern distinctive for larva retained in juvenile. Following initial changes that occur in early juvenile, growth proceeds at smooth continuous rate that makes definition of distinct adolescent stage impossible.

KEy TO THE SUBADULTS AND ADULTS OF TEUTHOWENIA la. Funnel-mantle fusion cartilages with multipoint tubercle (2-4 points; occasionally I or 5); gladius tail about 20-22% gladius length; males with ends of arms I and II modified with 2- 4 rows of specialized suckers; rings oflarge club suckers with about 26-32 teeth (circumglobal in area of Southern Subtropical Convergence) . ...._.. .__.... Teuthowenia pellucida Ib. Funnel-mantle fusion cartilages with single-point tubercle (rarely 0); gladius tail about 23- 29% gladius length; males with ends of arms I and II, or only II modified with 2-4 rows of specialized suckers; rings oflarge club suckers with about 19-26 teeth (Atlantic north of about 20OS) _.._ _ _.._ _.._.._ _ _ _.._.._..______.._ .._.._ _.._.._ _ _._...... 2 2a. Males with about 25-27 pairs of normal suckers on arms I and II prior to modified ends; diameter oflargest suckers on arms III about 3.2% (? or greater) gladius length (tropical eastern Atlantic) _.. _.._..__ _.._._..______._.._ _.._ _.__._ _.._._._.._ __Teuthowenia macu/ata2 2b. Males with about 15-19 pairs of normal suckers on arms n prior to modified end; arms I without modified ends, but elongate with marked increased robustness in midportion in mature males; diameter oflargest suckers on arms III about 2.0-2.8% gladius length (Atlantic north- temperate and subarctic waters) _.._ _._ _ _ _..__..__._._.._._. __ ._._.._. Teuthowenia mega/ops

KEy TO THE LARVAE AND JUVENILES OF TEUTHOWENIA lao Chromatophores densely spaced on mantle, approximately 16 chromatophores on 5-mm square on mediolateral dorsum at about 45-64 mm GL; funnel-mantle fusion cartilages with single-point tubercle usually lacking in larva, but appearing in early juvenile of about 65-70 mm GL; arms relatively long and noticeably disproportionate in length in larvae of 5-44 mm ML, with arms III usually 12-22% mantle length; largest suckers on arms III display marked enlargement with growth from about 1.2-2.0% mantle length in larvae of 5-44 mm ML, to 1.3-2.6% gladius length in larvae and juveniles of 45-84 mm GL, to as much as 3.2% gladius length in juveniles of 105-124 mm GL; gladius tail well developed to about 11-17% mantle length in larvae of 25-44 mm ML (tropical eastern Atlantic) __. Teuthowenia maculata (Leach) lb. Chromatophores widely or moderately spaced on mantle, approximately 4 or 8 chromatophores on 5-mm square on mediolateral dorsum at about 45-64 mm GL; funnel-mantle fusion cartilages with single or multipoint tubercle present in larva of about 10-60 mm GL; arms relatively short and less noticeably disproportionate in length in larvae of 5-44 mm ML, with arms III usually 3-12% mantle length; largest suckers on arms III display lesser degree of enlargement with growth from about 0.6-1.3% mantle length in larvae of 5-44 mm ML, to about 0.8-2.0% gladius length in larvae and juveniles of 45-84 mm GL, to about 1.2-2.2% gladius length in juveniles of 105-124 mm GL; gladius tail moderately developed, about 5- 12% mantle length in larvae of 25-44 mm ML (Atlantic north of about 32°N or circumglobal in the area of the Southern Subtropical Convergence) .._ _.._.._ _.._.._.._.._.._ _ _ _.._.._.... 2 2a. Chromatophores widely spaced on mantle, approximately 4 chromatophores on 5-mm square on mediolateral dorsum at about 45-64 mm GL; funnel-mantle fusion cartilages with single tubercle present at about 30-60 mm GL (Atlantic north-temperate and subarctic waters) . ...._._.._.._..______.._ _ _._.._.._ _.._ _._.._.._._ _._._._.._.._.__ Teuthowenia megalops (Prosch) 2b. Chromatophores moderately spaced on mantle, approximately 8 chromatophores on 5-mm

, Adults not known; only early subadult male known. voss: REVISIONOF TEUTHOWENIA 15

square on mediolateral dorsum at about 45-64 mm GL; funnel-mantle fusion cartilages with multiple tubercles usually present at about 25-30 mm ML (single-point tubercle usually present at about 10-20 mm ML) (circumglobal in area of Southern Subtropical Convergence) _.._ . ______. Teuthowenia pellucida (Chun)

Teuthowenia megalops (Prosch, 1847) Figures 1-3, 4a-e, 5-7

Cranchia (Owenia) megalops Prosch, 1847: 71, figs. 4-6 (west of Faeroes; type in University Zoo- logical Museum, Copenhagen; examined). Cranchia megalops. Morch, 1850: 57 [Prosch's description of type amended);-Steenstrup, 1861: 77 (p. 27 in English translation; discussion); 1881: 20 [po 121 in English translation); discussion; -Tryon, 1879: 162 [listed);-Hoyle, 1886: 44 [listed); 1909: 276 [listed); 1910: 411 [listed). Leachia hyperborea Steenstrup, 1856: 200 (Baffin Bay, off North Greenland; type in University Zoological Museum, Copenhagen; examined) [po 96 in English translation). Taonius hyperboreus. Steenstrup, 1861: 83 (p. 37 in English translation; discussion]; - Hoyle, 1885: 321 [type described]; 1886: 45, 191, pI. 32, fig. 12, pI. 33, figs. 1-10 [part: (?) PORCUPINEspecimens (Carpenter et aI., 1870); type figured). Loligopsis hyperborea. Tryon, 1879: 162 [Iisted);-Rochebrune, 1884: 12 [listed). Desmoteuthis tenera Verrill, 1881: 412, pI. 55, figs. 2-2d, pI. 56, fig. 3 (39°55'N, 70028'W; 116-mm ML syntype in Yale Peabody Museum, YPM 12462; 72-mm ML syntype in National Museum of Natural History, Washington, USNM 574850; examined) [date and depth of capture in error]; 1882a: 426, pI. 55, figs. 2-2d, pI. 56, fig. 3; 1882b: 216, pI. 55, figs. 2-2d, pI. 56, fig. 3; 1884: 245 (39°27'N, 69°56'W). Desmoteuthis hyperborea, Pfeffer, 1900: 192 [listed); 1908: 104, fig. 119 [listed); 1912: 708 [part: (?) synonymy citations Carpenter et aI., 1870: 423];-Degner, 1925: 61, figs. 42-44 (36°-66°N, 10-28°W) [development of young). Owenia megalops, Pfeffer, 1900: 193 [listed); 1908: 106, fig. 120 [part: (?) synonymy citation Hoyle, 1906: 161; listed]. Desmoteuthis hyperboreus, Hoyle, 1909: 277 [listed]; 1910: 408 [listed). Teuthowenia megalops. Chun, 1910: 376;-Pfeffer, 1912: 742, pI. 48, figs. 9-11 (41°35'N, 14°19'W) [part: not specimens from stations IN 180, 264 and 274, figs. 5-8, 17, 18);-Grimpe, 1922: 51 [listed); 1925: 98 [Iisted);-N. Voss, 1980: 401, fig. 13;-Stephen, 1982: 175, fig. 126 (Canadian Atlantic to 46°N; numerous specimens examined);-Nixon, 1983: 233, figs. 14.1-14.5 [life history); - Young, 1984: I fig. 5e-f, pI. IVe-h [morphology and discussion). Megalocranchia hyperborea. Berry, 1912: 644 [discussion];-Joubin, 1924: 94, pI. 7, fig. 7 (46°15'N, 10°11'W; 127, 128-mm ML specimens examined). Taonidium pfefferi. Massy, 1913: 8 (60003'N, 03°53'W; examined). Verrilliteuthis tenera. Berry, 1916: 64 [discussion]. Verrilliteuthis hyperborea, Grimpe, 1922: 51 [listed]; 1925: 98 [Iisted];- Thiele, 1934: 980 [Iisted);- Clarke, 1966: 237 [part]. Desmoteuthis thori Degner, 1925: 68, figs. 45-48 (46°30'N, 07000'W; type in University Zoological Museum, Copenhagen; examined). Verriloteuthis hyperborea. Massy, 1928: 35 [listed; erroneus spelling of Verrilliteuthis Berry, 1916]. Hensenioteuthis (Teuthowenia) mega lops, Thiele, 1934: 981 [listed]. Desmoteuthis megalops. Muus, 1956: 8, figs. 1-9 [part: not synonymy citations Helicocranchia pfefferi Massy, 1907, 1909; Pfeffer, 1912; Desmoteuthis pellucida Chun, 1910; Teuthowenia antarctica Chun, 1910; Massy, 1916; Megalocranchia pellucida, Pfeffer, 1912; Teuthowenia megalops, Joubin, 1920 (examined); Teuthowenia (Helicocranchia) pfefferi. Bouxin and Legendre, 1936. (part) Desmoteuthis hyperborea Pfeffer, 1908, 1912. (?) Leachia ellipsoptera. Carpenter et aI., 1870; Desmoteuthis pellucida. Chun, 1913; development and discussion]; 1962: 7 [part; discussion]. Megalocranchia megalops. Zuev and Nesis, 1971: 299 [part];-Lu and Roper, 1979: 16 (38°40'- 39000'N, 72000'-72°30'W; 107-mm ML specimen examined) [distribution]. Taonius megalops, Clarke, 1962: 41; 1966: 234 [part];-Dilly, 1972: 403, fig. I;-Lu and Clarke, 1975a: 150, 153, fig. 6 (53° and 60oN, 20°W) [vertical distribution);-Dilly and Nixon, 1976: 19, pis. 1-22, figs. 1-22 [growth and development]. Taonius thori, Clarke, 1966: 236 [listed]. Verrilliteuthis megalops megalops. Nesis, 1974a: 10 [part: (?) synonymy citations G. Voss, 1960; listed]. Teuthowenia megalops megalops. Imber, 1978: 470, fig. 6c [part: not synonymy citations Bouxin and Legendre, 1936; Megalocranchia megalops Voss, 1960: 433. (?) Teuthowenia megalops, Chun, 1913: 9; Desmoteuthis pellucida, Chun, 1913: 9; "Unknown oegopsid" Clarke and MacLeod, 1976: 744, fig. 3. Beaks of Degner's type of Desmoteuthis thori figured]. 16 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

Materials Examined.-257 specimens (4-380 mm (GL (ML)) from 113 lots, including holotype. Holotype.-?s, 25± mm ML, off Faeroe Islands, 1844, ZMC. Hypotype. -9,215 mm GL, Baffin Bay, offN. Greenland, 1848, ZMC (holotype of Leachia hyperborea Steenstrup).

Other Types. -I 9, 380 mm GL, THOR Sta. 19006d, 46°30'N, 07"00'W, 1,350 m, II IX 1906, ZMC (holotype of Desmoteuthis thori Degner).-2 cScS, 116, 72 mm ML, FISH HAWK Sta, 952, 39°55'N, 70028'W, 724 m, 23 VIII 1881, YFM 12462 (116 mm ML), USNM 574850 (72 mm ML) (syntypes of Desmoteuthis tenera Verrill). Other Material.-See Appendix. The following diagnosis is based on 31 adults and subadults (145-380 mm GL), including the hypotype (215 mm GL), from over the full geographic range of the species. Indices for all specimens selected for detailed measurements are summarized in Table 2. Diagnosis.-Moderately large species (males mature at about 182-244 mm GL; females at greater than about 350 mm GL). Funnel-mantlefusion cartilages with I single-point, usually stout tubercle (rarely 0) at mantle margin (Fig. 19). Gland- like organs on middorsal line of mantle in mature and near-mature females not known. Fins long (FLI approx. 39-55), narrow (FWI approx. 17-25). Eye large (EDI 12-16) (Fig. Ie). Attachment formula for buccal membrane usually DDVV, but sometimes second support, either right, left (hypo type) or both, bifurcate, ventral to I (occasionally to web between I and II) and dorsal to II. Arms with length formula III> II> I> IV except in adult males in which it is I> III> II> IV or III> I=II> IV; on arms III sucker pairs 8-14 usually largest, with greatest diameter (ASI, eIII 2.0-2.8) about 3 times diameter of basal suckers (Fig. 2h); in sub adults and adult females, rings of basal and midarm suckers usually smooth; rings ofsmall terminal suckers with about 6-10 low, flat, round, triangular or truncate teeth on distal and lateral margins except on modified ends of II in males. In mature males, all arms show some modification. Arms I elongate with increased overall robustness; mid portion markedly swollen with expanded protective membranes, terminal portion with 2 rows of normal suckers (Fig. 2a, b). Arms II with 15-19 pairs of normal suckers prior to elongate, usually swollen, modified terminal portions (about 23-32% of arm length) (Fig. 2c, d). Terminal portions with 2 rows (sometimes appearing as 3-4 rows because of crowding) of modified, slightly swollen suckers (often missing) bare, whiplike tip not present; protective membranes reduced. Arms III and IV with attenuate, whiplike ends. Basal and medial suckers on all arms with reduced apertures and swollen, fleshy collars (only traces remain in available specimens). Suckers on modified ends of II with minute, pointed teeth on entire ring margins (Fig. 2f); all other suckers on all arms with numerous small, irregular, rounded or triangular teeth on distal and lateral ring margins with teeth usually becoming progressively more pointed on midarm, especially on arms I (Fig. 2e, g). In large adult females (no mature females available) terminal 6-9% of all arms modified to form brachial end-organs, with about 28-43 pairs of normal suckers prior to end-organs. Small, irregular- shaped processes, partly comprised of glandular tissue, scattered on oral surface between largest suckers (approximate sucker pairs 8-15) on all arms (Fig. 3e). Largest suckers (CISI 0.8-1.1) of tentacular club with about 19-24 teeth (Fig. 1c, d). Spermatophore (9 spermatophores from 3 specimens, 199-215 mm GL) moderately large (SpL 35.7-56.4 mm, SpLI 16.5-26.2), with long sperm mass (SpMI voss: REVISION OF TEUTHOWENIA 17

Figure 1. Teuthowenia megalops. a. Subadult female, ventral view, hypotype, ZMC, 210 mm ML. b. Dorsal view, of same. c. Right tentacular club, WALTHER HERWIG 712/73, female, 187 mm GL. d. Largest manus sucker, of same. e. Right eye, ventral and anterolateral views showing photophores, USNM 816155, male, 199 mm GL. f. Nuchal fusion cartilage, WALTHER HERWIG 695/73, male, 215 mm GL. g. Right funnel-mantle fusion cartilage, USNM 816155. h. Funnel organ, WALTHER HERWIG 695/73. 18 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

a J d

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o ,"~' ,t" ['.

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r:, I. \. J \

Figure 2. Teuthowenia mega/ops. a. Entire right arm I of mature male showing sexual modifications, USNM 816155, 199 mm GL. b. Lateral view of distal half of same arm in frequently encountered state of contraction. c. End of right arm II of mature male showing sexual modification, USNM 816155,199 mm GL. d. End of right arm II of immature male showing sexual modification, UMML 31.1845, 184 mm GL. e. Sucker from last pair prior to modified terminal portion of right arm II, of same. f. Proximal sucker from modified terminal portion of right arm II, of same. g. Sucker (from 25th pair) from midportion of right arm I of mature male, WALTHER HERWIG 695/73, 215 mm GL. h. Brachial circlet with tentacle and buccal membrane elements of immature female, right half, hypotype. voss: REVISION OF TEUTHOWENIA 19

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Figure 3. Teuthowenia megalops. a. Gladius, ventral view, with enlarged cross sections and detail of pseudo conus, BRANDAL 6, subadult female, 254 mm GL. b. Upper mandible, single view, and lower mandible, three views, BRANDAL II, subadult female, 309 mm GL. c. Radula, of same. d. Spermato- phore, with detail of oral end showing ejaculatory apparatus, USNM 729972, 185 ± mm GL. e. Section of oral surface of left arm III showing glandular processes, UMML 31.1845, maturing female, 352 mm GL. f, End of left arm IV showing developing brachial end-organ, of same. voss: REVISION OF TEUTHOWENIA 21

Table 3. Teuthowenia mega lops. Spermatophore lengths and indices

WH 695/73 B 15 USNM 816155 Range GL(mm) 215 212 199 Number of spermatophores 2 6 2 SpL (mm) 54.8,56.4 35.7-41.2; 25.3* 39.2,40.7 35.7-56.4 42.5 SpLI 25.5,26.2 16.5-19.4; 11.9 19.7,20.5 16.5-26.2 20.2 SpWI 3.2,3.5 1.9-2.5; 2.4 2.2,2.3 1.9-3.5 2.5 SpMI 55.1,56.7 53.2-58.5; 45.1 53.1, 55.8 53.1-58.5 55.2 CBI 24.1,24.8 22.2-26.1; 29.3 24.1,24.7 22.2-26.1 24.2 EjAI 18.4, 19.3 18.0-20.6; 22.1 19.2,20.9 18.0-20.9 19.4 • Aberrant spermatophore; not included in ranges and means.

53.1-58.5), cement body of medium length (CBI 22.2-26.1) and moderately short ejaculatory apparatus (EjAI 18.0-20.9) (Fig. 3d; Table 3). Gills with about 23-26 lamellae on outer demibranch. Lower beak with rostral lengths of 4.8, 4.2 mm in mature males (2 specimens, 215, 244 mm GL), 5.6-7.0 mm in maturing adult females (4 specimens, 309- 380 mm GL); wing fold moderately low; shoulder ridge developed into distinct, broad-based, triangular tooth; hood set moderately high above straight crest; distal edge of hood with moderately deep median notch. Upper beak with rostral lengths of 4.8, 4.1 mm in mature males (2 specimens, 215, 244 mm GL), 5.0-6.0 mm in large maturing adult females (3 specimens, 309-335 mm GL); jaw angle slightly obtuse to slightly acute (Fig. 3b; Table 4). Radula with median cusp of rachidian tooth, inner cusp of first lateral, and single cusps of second and third laterals, moderately long and slender; rachidian and first lateral with broad, shallow base and small, stout outer cusps; small, marginal plates poorly developed (Fig. 3c). Gladius (5 specimens, 187-311 mm GL) with vanes bordering approximately posterior half(VLI 24-54); gladius tail about 1f4or somewhat in excess Ofl/4gladius length (GTILI 24-29); lanceola very thin in cross section, with moderately developed dorsal keel (Fig. 3a; Table 5). Larva.-Mantle or gladius lengths, indices and selected characters of young are summarized in Tables 2 and 15. Morphometric relationships with growth of selected characters are shown in Figures 21-24. Ninety-seven larvae were available, of which 30 were selected for detailed measurements and counts. The selected specimens are from the full distributional range of the species except for the northernmost section, off Iceland and Greenland, an area from which no larvae are available.

Table 4. Teuthowenia megalops. Beak measurements, indices and ratios

USNM T 190"d B 15 B9 B II 729989 WH 695/73 Range

Sex 'i' 'i' 'i' 'i' cSM cSM GL (mm) 380 335 311 309 244 215 URL(mm) 6.0 5.0 6.0 4.1 4.8 LRL (mm) 7.0* 6.0 5.6 5.8 4.2 4.8 LRLI 1.8 1.8 1.9 1.7 2.2 1.7-2.2 1.9 WiULREL 1.7 1.9 1.8 2.1 2.0 1.7-2.1 1.9 LCrULHoL 1.9 1.8 1.9 1.7 2.1 1.7-2.1 1.9 • From Imber, 1978: 472. 22 BULLETINOFMARINESCIENCE,VOL.36, NO.1, 1985

.', ;..•• ~J 'f •.- ~ ~ ) ,t ~~\

Figure 4. Teuthowenia megalops. a-e. a-c. Developmental series of young, dorsal views. a. BELO- GORSK7903-IB-34-I(S), II mm ML. b. BELOGORSK7903-41-2, 29 mm ML. c. BELOGORSK7905-86- 2, 60 mm ML. d. Right funnel-mantle fusion cartilage, showing tubercle, of same late larva. e. Left arm III, of same. Teuthowenia maculata. f-j. f-h. Developmental series of young, dorsal views. f. GALATHEA8, II mm ML. g. USNM 816157, 27 mm ML. h. DISCOVERY7824-22, 56 mm ML. i. Right funnel-mantle fusion cartilage, showing absence of tubercle, of same late larva. j. Left arm III, of same. Teuthowenia pellucida. k. Left arm III, SAM A4981-W5-3, 57 mm ML. (e, j, k drawn to same scale. Chromatophores on mantle shown expanded in a-c, g, h; partially contracted in f.) voss: REVISION OF TEUTHOWENIA 23

Table 5. Teuthowenia megalops. Gladius indices

USNM USNM 89 86 WH 712/73 729989 816155 Range

Sex Q Q Q ~M ~M GL(mm) 311 254 187 244 208 VLI 45 50 46 54 48 45-54 49 GWI 10 10 10 II 12 10-12 II GTILI 29 29 25 24 25 24-29 26 FrRaWI, ant 0.6 0.7 0.6 0.7 0.8 0.6-0.8 0.7 FrRaWi, mid 0.6 0.6 0.5 0.6 0.5 0.5-0.6 0.6

End of larval stage occurs between about 75-95 mm GL. Larva specifically recognized by large, oval chromatophores widely spaced on mantle (Fig. 4a-c); average of 4 chromatophores on 5-mm square on mediolateral dorsum of larvae of 55-60 mm GL. Characteristic pattern seen on smallest, well-preserved specimen of 8 mm ML. Single tubercle on funnel-mantle fusion cartilages, typical for species, usually appears during midlarval development between about 30-60 mm GL (Fig. 4d). Disproportionate arm growth does not occur in early larva. In specimens of 8- 11 mm ML, arms of about uniform length with mean length index of 5 (range, 4-6). During subsequent growth arms change to length formula III> II~IV> I which is maintained throughout remainder of larval period. Because of gradual increase in size of suckers as they extend out arms, sucker enlargement on distal third quarter of arms III noticeably, but not markedly, disproportionate (Fig. 4e). Gills small, with mean length index of 15 (range, 13-16) for larvae of about 8- 21 mm ML; proportionally decrease in length with growth to mean index of 11 (range, 10-12) for late larvae of 82-92 mm GL. Positive allometric growth of pseudoconus of gladius moderate. Gladius vanes, throughout larval stage, very thin in cross section resulting in exceptionally delicate lanceola, with faint sug- gestion of dorsal keel in late larva. Juvenile. -Juveniles were available from across the range of the species in the western Atlantic and in the eastern Atlantic south of 50oN. Over distribution of species, size at which eyes become sessile, marking begin- ning of juvenile stage, varies between about 75 and 95 mm GL. Single, small tubercle on anterior margin offunnel-mantle cartilages, typical of species, usually developed, rarely lacking.

Table 6. Teuthowenia megalops. Comparison of indices of selected characters showing morphological differences between early-metamorphosed juveniles and late-maturing larvae at the same approximate gladius length

Early-metamorphosed Late·maturing juveniles (79-94 mm GL) larvae (76-92 mm GL) N = 5 N=4

Index Range Range ALI, I 14-18 15 10-13 12 II 19-24 21 13-16 15 III 24-28 25 17-22 19 IV 15-22 18 12-15 14 ASI, ellI 1.8-2.0 2.0 1.2-1.7 1.4 EDI 13-14 13 9-11 10 24 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

Immediacy of increased growth of arms and disproportionate enlargement of suckers on third quarters of arms III that occur subsequent to metamorphosis evident when early juveniles of 79-97 mm GL are compared with young still in late, stalked-eye stage at 76-92 mm GL, approximately same size range (Table 6). Modifications on ends of arms II in male first detected in early juvenile stage. By 125-130 mm GL, modified ends may have elongated to about 8-16% of arm length. Arms I elongated in both sexes until they equal or slightly exceed length of arms IV. Modifications of midportions of arms I in male do not appear in juvenile but develop at much later stage when animal nears maturity. Proportional length of gills in juvenile approximately same as found in late larva, Marked changes in posterior end of gladius occur in early juvenile stage. Pseudoconus elongates rapidly, resulting in mean gladius tail length index of 25 (range, 23-28) in juveniles of 79-94 mm GL, approximately same proportional length as found in adult male, At the same time, vanes slightly thicken and dorsal keel gradually becomes more pronounced. Maturity.-Size at maturity shows a marked degree of sexual dimorphism, with females maturing at a much larger size than the males. Unfortunately, no mature females are available. Thirteen of the largest females, 248-380 mm GL, are in the early stage of maturing: the body texture is firm, and the brachial end-organs on all the arms are at the initial stage of development (Fig. 3f), lacking heavy pigmentation and with only slightly expanded protective membranes. The suckers remain on the forming end-organs, and the sucker pedestals are swollen and expanded basally to form alternating ridges, each extending to the midline of the arm. The oviducal and nidamental glands of these females are slightly to moderately enlarged to about 3-11 % of gladius length, and the egg mass shows only moderate enlargement. In the 380-mm GL female (type-specimen of Desmoteuthis thon), the eggs (now long preserved) are very small (about 0.6 x 0.5 mm). As the brachial end-organs develop in the maturing females, the numerous glandular processes (Fig. 3e), described and figured by Degner (1925: 70), appear between the largest suckers on all the arms, usually appearing last on arms IV. The processes, found in all available females over about 270 mm GL, are swollen, irregular-shaped papillae whose smooth or folded surfaces are largely covered with glandular epithelium. Eight of the largest males, 182-244 mm GL, including all in excess of 196 mm GL, are mature. The body texture shows no signs of deterioration; the reproductive organs are greatly enlarged; and the long, muscular-walled penis extends often as much as 30 to 50 mm out of the mantle opening. One to about 28 moderately large spermatophores were found in the spermatophoric sac and/or penis in each individual, with an additional I or more in varying stages of completion in the spermatophoric organ. Spermatophores extracted from 3 males of 215,212 and 199 mm GL display a 3-15% variation in length within an individual, with one exceptional, small spermatophore from the 212-mm GL male being reduced by about a third in length (Table 3). Despite the variation, the proportional size of the internal parts of the spermatophores remains approximately the same, except in the aberrant small one in which the sperm mass is somewhat reduced. The spermatophores from the 212-mm GL male in which the greatest length variation occurred, and some or all of those from several additional males display varying degrees of imperfections in folding and spiral features of the ejaculatory apparatus. The spermatophores showing the most imperfections tend to be the smallest and were found in the penis, while those taken from the spermatophoric sac tend to show progressively less imperfections with increased length. The evidence suggests voss: REVISION OF TEUTHOWENIA 25

that the imperfect spermatophores may, at least in some cases, not be the result of internal rupture due to preservation or disturbance but rather represent the early spermatophores produced by the still imperfectly functioning spermatophoric organ of the maturing male. Besides the sexual organs, maturation in males involves various changes in all the arms. In the mature male, all of the arms are proportionally longer than in the female, particularly arms I which are very robust, especially in the midportion where the protective membranes are markedly expanded (Fig. 2a, b). The ends are normal on arms I and modified on arms II. The modified ends comprise about 23-36% of the arm length, are swollen, and often lack most of the small, specialized suckers (Fig. 2f) leaving only the palisadelike formation of the enlarged pedestals (Fig. 2c). Additional changes in the mature male involve the whiplike attenuation of the ends of arms III and IV and the modification of the basal, and possibly the medial suckers, of all the arms. The modified suckers have reduced apertures and swollen, fleshy collars. Only fragments of these collars remain in the available animals, but they appear to be similar to those found intact in the mature males of pellucida (Fig. 13f). In addition, all of the sucker rings of the arms of the mature male, with the exception of those on the modified ends of arms II, display increased dentition on the distal and lateral margins with the teeth usually becoming progressively sharper on the midportions especially on the markedly enlarged arms I (Fig. 2g). Four males, 184, 184, 155 and 145 mm GL (all from off Newfoundland) are judged to be near-mature. They display all the features of the mature male (developed to a slightly lesser extent) with the exception of the modified ends of arms II (17-25% of arm length) which are not swollen and usually retain the specialized suckers set on slender, moderately long pedestals (Fig. 2d). Addition- ally, the sexual organs are but moderately large with no signs of forming spermatophores. Two of the largest males, 190 mm (Newfoundland) and 189 mm GL (eastern Atlantic) are still clearly immature. They show no enlargement or expanded membranes on the third quarter of arms I; the modified ends of arms II measure 22% (western Atlantic specimen) and 9% (eastern Atlantic specimen) of the arms' lengths; the ends of arms III and IV are not whiplike; and the arm suckers show no signs of either swollen collars or increased dentition and reduction of the apertures of the rings. In addition, the sexual organs are only slightly (eastern Atlantic specimens) to moderately enlarged. The sequence of appearance of the various maturity-related changes just described for the male begins with the first detection of the modification of the ends of arms II in the early juvenile, followed by the onset of disproportionate growth of all the arms at about 120-140 mm GL. The various other features appear over a short period close to the sexual maturity of the animal. Over the geographic range of megalops. mature animals were taken in the months of April, July, September and December. Captures oflarvae less than 10 mm ML were made in February, April, May and July. The evidence indicates that maturity and spawning occur over an extended period, possibly year-round. Geographic Distribution. - The zoogeographic map and terms for pelagic faunal regions and provinces in the Atlantic as given and defined by Backus and Craddock (1977) and Backus et al. (1977) are used for mega lops and maculata. Teuthowenia megalops inhabits the highly productive, subarctic and northern temperate waters of the Atlantic Ocean (Fig. 5). All captures, with the exception of scattered larvae, were made over bottom depths in excess of 1,000 m. Based 26 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

70 60 50 40

10 10

o o

-10 10

90 80 70 60 50 40 30 o -10 -20 -30

Figure 5. Geographic distribution .• Teuthowenia megalops; .•. Teuthowenia maculata. Each symbol in this and the subsequent map indicates the locationof one or more specimens reported in this paper; some stations that overlap greatly are not shown. See text for exact number and location for each record. The scheme of pelagic faunal regions and provinces shown are from Backus and Craddock (1977). The heavy boundary lines define the regions, the light lines, the provinces, Only those areas referred to in this paper are named. ATLANTIC SUBARCTIC: 1- Atlantic Subarctic. NORTH ATLANTIC TEMPERATE: 2-North Gyre, 3-Slope Water, 4-Azores-Britain, S-Mediterranean Outflow, 6-Western Mediterranean Sea, 7-Eastern Mediterranean Sea. MAURITANIAN UP- WELLING: I8-Northern Mauritanian Upwelling, 19-5outhern Mauritanian Upwelling. ATLAN- TIC TROPICAL: I6-Guinean.

on the 257 specimens from this study, the southern boundary of distribution in the eastern half of the Atlantic is at about 44°-45°N. This boundary is supported by (1) the absence of mega lops in the day and night series of hauls made with closing RMT combination nets between the surface and 2,000 m at 30oN, 23°W and 40oN, 200W reported by Clarke and Lu (1974) and Lu and Clarke (1975a); (2) the lack of the species in the collections made by the WALTHERHERWIG using the 1,600-mesh Engel trawl on the 1971 Cape Town-Madeira transect; and (3) the first encounter of the species at 46°N, 08°W by the DANAon the Cape Town- Denmark transect of the 1928-1930 Round-the-World cruise during which nonclosing, stramin nets were largely used. On the 1931 DANAcruise to the area of Madeira-Azores and to the north, megalops was taken at 48°N, 14°W on the voss: REVISION OF TEUTHOWENIA 27

southward leg and was not encountered again until 48°N, 29°W on the northward leg. It was subsequently taken at 3 out of 4 stations (75%) to the northernmost station at which the successful catching gear was used at 55°N, 300W. The species was not captured on the ANTONDOHRN1979 cruise in any of the numerous hauls using the 1,600-mesh Engel trawl in the Sargasso Sea. Along the diagonal transect from the southwest, the first capture, a 71-mm GL larva, was made at 42°N, 24°W, after crossing into temperate waters. This was the only capture in 5 stations to 45°N, 210W. Thereafter captures, which included young as well as maturing females and large subadult males, were made at 5 out of the 7 stations (71%) between 44°-48°N, 09°-18°W, an area higher in productivity than that to the southwest. A hydrographically ill-defined boundary in the region of 44°N in this area off the Iberian Peninsula is suggested by preliminary results from recent DISCOVERYcruises (Angel, 1979), and confirmed by the limits of distribution of certain species of euphausiids (Brinton, 1975) and myctophids (Hulley, 1981). The single capture of the larva at 42°N by the ANTON DOHRN together with Degner's (1925) record of 5 larvae (under the name Desmoteuthis hyperborea; not examined but probably of this species) taken by the THORbetween 36°-400N, 8°_ 100W indicates that at least the larvae of megalops are scattered southward, possibly to the northern limit of subtropical waters at about 32°-35°N (Worthing- ton, 1976). There are no records of mega lops from the Mediterranean. The northern boundary of distribution in the eastern half of the Atlantic is at about 65°-66°N between Greenland and Iceland and 62°-63°N in the Norwegian Sea, just south ofthe secondary Polar Fronts. Since the front occurs in both areas over broad oceanic ridges considerably shallower than 1,000 m, it is problematical whether the front or the shallow bottom depths, or both serve to limit the northward distribution. All size stages from larvae to near-mature or mature adults were found over the range between 44°-45° and 62°-66°N, but not in equal abundance. Lu and Clarke (1975a) reported that mega lops is the commonest of 7 species of cephalopods captured, 195 of 373 specimens, or 52% of the total catch, in a series of hauls at 53°N, 200W. A similar series of hauls at 600N, 200W caught 281 cephalopods comprised of 3 species of which 54, or 19%, were megalops. At the former station, Gonatusfabricii and Brachioteuthis riisei were the second and third commonest species, comprising 25% and 17% of the total catch, while at the latter, higher latitude station, G. fabricii comprised 74% of all cephalopods taken, dis- placing mega lops as the commonest species. (A part or all of these specimens of Gonatus may be identified to G. steenstrupi (Kristensen, 1981a).) In the western half of the Atlantic, mega lops is primarily confined to the subarctic Labrador-Coastal Water and the temperate Slope Water over bottom depths in excess of 1,000 m. The southern boundary occurs at about 36°N, the southern limit of the Slope Water (Iselin, 1936). Scattered captures of single larvae, however, have been made in subtropical waters to the south. The Ocean Acre Study took 4 larvae, 8-43 mm ML, using the 3-m IKMT in the area of Bermuda, between 310-33°N, 63°-64°W, and 2 additional larvae, 33, 57 mm ML, including one from the stomach of a lanceletfish Alepisaurus caught at 300N, 69°W, have also been identified from other collections. The lack of mega lops in the collections from 5 hauls made by the ANTONDOHRNon the 1979 Sargasso Sea-Cruise using the 1,600-mesh Engel trawl in depths of 170-~ 1,800 m in approximately the same area of Bermuda, between 28°_32°N, 60°-67°W, suggests that there probably is not a breeding population in these warmer, less productive waters. These expatriated larvae may have been transported by cold-core rings that have separated offfrom the Gulf Stream. Such rings are known to carry cephalopods (Lea, 1978) 28 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.1, 1985 and other organisms of Slope Water origin into the Sargasso Sea (Wiebe et al., 1976; Backus and Craddock, 1982). The eastern boundary of the range in the western Atlantic is marked by the coastward margin of the Gulf Stream in the south and by the Polar Front in the north. The available material is inadequate to determine the northern distribution boundary. Stephen (1982) shows the northern limit for the species at about 46°N. However the BRANDALcaptured 42 specimens ranging in size from larvae to mature adults, 62-352 mm GL, in 14 hauls using a large Engel trawl in an area largely to the north, between 42°-51 oN, 44°-52°W. It is not known whether the capture area was the northernmost fished because the complete data for the cruise was apparently destroyed in a fire in 1979 at the Bedford Institute of Oceanography (S. Stephen, pers. comm.). Considering the tolerance of megalops for the colder waters, the common occurrence of the species in the collections made at 50°- 51oN and the presence of suitable bottom depths to the north, I would expect the normal range of the species to extend to higher latitudes. The only specimen of mega lops from far northern waters of the western Atlantic that I have examined or can accurately identifY from the literature is the 215- mm GL hypotype taken in 1848 by Kielsen from off North Greenland. Though the precise location of capture is not known, Steenstrup (1861) referred to the locality as "high up in Baffin Bay." The subarctic region extends well up into the bay and a considerable portion of the central sector has bottom depths in excess of 900 to over 2,000 m. The area, however, lies north of, and is separated from, the known range of the species to the south by the broad, shallower depths of Davis Strait (~300-900 m). Whether the specimen is an expatriate transported northward by the West Greenland Current or is a representative of an unreported population in the Baffin Basin is not known. There is also the possibility that the locality is inaccurate. Captures oflarvae and mature adults indicate that the species breeds and spawns in the western half of the Atlantic between about 36°N to about 5loN. In this area, the major concentration of mega lops appears to occur in the subarctic waters of the northern sector. Stephen (1982) found that the species is one of the commonest captured in Canadian waters to about 46°N. To the south, occurrence and abundance in the region of the Slope Water may be affected by intrusions of shelf water and warm-core eddies from the Gulf Stream (Lu and Roper, 1979). The gap in distribution shown on the map between the populations in the eastern and western Atlantic probably reflects, in part, lack of sampling. To my knowledge, there are no collections of cephalopods from this area, which contains 2 distinct regions, the temperate Northern Gyre to the south (Worthington, 1976) and the subarctic waters to the north. I would not expect a continuous distribution across the gyre. The sharp gradient of the Polar Front would probably deter crossing from the west and north; to the east the lower productivity of the adjoining temperate waters, which appears to be less preferred by the species, would diminish the likelihood of crossing from that direction. The distribution is probably continuous across the central portion of the Atlantic in the subarctic region north of the gyre, at about 500-52°N. Teuthowenia megalops displays the Northern Subpolar-Temperate distributional pattern described by Backus et al. (1977), based on mesopelagic myctophid fishes. Restricted to, but not distributed throughout the Atlantic Subarctic and North Atlantic Temperate Regions, the species is absent, or virtually so, from the temperate provinces of the west and east Mediterranean Sea and Mediterranean Outflow, and probably from the Northern Gyre. The distribution in the temperate voss:REVISION OF TEUTHOWENIA 29

Imm.lulesp,clm.n$ _ M.lunni '.m~lu,rill oz;a;ao nut-muur. miles

Mature mllu 60. ,..

.c 1000 C.

!JOG ".. - 2600

2700 60 80 1 0 •• 0 1/:10 10 200 2tlO 20 :I 0 320 3AO 30 J 0 Gladius length (mm)

Figure 6. Teuthowenia megalops.Vertical distribution with respect to stage of maturity. Data used in this and the subsequent vertical distribution graphs, except when otherwise stated, are from specimens collected with closing and nonclosing nets.

Azores-Britain Province is largely affected by productivity and depth of bottom; the distribution is poorly known in the Subarctic Region. The distribution pattern of megalops also conforms to the Subarctic-North Temperate Pattern described by Krefft (1976) on the basis ofa number offamilies of mid water fishes, and the Boreoarctic Pattern of Hulley (1981), which is based on the myctophid fishes. Vertical Distribution. - Figures 6 and 7 summarize the vertical distribution data on all specimens included in this study for which information is available. The majority of the data for this and the subsequent species is based on captures made with non-closing nets, therefore the exact depth of capture for most of the specimens cannot be determined. Additionally, the amount of sampling, and types of nets used are not the same for all depths, and fishing time and depth were selected in many cases in anticipation of maximum catches. With these limitations, certain cautious conclusions can be drawn. Teuthowenia megalops has been taken by nets fishing in the upper 40 m to depths of 2,650 m, over temperature ranges in the vertical column of 21°_5°C in the Slope Water (0-800 m; Lu and Roper, 1979) and 13°-3°C in the temperate eastern Atlantic (85-2,650 m; WH/73). The pattern of vertical distribution appears to be similar throughout the known geographic range of the species. The few captures oflarvae smaller than 10 mm ML were scattered in the water column to depths as great as 1,000 m. Larvae from about 10 to 60 mm GL appear to be concentrated, both day and night, in the upper 200 m. Those larger than about 30 mm ML appear to display an expanded range extending down to about 500 m but are concentrated in the upper 300 m. Older larvae of about 60 to 80 mm GL also appear to concentrate, at least at night, in the upper 200 m. These distribution patterns are supported by Lu and Clarke (1975a) using closing nets at 53° and 600N at 200W. Though I did not examine their specimens, the material (under the name Taonius megalops) was described and figured, in part, by Dilly 30 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.1, 1985

.,. N1Rhl _ D.,T"'llIlllt~

BO. E ;1000 ==- 0. Q)

°1200

'2000 - ~

7600

2700+D--r-.",--,...... "~-,.:-...... -,l'==t---:!7"-.---,,.,-,-..,,'.F=D =F""f:-DD-'---"E. ='-:,r.,.~,'E,.=t-""'-'-,:-r:,.--r-T"i' .'-'-T3 ••~""'",:-r-""'3.'-a .,...... "",-.~3' • Glad,us length lmm) Figure 7. Teuthowenia rnegalops. Vertical distribution with respect to time of capture.

and Nixon (1976). How many of the captures of mid and late larvae in the deeper waters shown in Figures 6 and 7 are contaminates, caught in the upper waters during the descent or retrieval of the open nets, cannot be determined. The likelihood increases with the depth of the tow. Following metamorphosis at about 75-95 mm GL, the juveniles appear to extend their vertical range possibly to 1,000 m. Over this range, the juveniles may show diel migration. Lu and Roper (1979) recorded 2 specimens, 65 and 92 mm ML (described as subadults under the name Megalocranchia megalops) taken during the day with closing nets fishing at 600-800 m at approximately 39°N, now. Because of preservation contraction, the above specimens together with those recorded by Lu and Clarke (1975a) are probably larger by as much as 20% than the given mantle lengths. The catch data is meager for the larger sizes because of the apparent increasing elusiveness of the animals and the limited use of the larger collecting gear at all depths, particularly in the deeper waters. The data, though scanty, does suggest that with growth the immature animals seek greater depths possibly to 2,000 m or more. The few relatively shallow night captures suggest that some vertical migration may occur. Even though no mature females have been caught, the recorded captures of larvae smaller than 10 mm ML scattered to about 1,000 m, using both closing (Lu and Clarke, 1975a) and non-closing gear (this study), and the captures of mature males only below about 1,000 m combine to suggest that the mature female mates and spawns in the deep midwaters. The evidence suggests that hatching occurs in the greater depths. Subsequently, the larvae may float or swim up to shallow waters. Geographic Variation.-Litt1e morphological differentiation has occurred in Teu- thowenia megalops across its range in the highly productive waters of the North Atlantic. Though substantial differences are found in the larvae and juveniles from the two sides of the ocean, the majority of the differences are submerged with subsequent growth and are not exhibited in the subadults and adults. A comparison of young from between 42° and 51oN, 44° and 600W in the western Atlantic with voss: REVISION OF TEUTHOWENIA 31

Table 7. Teulhowenia megalops. Comparison of indices of selected characters showing geographic variation found in young of75 to 95 mm GL between 2 areas: area A, western North Atlantic, between 42° and 51°N, 44° and 600W; and B, eastern North Atlantic, between 44° and 52°N, 7° and 200W

Area A Area B N =4 N = 5 Index Range Range ALI, II 14-21 17 17-24 20 ALI, III 17-25 21 21-28 25 ASr, elII I.I-I.3 I.2 1.5-2.0 1.9 ED! 9-13 11 13-14 13 GTILI 17-18 18 23-28 25

young from between 44° and 52°N, 7° and 200W in the eastern Atlantic shows that at a given size, particularly between about 75-95 mm GL, the eyes and the enlarged suckers on arms III are usually larger, and the gladius tail and the arms are usually longer in the latter area (Table 7). The variations observed may be the results of differences in either the length of the larval stage or the growth rates in the two areas. In the eastern area metamorphosis occurs between about 75-80 mm GL while the western area it occurs between about 88-95 mm GL. The timing appears to relate to the temperature in the shallow waters where the young develop. The event is accelerated in the relatively warm temperate waters in the former area and is retarded in the cold, largely subarctic waters in the latter. Once metamorphosis occurs, the above morphological differences gradually disappear with growth. A comparison of the subadults and adults shows that such characters as the size and dentition of the suckers on the tentacular club and the arms, the exact pairs of suckers on arms III that show maximum enlargement, and the number of normal suckers prior to the modifications on the ends of arms II in the males appear to vary to the same degree in all areas of the distribution. Differences are found, however, in the degree of development of the modified ends of arms II and the lengths of all the arms at a given size in the subadult and immature adult males from the eastern and western areas. In the eastern area, immature males of 143-189 mm GL (3 specimens) have the terminal 7-9% of the arms modified, while 13-25% of the arms are modified in immature males of 145-190 mm GL (6 specimens) from the western area. In addition, the arms in the eastern males may be as much as 10 to 30% shorter than in the western males. Though the sometimes stretched condition of the arms in the western material might account for some of the differences found, the degree of development of the internal sexual organs supports the conclusion that sexual maturity is delayed and the males may mature at a larger size in the warmer waters of the eastern half of the Atlantic. Unfortunately, no mature males are known from this area. Comparable immature male material is lacking from other sectors of the distribution, but in the central sector and in the slope water, males are known to be mature over the same approximate size range of 182-244 mm GL as is found in the western area. The material available is too meager to show if similar growth differences occur in the females of the species. Remarks. - The type-specimen of Cranchia (Owenia) megalops Prosch is in poor condition, soft, browned, considerably cut up and missing various bodily parts. However the characteristic 4 rows of carpal suckers can be seen on the single remaining tentacle, and the pattern of widely-spaced, relatively large, oval chro- 32 BULLETINOFMARINESCIENCE,VOL.36, NO. I, 1985

matophores is still discernible on the mantle. Both oval eyes on short, stout stalks are intact, as well as the nuchal and funnel-mantle fusion cartilages, but only 4 of the small arms remain, mostly devoid of suckers, and the posterior end of the mantle is partially inverted, obscuring a clear view ofthe fins and their insertions. Since there was no need for confirmation of additional characters for specific identification, and considering the delicate condition of the specimen, I refrained from further dissection. My measurement for the mantle length is 25± mm. The type-specimens of the junior synonyms, Leachia hyperborea (designated as hypotype of mega lops in present paper), Desmoteuthis thori and Desmoteuthis tenera were all examined and found to be in good, specifically identifiable condition. In addition to the 72-mm ML syntype of D. tenera deposited in the National Museum of Natural History, Washington, I located Verrill's second syntype in the collections of the Yale Peabody Museum. The deposition of the latter specimen, the 116-mm ML male originally figured by Verrill (1881), was listed as unknown by Roper and Sweeney (1978). In addition to the literature records of the various synonyms of megalops, confirmed or unconfirmed, noted in the Historical Review, I examined in the British Museum the 37±-mm ML larva from off Ireland which Massy (1913) misidentified to Taonidium pfefferi Russell, 1909. Massy's specimen has the typical ocular photophores, single tubercle on the funnel-mantle fusion cartilages, funnel organ, and arrangement of carpal suckers of megalops. Because Russell's description of his 6l-mm ML type (no figure given) bears some resemblance to megalops, I attempted to locate the specimen but with no success. In view of its apparent loss (D. Heppell, Royal Scottish Museum, pers. comm. to M. J. Sweeney, USNM) and the inadequacy of the description, I believe the name should be considered a nomen dubium.

Teuthowenia maculata (Leach, 1817) Figures 4f-j, 5, 8-11

Cranchia maculata Leach, 1817: 140 (type in British Museum (Natural History); examined); 1818a: 410; 1818b: 395;-Tryon, 1879: 162 [listedl;-Hoyle, 1886: 44,186 [listed; type described];- Monod, 1970: 56, pi. 1, fig. 6 [Cranch's previously unpublished journal reference and aquarelle of type; documents collecting date, and type-locality as 0Io36'S, 08°47'El Cranchia (Cranchia) maculata, Prosch, 1847: 71. ? Verrilliteuthis hyperborea, Adam, 1962: 44 (Baia de Luanda, Angola) [description and discussion]. Taonius megalops, Lu and Clarke, 1975b: 382, fig. 9 (11ON, 20oW; specimens from 9 of 10 hauls examined) [vertical distribution]. Teuthowenia megalops, Morales and Guerra, 1977: 300, figs. 2-3 (17°_20oN, 17°-19°W) [description];-Roeleveld, 1977: 129, fig. 5 (16°58'N, 18°16'W; examined) [description]. Teuthowenia sp. C, Clarke (in prep.l) [beaks]. Material Examined.-54 specimens (5-143 mm GL (ML» from 38 lots, including holotype. Holotype.-?s, 32+ mm ML, 01036'S, 08°47'E, 30 V 1816, coli. J. Cranch, BMNH 1983100. Hypotype.-d, 123 mm GL, GERONIMOsta. 2-240, 04°08'S, 10OO8'E,0-'" 1,135 m, 8 IX 1963, USNM 729975. Other Material.-See Appendix. The diagnosis is based on the two largest known individuals, both immature males, the hypotype (123 mm GL), a late juvenile, from the Guinea Basin in the vicinity of the type-locality and a l43-mm GL subadult taken off the Cape Verde Islands in the northern end of the geographic range of the species. Indices for all specimens selected for detailed measurements are summarized in Table 8. Diagnosis.-Size of adults not known. Funnel-mantle fusion cartilages with 1 voss: REVISION OF TEUTHOWENIA 33

d ':":.'.-)1<;>':- ';~0::~ . ; .•..

Figure 8. Teuthowenia maculata. a. Late juvenile male, ventral view, hypotype, USNM 729975, 123 mm GL. b. Dorsal view, of same. c. Left eye, ventral and anterolateral views showing photophores, of same. d. Nuchal fusion cartilage, of same. e. Left funnel-mantle fusion cartilage, of same. f. Funnel organ of same. g. Left tentacular club, USNM 729962, male, 143 mm GL. h. Largest manus sucker, of same. single-point, poorly developed tubercle at mantle margin (Fig. 8e). Glandlike organs on anterior portion of dorsal midline in mature females not known. Fins long (FLI 46+,46+), narrow (FWI approx. 26; hypotype). Eye large (EDI approx. 15; hypotype) (Fig. 8e). 34 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

Figure 9, Teuthowenia maculata. a. Brachial circlet with tentacle and buccal membrane elements of subadult male, right half, USNM 729962,143 mm GL. b. End ofarm II showing sexual modification, of same, c, End of arm I showing sexual modification, of same. d-f. Suckers from arm III, of same: d, basal sucker (from 2nd pair); e, largest sucker (from II th pair); f, terminal sucker (from 19th pair). g, Sucker from end of arm I (from 25th pair) prior to modification, of same.

Arms about one-quarter to somewhat less than one-half length of gladius, with arm length formula III> II> IV=I, or III> II~IV> I (hypotype); on arms III, sucker pairs 8-12 largest with greatest diameter (ASI, elII 3.2, 3.2) about 3 times diameter of basal suckers (Fig. 9a); basal 8-10 pairs of suckers on all arms with voss: REVISION OF TEUTHOWENIA 35

smooth ring margins; distal suckers with about 6-15 low, flat, rounded or truncate teeth (10-15 rounded teeth ..in hypotype) on distal and lateral margins (Fig. 9d- f) except on modified ends of I and II in males; extent of arm modifications in mature males not known. In available immature males, arms I and II with 25- 27 pairs of normal, wide-aperture suckers prior to modified, slightly swollen, terminal portions (5% (hypotype), 8% of length of arms I; 3% (hypotype), 7% of length of arms II); terminal portions of both arms with modified suckers arranged in 3-4 rows, changing to 2 rows prior to short, bare tip on arms I, and in 2 rows, followed by 3-4 rows on the midportion, changing to 2 rows prior to very short, bare tip on arms II; protective membranes reduced (Fig. 9b, c, g). Largest suckers (CISI 1.0 (hypotype), 1.3) of tentacular club with about 22-26 teeth (Fig. 8g, h). Spermatophores, brachial end-organs and mature eggs not known. Gills with about 23-24 lamellae on outer demibranch. Lower beak (143 mm GL) with low wing fold; shoulder ridge developed into distinct, broad-based, triangular tooth (may be broken and appear as low ridge); hood set moderately high above straight crest; distal edge of hood with moderately deep median notch. Upper beak with rostral length of 2.8 mm; jaw angle slightly obtuse (Fig. IDb).Radula with median cusp of rachidian tooth, inner cusp of first lateral, and the single cusps of second and third laterals, long and slender; rachidian and first lateral with broad, shallow base and small, strong, slender outer cusps; small marginal plates poorly developed (Fig. IOc). Gladius (143 mm GL) with vanes bordering somewhat more than posterior half (VLI 57); gladius tail about one-quarter gladius length (GTlLI 23); lanceola with strong dorsal keel (Fig. IDa). Larva-Mantle or gladius lengths, indices and selected characters of young are summarized in Tables 8 and 15. Morphometric relationships with growth of selected characters are shown in Figures 21-24. Forty-four larvae were available. Of the 13 selected for detailed measurements and counts, II are from the northern, and I each from the central and southern portions of the geographic range of the specIes. Larval stage terminates at about 55-60 mm GL. Larva specifically distinguished by dense pattern of small oval chromatophores on mantle (Fig. 4f-h). At about 55-60 mm GL, chromatophores number about 16 on 5-mm square on mediolateral dorsum. Distinctive dense arrangement seen in well-preserved specimens as small as 8 mm ML (and probably smaller). Early larva has noticeably large buccal mass. Funnel-mantle fusion cartilages (Fig. 4i) usually lack single, small tubercle characteristic of older animal. Tubercle sometimes appears in late larval stage. Species additionally characterized by early, disproportionate arm growth, particularly of arms III. As early as about 10-11 mm ML, arms III usually markedly longer than other arms. Disproportionate enlargement of suckers on third quarter of arms III first apparent at about 20-30 mm ML. By late larval stage, largest suckers, usually pairs 8-10, markedly enlarged to mean diameter index of 1.7 (range, 1.3-2.1) (Fig. 4j). Gills of moderate length, with mean length index of about 26 (range, 22-29) in larva of about 10-18 mm ML decreasing to 15 (range, 13-18) in late larva of about 45-55 mm ML. Development of gladius distinguished by early elongation of pseudoconus. Positive allometric growth of pseudoconus usually noticeable after about 20 mm ML. By about 27-42 mm ML, resulting tail of gladius with mean length index of 14 (range, 11-17); vanes comparatively broad and slightly thickened in cross section. Latter 2 features particularly apparent in older larva. Juvenile.-Only 8 juveniles, 55-77 mm GL, were available (excluding 123 mm 36 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

N ,.-, ...J 0011 r---NNf""--C"'iO\t""')- (J Z Nf"'l"')'VN V'l- E E '" ~ o '" II Z

o r-r-'-OI/'lOO-

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~ ~ ....l 8S'SN~:::-V)r--::;;'S~8 o ,.-,-N--N--,.-,-O- E'" '-''-'''-'''------r./) Ell NV)vOOC"'iV"lO'\~OOt"'io;t""') •..• ~z 7'r'r"i'r'r'i'r7'i1'"iiJ) J, OOf""-\.OO\M\O-("t')t"--O\\OOO '" N-- ---~N <::>

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---~ b

.\ I--~ r \ / ./ fj' - - -~ - - ~ ~ VI':'

I .,~ r I , I ·1,. ~ ,------U . -~~(j

Figure 1o, Teuthowenia maculata. a, Gladius, ventral view, with enlarged cross sections and detail of pseudo conus, USNM 729962, subadult male, 143 mm GL. b, Upper mandible, single view, and lower mandible, three views, of same, c. Radula, of same,

GL, late juvenile designated and described above as hypotype); all came from an area between 110 to 19°N and 18° to 22°W in the northern portion of the distribution range. At least in above area, eyes become sessile between about 55-60 mm GL. By 64-77 mm GL, eyes enlarged to mean diameter index of 13 (range, 13-14) and 38 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

3 ocular photophores have separated. Single, small tubercle, typically found on funnel-mantle cartilages oflarger animal developed in only 2 of juveniles, 55 and 70 mm GL. Change in growth of arms abrupt; by 57-77 mm GL, mean length indices show average increase of 31% from late larval stage. Concurrent with disproportionate arm growth, largest suckers on arms III markedly enlarge to mean diameter index of2.2 (range, 1.9-2.6). Rings of suckers smooth on basal portions of arms; incised with small, low, flat, irregular to truncate teeth on mid and distal portions. Ends of arms I and II in largest males of 64 and 77 mm GL lack modification. In contrast to congeners, maculata displays delayed development of male modifications of ends of arms. Mean length index of 14 (range, 13-16) for gills shows little change in proportional length from that in late larva. Posterior end of gladius develops rapidly. At 55-64 mm GL, lanceola fairly delicate with only weak dorsal keel. By 70-77 mm GL, vanes moderately thick, dorsal keel well developed, and pseudoconus stout. Combined features give overall appearance of robustness to posterior end of gladius elongated to mean gladius tail index of 20 (range, 18-20) in juveniles of 55-70 mm GL. Tail exceptionally long (GTlLI 31) in 77-mm GL specimen. Geographic Distribution. - Teuthowenia maculata is restricted to the moderately to highly productive waters of the tropical eastern Atlantic (Fig. 5). All known captures were made over bottom depths in excess of 1,000 m, Though collections of cephalopods from the tropical eastern Atlantic are relatively meager, the 54 specimens of maculata from this study together with the positive-negative capture results of the few cruises that have worked in the area and adjoining waters are sufficient to show that the species has its northern limit of distribution, east of about 200W, at about 200_22°N, in the vicinity of the frontal meeting of the subtropical North Atlantic Central Water and the tropical South Atlantic Central Water. The northernmost station at which the species was taken by the DANA along the Cape Town-Denmark transect during which the successful catching gear were used at about 1°_5° intervals, was at 18°N, 18°W. In a series of 6 stations made between 20° to 25°W at 7°_17° intervals from lION to 600N, Clarke and Lu (1974; 1975) and Lu and Clarke (1975a; 1975b) report (under the name Taonius megalops) the species only at lIoN, 200W. This station was occupied twice, once in February 1968 and again in March 1972; successful captures were made on both visits. The majority of their specimens are included here. The absence of the species in the two collections made in November 1969 and February 1972 at the station at 18°N, 25°W to the west of the Mauritanian Upwelling region suggests that maculata is confined in the northern end of its range to the physically distinct area of the upwelling. Krefft (1974) observed a distinct change in the midwater fish fauna at about 200N. Various midwater fishes (Krefft, 1974; Backus et aI., 1977; Hulley, 1981) have one of their limits at about this latitude in the eastern Atlantic. Additionally, Van Soest (1979) considered 200N to be one of the major N-S boundaries for epipelagic organisms. Unfortunately, material is insufficient to clearly define the southern boundary for maculata. The DANA,working northward from Cape Town first encountered the species at 80S, 9°W and subsequently made captures at 6 out of 8 stations (75%) to 18°N, 18°W. The DISCOVERY,however, fishing northward in the same latitudes but in the easterly more highly productive waters off the coast made its first capture at 19°5, II°E, followed by a second at the subsequent station at 16°S, II°E. Along the southern half of the Cape Town-Madeira transect of the 1971 voss: REVISION OF TEUTHOWENIA 39

cruise, the WALTHERHERWIGfished the less productive waters to the west of those fished by the DANAand DISCOVERYand caught no specimens. These results suggest that the distribution of maculata is strongly influenced by the productivity of the water. In the more inshore highly productive waters, the southern boundary of distribution may occur at about 20OS.The species may extend only to about 100S in the moderately productive waters to the west and be absent in the low productive waters furthest offshore. Some myctophids also have their southern limits in this area between about 10° and 21oS, the region where the Benguela Current turns westward (Hulley, 1981). Nafpaktitis et al. (1977) found two eastern tropical Atlantic species of myctophids, Diaphus vanhoeffeni and Lampanyctus isaacsi, that displayed the same N-S distributional limits as maculata. Van Soest (1979) considered 200S to be the site of one of the major N-S hydrographic barriers to the distribution ofepipelagic organisms and regarded it as the southern counterpart of the barrier at 200N. Krefft (1974), however, did not find this to be the situation with midwater fishes, where both tropical and subtropical species tend to find their southern limit at the Subtropical Convergence. Roper (unpubl. pers. journal3), observing the collections of cephalopods taken along the Cape Town-Madeira transect of the 1971 cruise of the WALTHERHERWIG,noted a faunal change at about 34°-35°S, and by 24°S the collections were clearly composed of tropical speCIes. The eastern limits for maculata are less distinct, and, again, appear to reflect the water productivity. The species is not known from west of23°W in the northern half of its range, and 100W in the southern half. Distribution appears to be continuous over the range between 200N and 200S, with the greatest abundance between 10° and 200N, in the region of the Mauri- tanian Upwelling. The lack of captures by the PILLSBURYon either the May-June 1964 or May 1965 cruises to the Gulf of Guinea (G. Voss, 1969), during which 34 IKMT hauls were made in an area between lOSto 6°N and 3°W to.9°E over bottom depths greater than 1,000 m, in the vicinity of the type-locality (01°36'S, 08°47'E), indicates that the species is scarce in these waters where the year-round productivity is less than to the north or south. Similarly, lower productivity correlates with the only 2 stations, mentioned above, at which the DANAfailed to capture the species between 80S and 18°N. The negative-capture stations occurred immediately to the west of the gulf, between 0° to 4°N and II ° to l3°W. Since maculata is only known from the larval to subadult stages, it cannot be exactly determined whether breeding occurs over its range, but the captures of early larvae of 5-7 mm ML in the northern, central and southern sectors suggest that there is a breeding population throughout the known range with the possible exception of the vicinity of the Gulf of Guinea. Other captures not included in this study but probably identifiable to the species are reported by Adam (1962) (under the name Verrilliteuthis hyperborea), of a juvenile from about 09°S, l3°E, and by Morales and Guerra (1977), of 5 larvae taken between 17° to 200N and 17° to 19°W. Additionally, the undescribed 26- mm ML larva taken in the Gulf of Guinea at 02°35'N, 03°48'W mentioned by Nesis (1974a: 11) is probably also of this species. In brief, Teuthowenia maculata is found in the northern and southern provinces of the Mauritanian Upwelling Region, and the Guinean Province of the Atlantic Tropical Region, with distribution in the Guinean Province influenced by productivity. Similar distributional patterns which closely relate to land affected

'c. F. E. Roper. Personal journal from the March/April 1971 cruise of the WALTHER HERWIG (unpubl. manuscriPt). 40 BULLETIN OF MARINE SCIENCE. VOL. 36. NO.1, 1985 o

10 Number of [ specimens L 0

Night Twilight = 400 Day = 600 - 800

.§. 1000 ~ = -Q. Q) 1200 Cl

1400

1600

1800 =

2000

o 20 40 60 80 100 120 140 160 Gladius length (mm)

Figure II. Teuthowenia maculata. Vertical distribution with respect to time of capture. All specimens are immature. environments have been reported for some midwater fishes by Baird (1971) and Nafpaktitis et al. (1977) among others, and are referred to as pseudoceanic by Krefft (1974). Teuthowenia maculata may be best described as an eastern-tropical, pseudo-oceanic species. Vertical Distribution. - Vertical distribution data for 52 specimens are summarized in Figure 11. Teuthowenia maculata, ranging in size from larva to early subadult 5-143 mm GL, has been taken by nets fishing in the upper 25 m to about 2,000 m depth. Thirteen of the 17 specimens caught by the DISCOVERY with closing nets and reported (under the name Taonius mega lops) by Lu and voss: REVISION OF TEUTHOWENIA 41

Clarke (1975b) were examined and included in the study. The remaining 4 specimens which could not be located are also probably this species. The captures were made over a temperature range in the vertical column of 23°-7°C (0-700 m). The closing-net captures suggest that larvae of 6-20 mm ML are concentrated, both day and night, in the upper 100 m. The daytime capture of 2 larvae of 10 and 16 mm ML in the upper 25 m indicates that the distribution extends to the near-surface layer. The data also show that at about 40 mm ML, the larva descends to depths of about 300-600 m. Juveniles of 50-60 mm ML, were taken during the day and night in depths of 600-700 m. The latter are the largest specimens taken and the deepest captures made of maculata by the DISCOVERYusing a closing rectangular midwater trawl (RMT 8 and combination net). The captures made with nonclosing nets over the same depth range generally describe a similar distribution pattern. The nighttime capture of a 70-mm GL specimen by the AT- LANTISII fishing at 115-125 m indicates that juveniles are found, at least occasionally in the shallower waters at night. The nighttime presence of early subadults in the approximate upper 600 m is indicated by the capture of the 143-mm specimen by the WALTHERHERWIGfishing at 596-610 m. The captures oflarvae under 10 mm ML from below 100 m, and the various captures from 1,000 m and below shown on the graph were all made with nonclosing nets and are regarded as possible contaminants from shallower depths. The occurrence of ontogenetic descent at a smaller size in maculata as compared with megalops, and noted by Lu and Clarke (1975b), correlates with the abbre- viated larval stage found to occur in the former species. Remarks. - Leach's Cranchia maculata has been regarded as a doubtful species nearly from the time of its proposal. The type-specimen was known only as a Cranchia-like squid having a mantle with a smooth surface and mottled with discrete, oval chromatophores until Hoyle (1886), on reexamination of the specimen, described the mantle as totally devoid of head and arms but with the funnel attached. He further described the nuchal and funnel-mantle attachments and gave the spacing of the chromatophores as about 2 mm apart and the mantle length as about 3 em. Monod's (1970) publication of the expedition journal and original aquarelles of John Cranch gave the collecting date for the type as 30 May 1816 and the locality as 01036'S, 08°47'£, and provided the original figure which showed the densely spaced chromatophore pattern and the small terminal fins. Though over the years, the type-specimen has virtually been dissected to pieces, I found, on reexamination, the additional features of a diamond-shaped lanceola with small oval, separate fins inserted on the posterolateral margins and a small, smooth, spindle-shaped funnel-mantle cartilage. The combined characters viewed with an overall knowledge of the family confirm the identity of Leach's species maculata to the series of larva through early subadult from the tropical eastern Atlantic I described above. The present measurement of the type-specimen is 32± mm ML.

Teuthowenia pellucida (Chun, 1910) Figures 4k, 12-20

Desmoleulhis pellucida Chun, 1910: 357, pI. 53, fig. 1, pI. 54, figs. 1-17 (33°20'5, 15°5S'E; type in Zoological Museum, Humboldt University, Berlin; examined). Megalocranchiapellucida. Berry, 1912: 644 [Iistedl;-Pfeffer, 1912: 716 [type described];-Berry, 1916: 64 [listed]. Megalocranchia pardus Berry, 1916: 61, pI. 9, fig. 2 (Sunday Island, Kermadec Group; type in Berry's private collection, Redlands, California; examined);-AlIan, 1945: 336, pI. 27, figs. 1-4 42 BULLETINOF MARINESCIENCE,VOL.36, NO.1, 1985

(off Port Hacking, N.S.W., Australia; 43'13'S, 148'I7'E) [description];-Dell, 1952: 135, pis. 31- 32 (Island Bay, New Zealand) [description];-Clarke, 1966: 231 [listed]. Anomalocranchia impennis Robson, 1924a: 8 (Cape Town, South Africa; type in British Museum (Natural History); examined); 1924b: 628, pI. I, fig. 3, text-figs. 17-20 [type figured; internal features of type described]; - Thiele, 1934: 979 [listed]; - Clarke, 1966: 232 [listed]. Megalocranchia richardsoni Dell, 1959: 10, figs. 7-10 (37'-43"S, 174'-1 78'E; types in Dominion Museum, Wellington; paratypes VUZ 57 and VUZ 110 examined). Taonius pellucida, Clarke, 1966: 236 (part: not Chun, 1913; Grieg, 1920; 1924) [listed]. Taonius richardsoni, Clarke, 1966: 237 [listed]. Megalocranchia megalops australis Voss, 1967: 82, pI. 8, figs. a-e, pI. 9, figs. a-f(31'-39"S, 33'- 45'E; types in South African Museum; examined). Verrilliteuthis megalops australis, Nesis, 1974a: 10, fig. 2, upper left (44'-45"S, 55'-56'W; 41°-44"S, 26°-27°W); 1974b: 66 [distribution]. Verrilliteuthis richardsoni, Nesis, 1974a: 10, fig. 2, lower left (36'53'S, 17'31 'E). Vossoteuthis pardus, Nesis, 1974a: 13 [listed]. Teuthowenia megalops impennis, Imber, 1978: 471, fig. IU [part: (?) fig. 6D; not synonymy citations Allan, 1940: 321; 1945: 337; Adam, 1962: 44. (7) Massy, 1916: 170; Imber, 1973: 650; 1976: 125; Imber and Russ, 1975: 31). Fusocranchia pellucida, Imber, 1978: 474 [part: (7) fig. IV; not synonymy citations Joubin, 1920: 73; Clarke, 1966: 232; Sasaki, 1929: 326; Clarke, 1966: 230; Nesis, I974a: 16; Dilly, 1972: 403; Lu and Clarke, 1975a: 150; 1975b: 382; Dilly and Nixon, 1976: 19; (?) Imber, 1976: 125]. Teuthowenia sp. B, Clarke (in prep. ') [beaks]. Material Examined.-235 specimens (4-201 mm GL (ML» from 101 lots including holotype. Holotype.-9, 77 mm ML, German Deepsea Expedition Sta. 90, 33'20'S, 15'58'E, 0-1,000 m, 2MB. Hypotype.-9, 140 mm GL, WALTHERHERWIGSta. 358 II/7I, 39'47'S, 43'38'W, 200-212 m, 7 III 1971, USNM 816160. Other Types.-?s, 50 mm ML [70± GL] Sunday Island, Kermadecs, beach, 1910, SSB 415 (holotype of Megalocranchia pardus Berry).-7s, 57 ± 3 mm ML, Sta. 540, off Cape Town, 765 m, coll. J. Gilchrist, BMNH 1924.9.9.38 (holotype of Anomalocranchia impennis Robson).-I .s, 130 mm ML [145± mm GL], VUZ 110, Cook Strait, "Middle ground," "'128 m, fish stomach, colI. R. Greco, DMNZ M12943; I .s, 86 mm ML [100+ mm GL], VUZ 57, 42'02'S, I74°50'E, "'549 m, DMNZ MI2941 (paratypes of Megalocranchia richardsoni Dell).-.s, 89 mm ML [109 mm GL], IKMT 40, 38'52'S, 33'90'E, 500 m, 13-14 XI 1962, SAM A29733 (holotype of Megalocranchia megalops australis G. Voss).-2 ?s, 39,29 mm ML, IKMT 32, 31'41'S, 44'35'E, 500 m, 15 VIII 1962, stomach of Alepisaurus ferox, SAM A29714; I ?s, 24 mm ML, IKMT 31, 32°30'S, 35'08'E, 500 m, 12 VIII 1962, SAM A29723 (paratypes of Megalocranchia megalops australis G. Voss). Other Material.-See Appendix.

The following diagnosis is based on 28 adults and subadults (128-201 mm GL), including the hypotype (140 mm GL), from over a wide part of the geographic distribution of the species. Twenty-four specimens came from across the broad range in the South Atlantic, 2 from the East Australian Current and 1 each from the southwestern Indian Ocean and the western South Pacific. Indices for all specimens selected for detailed measurements are summarized in Table 9. Diagnosis.-Medium-sized species (both males and females mature at about 141- 201 mm GL). Funnel-mantlejusion cartilages with multipoint tubercle at mantle margin (usually with 2-4 points, occasionally 1 or 5) (Fig. 12e). Anterior portion of middorsal line of mantle in mature and near-mature females with row of about 6-15 small, glandlike organs with 1 (occasionally 2-3) central pore(s) (Figs. 14d, e; 17). Fins long (FLI approx. 33-50; 43 ±, hypotype), narrow (FWI approx. 19- 27; 27, hypotype). Eye large (EDI 14-18) (Fig. 12c). Arms with length formula III> II> IV~I, except in adult males in which it is III>II>I>IV; on arms III, sucker pairs 10-14 usually largest, with greatest diameter (ASI, eIII 2.0-2.3; 2.1, hypotype) about two and a halftimes diameter of basal suckers (Fig. 13h); chitinous rings of basal and midarm suckers smooth to voss: REVISION OF TEUTHOWENIA 43

~tl·\ t? 1 a !A 'i 'I ,1

d ~. -. -"'.-'.--,.~- .~.,_., "P_·r:'~·· ,. ~ ~ >, I:

/ ' I \ \ e \

- - \ . -. . .:~ :.,~ .-•. I - -, ,I ~':~~, I

I I I' I \ I \ \ \ ~~-~.r-"._~~ ' •.._ •. J \'.~\~~J~

Figure 12. Teuthoweniapellucida. a. Subadult female, ventral view, hypotype, USNM 816160,140 mm GL. b. Dorsal view, of same. c. Left eye, ventral and anterolateral views showing photophores, USNM 816193, male, 138 mm GL. d. Nuchal fusion cartilage, hypotype. e. Right funnel-mantle fusion cartilage, of same. f. Funnel organ, USNM 816191, female, 20 I mm GL. g. Left tentacular club, USNM 816193, female, 143 mm GL. h. Largest manus sucker, of same. irregular; small distal suckers with about 6-14 small, flat, irregular, round or truncate teeth on distal and lateral margins (Fig. 13i, j) except on modified ends of arms I and II in males. In mature males, all arms, especially arms IV, with increased robustness; suckers on basal portions with swollen, fleshy collars and 44 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.1, 1985

~"~"~)'"," { ~ l ~ ••~••• -j'

d .J g

b ,"

,.. " \ , , .~, ~."

, I I I I I I I I I /

\ \ \ I

Figure 13, Teuthowenia pellucida. a. End ofleft arm IV of near-mature female showing brachial end- organ, USNM 816199, 176 mm GL. b. End of left arm II of immature male showing sexual modification, USNM 816193, 127 mm GL. c. Sucker removed from surface ofnidamental gland of near- mature female, USNM 816198, 179 mm GL. d. Sucker (from 17th pair) from sexually modified end of left arm I of immature male, USNM 816193, 138 mm GL. e, Entire left arm II of mature male showing sexual modifications, USNM 816198, 201 mm GL. f, g. Suckers from same left arm II: f, modified basal sucker (from 2nd pair), side and oral views; g, next to last sucker (from 12th pair) prior to modified end. h. Brachial circlet with tentacle and buccal membrane elements of subadult female, left half, hypotype. i, j. Suckers from left arm III, of same: i, largest sucker (from 12th pair); j, sucker (from 19th pair) from end of arm, voss: REVISION OF TEUTHOWENIA 45

rings with reduced apertures (Fig. 13f); arms I and II with 11-18 pairs of normal suckers prior to elongate, often swollen, modified terminal portions (33+-44+% of arm length). Terminal portions with abruptly reduced, modified suckers (present in near-mature, but missing in available mature specimens) arranged in 2 rows, changing to 3-4 rows on midportions, followed by 2 rows prior to bare whiplike tip; sucker pedestals swollen; protective membranes reduced (Fig. 13b, d, e). In mature and near-mature females, arms IV (only arms intact in availab]e specimens) with 17-21 pairs of suckers prior to terminal 13-] 9% of arm modified to form brachial end-organ. Brachial end-organ with entire oral surface rugulose; small, biserial suckers retained to tip; expanded, trabeculate protective membranes fleshy and heavily pigmented together with enveloped oral surface (Fig. 13a). Small, glandular processes not present between largest suckers on arms. Tentacular club (in specimens of 128-154 mm GL; missing in larger animals) with poorly defined carpal group; largest manus suckers (CISI 0.9-1.]; 0.9, hypotype) with about 26-32 teeth (Fig. 12g, h). Spermatophore (5 spermatophores from 2 specimens, 16],201 mm GL) small (SpL 12.3-17.7 mm; SpLi 8-9), with long sperm mass (SpMI 46-52), long cement body (CBI 34-40), and short ejaculatory apparatus (EjAI 12-14) (Fig. 14g; Table 10). Mature eggs small (2.2 x 1.6-3.0 x 2.2 mm); total egg count (2 specimens, 177, 187 mm GL) 7,900±, 6,300±. Gills with 18-22 lamellae on outer demibranch. Lower beak with rostral length of 4.3, 4.9 mm in mature males (2 specimens, 186,20] mm GL), 4.9-5.5 mm in mature and near-mature females (3 specimens, 176-184 mm GL); wing fold prominent; shoulder ridge low, sometimes developed into low, broad-based, triangular tooth; hood set low above straight or slightly curved crest; distal edge of hood with shallow median notch. Upper beak with rostra] length of 4.3 mm in mature male (186 mm GL), 4.8-5.2 mm in mature females (3 specimens, 176-184 mm GL); jaw angle usually obtuse, sometimes slightly recessed (Fig. 14b; Table 11). Radula with median cusp of rachidian tooth, inner cusp of first latera], and single cusps of second and third laterals moderately long and stout; rachidian and first lateral with broad, stout base and small, stout outer cusps; small marginal plates distinct (Fig. 14c). Gladius (4 specimens, ]6]- 20] mm GL) with vanes bordering somewhat more than posterior third (VLI 36- 40); gladius tail slightly less than one-quarter g]adius length (GT]LI 20-22); ]an- ceola with moderately developed dorsa] keel (Fig. ]4a; Tab]e ]2). Coloration. - In adults, thin epidermis on all surfaces of mantle, fins, funnel, neck, arms and tentacles with numerous, small reddish-brown chromatophores which give finely mottled, dark reddish-brown coloration to entire animal. Outer surfaces of bucca] membrane and bucca] mass similarly pigmented, but to lesser extent. Often head and aboral surfaces of arms appear more darkly colored; darker, median dorsa] line usually present on mantle overlying free rachis. Darkly pigmented outer layer nearly intact in all available mature and near-mature females but only fragments remain, or totally missing, in all adult males (may be related to change in texture of body in females as they near maturity). Under thin outer layer, second layer (more commonly retained in collected and preserved animals) bears fewer and larger reddish-brown chromatophores. Chro- matophore pattern of second layer commonly recognized as characteristic of species: numerous, regularly spaced, discrete, reddish-brown chromatophores over entire mantle; median dorsa] row over free rachis; and often row on dorsolateral margins of mantle along lines of fin insertions. Additionally, large chromatophores occur on ventral surface of head and dorsa] surface of funnel, with larger number of 46 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

~~ NV '0 '"II .•. Z \0'-"'-".,.., '0 " "IV ..J Z I 1 o -N 5 ..J Nv 5 o .•. 5 N .•.5 J.. .-.. .-.. o o N N~oo~oo6'~NS~~ I t'f"')("'>.l-t'lNNN"l:t-O- on '" '-"'-'------tnl.r\O-'f"'----~\o-~l,r'JCJ'j '" '" II Z 0- 11'711111'71''7OO~r---NLnO'.OOOO\""'('f"'l NN-NN---:~ 0- '" z"

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~\OM.t';;';;~"'lOM~"""CI) ooO1111117'1''7'7'7•.....-MO •.....No-oo.,.., .,; ..J MN-NNN"':v 0- c o '" ...o 5 c 5 .g .•. :,; c J o N o"~ o 0.

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---~ I' " !II'

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Figure 14. Teuthowenia pellucida. a. Gladius, ventral view, with enlarged cross sections and detail of pseudo conus USNM 816196, mature female, 177 mm GL. b. Upper mandible, single view, and lower mandible, three views, of same. c. Radula, of same. d. Anterior portion of mantle of mature female, dorsal view, showing maturity-related, g1andlike organs on middorsal line, embedded sperm reservoirs and arm sucker scars, USNM 816191, 201 mm GL. e. Enlargement ofa single g1andlike organ, of same. f. Posterior end of nidamental gland of near-mature female showing encysted sucker rings, USNM 816198, 179 mm GL. g. Spermatophore with detail of oral end showing ejaculatory apparatus, USNM 816194, 161 mm GL. 48 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, J 985

Table 10. Teuthowenia pellucida. Spermatophore lengths and indices

USNM 816198 USNM 816194 Range GL (mm) 201 161 Number of sperma tophores 2 3 SpL (mm) 17.5,17.7 12.3-14.5 12.3-17.7 14.9 SpLI 7.7,8.8 7.6-9.0 7.6-9.0 8.4 SpWI 2.3,2.3 3.2-3.5 2.3-3.5 2.9 SpMI 46.3,48.0 45.6-51.7 45.6-51.7 47.8 CBI 40,0,36.7 34.5-38.4 34.5-40.0 37.4 EjAI 12.4, 12.6 12.4-13.6 12.4-13.6 12.8

Table 11. Teuthowenia pellucida. Beak measurements, indices and ratios

UMML USNM USNM USNM USNM USNM 31.1850 816196 816199 816198 816199 816181 Range

Sex 2M 2M 20M oM oM 0* GL(mm) 184 177 176 201 186 119 URL(mm) 5.2 5.0 4.8 4.3 2.0 LRL (mm) 5.5 5.0 4.9 4.9 4.3 2.0 LRLI 3.0 2.8 2.8 2.4 2.3 1.7 2.3-3.0 2.7 WiL/LREL 2.0 2.4 2.1 2.0 2.5 2.5 2.0-2.5 2.2 LCrULHoL 1.9 1.9 1.8 2.0 1.9 1.9 1.8-2.0 1.9 • Immature; n01 included in ranges and means.

smaller chromatophores on dorsal surface of head, and row of large chromatophores extending length of arms and tentacles, often giving banded effect on tentacles. Larva.-Mantle, or gladius lengths, indices and selected characters are summarized in Tables 9 and 15. Morphometric relationships of selected characters with growth are shown in Figures 21-24. Sixty-five larvae were available from over the full circumglobal range of the species. Because of the relatively poor condition of the specimens from the western Atlantic and Indian Oceans, these areas are not represented by the 17 larvae selected for detailed measurements and counts. End oflarval stage occurs between about 53 and 95 mm GL. Larva characterized by moderately dense pattern of medium size, oval chromatophores on mantle (Fig. 15a-d), about 8 on 5-mm square on mediolateral dorsum of a 55-60-mm GL animal. Characteristic pattern seen in specimens as small as 5 mm ML.

Tab]e 12. Teuthowenia pellucida. G]adius indices

USNM USNM USNM USNM 816196 816199 816198 816194 Range x Sex 9M 9nM dM dM GL(mm) 177 176 210 16] VLI 37 36 40 40 36-40 38 GWI 9 9 9 ]1 9-11 10 GTILI 2] 22 20 21 20-22 21 FrRaWI, ant 0.9 0.9 0.9 0.9 0.9--0.9 0.9 FrRaWI, mid 0.7 0.5 0.6 0.6 0.5--0.7 0.6 voss: REVISIONOF TEUTHOWEN1A 49

n' c'

, ·1" .~.) \, ..

Figure 15. Teuthowenia pellucida. a-e. Developmental series of young. a-c, e. Ventral views. d. Dorsal view. a'-e'. Posterior ends of mantle, dorsal views, showing development with growth of the fins and the posterior end of the gladius, of same. a, a'. USNM 816170, 7 mm ML. b, b'. USNM 816173, 10 mm ML. c, c'. USNM 816184, 27 mm ML. d, d'. SAM A4981 WS-3, 57 mm ML. e, e'. SAM A31421, 85 mm GL. f. Tentacle of larva, USNM 816173,10 mm ML. g. Posterior end of gladius of larva, "c" stage, showing early formation of pseudoconus, DISCOVERY968, 32 mm GL. (Chromatophores bleached in a, b; on mantle, shown contracted in c, e; expanded in d.) 50 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

a "f.

~\l~:'~>' .;. . . --,~~'>~,I"'" b

,..,.;.,~-.

d {'''':'-;?'. ~;

Figure 16. Teuthowenia pellucida. a-c. Developmental series offunnel-mantle fusion cartilages show- ingtuberc1es. a. USNM 816173,10 mm ML. b. USNM 816184, 27 mm ML. c. DMNZ, 67 mm GL. d-f. Developmental series offunnel organ. d. SAM A4217, 12 mm ML. e. USNM 816184, 27 mm ML. f. AM C109363, 61 mm GL. g-j. Developmental series of eye, anterolateral and ventral views of left eye, showing photophores. Scales = I mm. g. USNM 816170,7 mm ML. h. USNM 816184, 27 mm ML. i. DMNZ, 67 mm GL.j. SAM A31421, 85 mm GL.

Characteristic multipoint tubercle on funnel-mantle fusion cartilages first appears at about 10-20 mm ML as small, single tubercle; by about 25-30 mm ML second and often third tubercle usually appear at base of first. Additional I or 2 tubercles may develop in older larva (Fig. 16a-c). In later growth stages individual voss: REVISION OF TEUTHOWENIA 51

Table 13. Teuthowenia pellucida. Comparison of indices of selected characters showing morphological differences between early-metamorphosed juveniles and late-maturing larvae at the same approximate gladius length

Early~melamorphoscd juveniles Latc-maturing larvae (83-95 mm GL) (82-95 mm GL) N ~ 4 N=4

Index Range Range ALI, I 14-19 17 11-16 13 II 16-23 21 14-19 17 III 23-28 27 18-25 21 IV 18-21 20 13-15 IS ASI, eIIl 1.8-2.1 1.9 1.3-1.7 1.5 EDl 13-14 14 9-11 10

tubercles usually fuse basally to form multipoint tubercle. Number of tubercles often different on two cartilages of individual, e.g., 1/3,2/3,3/4,3/5, etc. Cartilage tubercle rarely lacking in specimens larger than about 30 mm ML. Over broad geographic range of species, relative growth of arms in early larva varies widely. Growth of arms III slightly disproportionate. Basal suckers of arms small with small apertures. By 53-63 mm GL, disproportionate enlargement of suckers on third quarters of arms III moderate, with largest suckers having mean diameter index of 1.0 (range, 0.8-1.2) (Fig. 4k). When larvae reach sizes of 72- 95 mm GL, as in some areas of distribution, mean diameter of enlarged suckers increases to about 1.4 (range, 1.3-1.7), more markedly disproportionate. Exact sucker pairs that show maximum enlargement varies from 8-9 to 12-13, depending on area of origin of larva. Gills small, with mean length index of 17 (range, 16-18) in larvae of 7-24 mm ML, decreasing to 11 (range, 10-12) in mid to late larvae of 57-63 mm GL. Relative length of pseudoconus of gladius in early stages of larval development varies considerably over distribution range. Juvenile. -Juveniles were available from throughout the circumglobal range of the species except for the central and eastern sectors of the Indian Ocean. Over the broad range of distribution, the size at which the eyes become sessile and the juvenile stage commences varies widely from about 53 to 95 mm GL. Because metamorphosis signals change in allometric growth of various parts of body, proportional size of eyes and length of arms usually noticeably greater in small metamorphosed juvenile in lower end of above size range than in specimen in upper end of range that is still in larval stage. Lesser disproportionate growth is displayed by suckers on third quarters of arms III. Proportional differences in eyes, arms and enlarged suckers marked when small metamorphosed juvenile and large maturing larva compared at same size in upper end of size range (Table 13). Metamorphosis initiates development of male modifications simultaneously on ends of arms I and II. As result of variations in size at metamorphosis over wide range of distribution, number of normal sucker pairs prior to modifications varies from about 11 to 20. Similarly, degree of development of modifications (i.e., proportional length of arm) at given gladius length varies widely; juvenile males of about 95-109 mm GL with about 6 to 22% of arms modified depending on geographic area of origin. Beaks of juvenile differ from those of subadult and adult in degree of development of various features: on lower beak (2 specimens, 119 and 70 mm GL, 52 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985 from Tasman Sea), rostrum lacks hooked tip of adult; wing fold poorly developed; hood set slightly higher on crest than in adult; broad fold on lateral wall low in larger juvenile, and nearly undetectable in smaller one; and dark coloration confined to rostrum and anterior half (70 mm GL specimen) to three-quarters (119- mm GL specimen) of hood. In smaller juvenile, light coloration extends along anterior half of exposed crest; in larger juvenile, area of coloration extended to include anterior half of exposed lateral walls. Wings colorless. Proportional length of gills (GiLl 11-12) shows no change from that of late larva. Vanes of gladius of juveniles of 75-105 mm GL moderately thickened; dorsal keel of lanceola weakly developed. Proportional length of gladius tail at given gladius length varies widely over range of species; indices of 13-21 occur in juveniles of 53 to 90 mm GL (maximum elongation of posterior end of gladius found in 75-mm GL specimen from Peru Current). Maturity. - The female undergoes conspicuous changes at maturity, with the changes occurring over an apparent short time span near the maximum size of the individual. At maturity, the female is distended with approximately uniform size, oval eggs (2.2 x 1.6-3.0 x 2.2 mm). The total number of eggs was estimated to be about 6,000-8,000 in 2 mature females of 187 and 177 mm GL. In the mature female, the swollen oviducal and nidamental glands are approximately coequally enlarged; the ends of arms IV (only arms intact) and probably the ends of all the arms are modified into darkly pigmented brachial end-organs (Fig. 13a); and a series of small, glandlike organs occurs on the mantle along the midline of the dorsum from the nuchal cartilage for about 7-16% of the mantle length (Fig. 14e, d). Additionally, the firm texture of the entire body is lost, giving the animal a general flaccid appearance. Sections of the mantle show an overall deterioration of tissue. The outermost skin layer and a thin layer of dense connective tissue is loosely attached to an underlying more gelatinous layer of connective tissue. Sections of the glandlike organs on the middorsum (Fig. 17) show them to be comprised of spongy tissue with 1 or more, complex epithelial-lined chambers which open to the exterior through a simple or branched, hollow papilla, or 1 (occasionally 2-3) central pore(s) surrounded by a circlet of solid papillae. No gland cells were found in the linings of any of the cavities. While some of the chambers are empty, others contain masses of an undefined nature. Of the 10 mature females available, the smallest measured 147 mm and the largest 201 mm GL. Three females of 176, 179 and 189 mm GL were judged to be near-mature: the brachial end-organs are deeply pigmented; the organs on the middorsal line of the mantle are developed; the egg mass is large, with the eggs moderately enlarged (1.2 x 1.0-1.3 x 1.2 mm); and the sex glands are moderately enlarged (NGILI 12-13 compared with 14-21 in mature females). Three specimens, 153, 154, and 154 mm GL, exceed the size of the smallest mature specimen of 147 mm GL but are still in the immature stage: brachial end-organs and the organs on the dorsum of the mantle are not developed; the sex glands are small (NGILI 1-2); the egg mass shows moderate enlargement; and the eggs are very small. Of 9 mature females available for detailed examination, 5 have 5-27 sperm reservoirs embedded in the external surface of the anterior half of the dorsum of the mantle (Fig. 14d), and 1specimen has 24 reservoirs embedded on both dorsum and ventrum. A number of the sperm reservoirs have penetrated the inner wall and their open, funnel-shaped ends project into the internal chambers. The remaining 3 mature and the 3 near-mature females contain no sperm reservoirs. All but 2 of the mature and 1 of the near-mature females have numerous detached voss: REVISION OF TEUTHOWENIA 53

nuchal cartilage

O.5mm

O.25mm

Figure 17. Teuthowenia pellucida. Transverse section of one of the g1andlike organs which occur along the midline of the anterodorsal surface of the mantle in mature and near-mature females, USNM 816200, 189 mm GL. • suckers with reduced-aperture rings encysted internally on the walls, usually of both the dorsal and ventral chambers, and on the exposed internal organs of the anterior half of the mantle, especially on the swollen sex glands (Fig. 14f). All of the sucker rings have uniform, minute, sharply-pointed teeth on the entire margins 54 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

(Fig. 13c). The 2 females that lacked these encysted suckers did have sperm reservoirs embedded in the mantle. The male displays marked, though somewhat less conspicuous, changes at maturity. The mantle remains relatively narrow, and there is no apparent deterioration of the texture of the body. In the mature male, the sex organs are greatly enlarged; the penis, whose muscular walls taper to a slender tip, is elongated and extends as much as 25 mm out of the mantle opening; and completely formed spermatophores are found in the spermatophore sac and/or penis. The number of spermatophores found in an individual was never large, ranging from 3 to about 10, often with 1 or 2 incompletely formed in the spermatophoric glands. The spermatophores varied in length within an individual with those taken from the penis in a 161-mm GL male being smaller than one taken from the spermatophore sac indicating an increase in size of the spermatophore with continuing growth of the individual. Increased size of the spermatophore with growth of the animal is further demonstrated by the spermatophores taken from a 20 1-mm GL male which were about 1/3 longer than those from the 161-mm GL male (see Table 10). In both specimens, the spermatophores display no imperfections and the proportional size of the component parts is similar. Another feature of male maturity in pellucida is the elongation of the modified ends of arms I and II to about 31-46% of the arm length. On the often swollen, modified ends, the specialized suckers that have reduced-aperture rings which are finely incised on all margins (found in all of the near-mature males) are often missing in the mature males, and only the swollen pedestals remain (Fig. 13b, d, e). Additionally at maturity, all of the arms, particularly arms IV, show increased robustness and the suckers on the basal portions have developed swollen, fleshy collars and rings with reduced apertures (Fig. 13f). The suckers on the basal portions that remain on the specimens display no substantial changes in dentition. The available 6 mature males (including 1 probable; sex organs missing) ranged in size from 141 to 201 mm GL. The fact that all available males over 141 mm GL are mature could be misleading, for the next largest males of 138, 128 and 127 mm GL have small, immature sex organs, show no increased robustness of the arms, nor modifications of the basal suckers. The modified ends of arms I and II of these subadults, however, have elongated to 15-23% of the arm length, but are neither swollen nor have the naked tips attenuated. It appears that the above described, male, maturity-related changes, with the exception of the multiple rows of modified suckers on the ends of arms I and II, take place over a relatively short period close to the maturity of the individual. The data are inadequate to determine if there is a limited breeding season or not, though the catches of mature animals in January and March in the Atlantic and May in the Indian and Pacific Oceans, and oflarvae of 5-10 mm ML in the Pacific in August to December suggest an extended maturity and spawning period for the species, perhaps year-round.

Observations on Possible Behavior During Mating and Spawning. - In the absence of observations of live animals, I can only speculate as to the probable functions of the maturity-related, morphological changes described above, and attempt to integrate them into a possible behavior pattern for courtship and copulation in pellucida. The various features suggest that the female may attract a mating partner using the bioluminescence of the brachial end-organs (R. Young, 1975a) and, perhaps, (a) pheromone(s) produced by the glandlike organs on the dorsum ofthe mantle. The bioluminescence would be effective at relatively long range (~50- voss: REVISION OF TEUTHOWENIA 55

100 m depending on water clarity, etc. (R. E. Young, pers. comm.» to cause the male to approach while the pheromone(s) would be optimally effective at short range to stimulate subsequent courtship and copulatory behavior. The attracted and stimulated male, using the more robust fourth arms and the basal halves of some or all of the other arms, may grasp the mantle of the female, insert the attenuated, modified ends of arms I and II into the mantle chambers and proceed to stroke the inner walls and swollen sex glands of the female. The small, specialized suckers on the modified ends of the arms of the male adhere to the surfaces of the inner linings and organs of the female, subsequently detach from the arms and become encysted. The encysted suckers may afford further stimulation for egg laying. If the female is ready to copulate, the mating process may proceed. The possible use described above of the modified arms of the male is supported by the report ofR. Young (1978) of an adult female Bathothauma sp. with two long arms (presumably from a male, which has elongate, modified ends on both arms I) attached in the mantle cavity. During copulation, it would appear that the assumption of a parallel position with the ventral surface of the male against the dorsal surface ofthe female would permit the male, with the long penis extending out of the opening of its mantle, to deposit the sperm reservoirs of the ejaculated spermatophores directly onto the anterodorsal mantle surface (occasionally misplacing them onto the antero- ventral surface or on the head or arms) of the female. The sperm reservoirs attach, become totally embedded in the mantle, then penetrate the inner wall and release the sperm into the mantle cavity where fertilization probably occurs as the eggs leave the oviducal glands. The large, swollen nidamental glands suggest that the eggs are probably deposited in (a) gelatinous egg mass(es). The use ofbiolumines- cence as an attractant and (a) pheromone(s) as close-range stimuli are of particular advantage to an animal that mates in the darkness of the midwaters in the possible absence of a mass breeding congregation. The largest number of mature animals taken by the WALTHER HERWIG in a single haul was 5 over a 3D-minute period. The remaining successful hauls took 1 or 2 individuals over periods of 30-90 minutes. The catches, however, probably give a poor indication of the actual concentration of the breeding population because far more animals are likely to have avoided the trawl than were caught. All of the mature males, but none of the large subadult males, showed recent loss of bodily parts, especially 1 to 3 pairs of arms. Interestingly, arms I, and usually arms II, the two pairs of modified arms, were always intact, suggesting that they may have been inside the mantle of the female during mating when the injuries were sustained. Most of the mature females have only about the basal quarter to mere stubs of all the arms remaining. A few individuals have one or two arms intact; in each case, they are the fourth arms. All of the mature male and female animals, suffered loss of the tentacles. An examination of the stomach contents of the specimens showed the stomachs of2 females distended with chunks of body parts, obviously recently consumed, and pieces of gladii and spermatophores identifiable to pellucida. An additional female and 2 males had chunks of outer body parts and digestive gland tissue tentatively identified to the species. It is, of course, impossible to determine exactly where, or to what extent, the cannabalism took place, whether during mating or in the trawl after capture. Possibly, some of the observed injuries could be attributed to predators, either at the mating scene or in the trawl. The evidence of (1) the marked deterioration of the overall texture of the body in the female; (2) the near-uniform size of the eggs at maturity; and (3) the possible 56 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

•• ••

BO 80 80 80 11. '0

Figure 18. Teuthowenia pellucida. Geographic distribution of specimens reported in this paper.

occurrence of cannibalism during mating suggest that the female probably sheds all of the eggs over a short period of time and does not survive beyond one spawning period. In the male, the evidence of (1) the apparent loss of the suckers on the modified ends of arms I and II during courting and mating; (2) the presence oflarge and small spermatophores in the spermatophoric sac suggesting the making and subsequent storage of spermatophores over a time period; and (3) the indications of possible cannibalism occurring during mating combine to suggest that though the male may mate more than once, mating is done over a short time span. Geographic Distribution. - Teuthowenia pellucida is concentrated circumglobally in a narrow belt of about 6 to 7 degrees width in the area of the Southern Sub- tropical Convergence (Fig. 18). Records of capture indicate that the species extends into adjoining boundary currents and is scattered southward in the fringing waters of the West Wind Drift and northward in the western Indian Ocean and the Pacific into the fringing subtropical waters. Lack of material from the central sector of the southern Indian Ocean precludes knowledge of the distribution in this area. The Subtropical Convergence is the line of convergence formed between the northward movements of subantarctic waters and the southward movements of subtropical waters. This convergence is found circumglobally in the vicinity of 400S, displaced to the south on the western sides of the oceans and to the north on the eastern sides. Across the convergence, the temperature changes from about 10°to 14°C in winter, and 14°to 18°C in summer (Deacon, 1966). The convergence is a relatively shallow feature whose position fluctuates under the influence of surface winds and currents. Along the Mar del Plata-Cape Town transect, the WALTHERHERWIGworking in March 1971 between 34° and 41°S in the area of the convergence captured 36 voss: REVISION OF TEUTHOWENIA 57 specimens of pellucida, ranging from larvae to adults, in 14 (38%) of the 32 hauls at the 16 stations occupied using the Engel MWT of 1,600 meshes circumference. Nine (56%) of the stations were positive for the species. On the northward Cape Town-Madeira transect, on which hauls were made at 3°_6° intervals, the WAL- THERHERWIGcaptured the species at 2 (67%) of the 3 stations occupied between 36° and 34°S, in 2 (33%) of the 6 hauls taken. No captures were made at the stations to the north. The 2 successful stations were made in the productive waters of the Benguela Current, where several specimens from the collections of the WILLIAMSCORESBYand the South African Museum were also taken. Larvae were found in collections of the South African Museum made in surface waters north to 29°S. The type-locality, 33°20'S, 15°58/E, is located in the Benguela Current. Working southward in November along 500W to antarctic waters, the WALTHER HERWIGII during the 1975/1976 cruise encountered pellucida first at 36°S and last at 41°S. Thirty-three specimens, including larvae to adults, were taken at 4 (100%) of the 4 stations occupied, and in 5 (100%) of the 5 hauls taken between these latitudes. On the northward return in January, along 400W, the species was found in the same approximate narrow belt. It was first collected at 42°S and last at 36°S. On the latter leg, 10 specimens, ranging from larvae to adults, were captured at 3 (75%) of the 4 stations occupied, in 4 (67%) of the 6 hauls made within the latitudinal range. Across the area of capture, from south to north, surface temperature and salinity changes of 14° to 20°C, 34.5 to 35.3%0 were recorded. T. pellucida was the most common cranchiid found in the collections from the area of the convergence made by both the WALTHERHERWIGand WAL- THERHERWIGII. Only one of the 53 hauls made to the south of the convergence on the 1975/1976 cruise captured the species. The exceptional haul took 2 specimens, a near-mature female of 179 mm GL and a mature male of 20 I mm GL while fishing at 2,000 m at 48°S, 400W (WH 101-11) below a pocket of warmer water (8°-10°C, 34.6-34.70/00) found in the upper 200 m (Hulley, 1981). The species was not taken in the shallower haul from 580-620 m at the same station or in the 2 hauls made under the pocket at a subsequent northward station. The adults are probably expatriates associated with the warm-water pocket. In the collections from the same stations taken by the WALTHERHERWIGin the area of the warm- water pocket, Hulley (1981) reports finding 7 species ofmyctophid fishes that are normally found to the north in the convergence. In contrast to the case in pellucida, however, he found that the specimens were "generally smaller and sexually immature." The total captures of 4 near-mature and II mature animals from stations in the highly productive waters of the Benguela Current and in the less productive waters of the convergence indicate that pellucida breeds and spawns throughout its range in the Atlantic. Across the convergence the species was found in greatest abundance in the western sector between 36° and 41oS, 48° and 5IoW, where the productivity is higher than in the central sector. Additional captures from the South Atlantic were made by the Russian R1V AKADEMIKKURCHATOVwhich similarly found the species (reported as Verrilli- teuthis mega lops australis) restricted to a narrow belt in the area of the convergence (Nesis, 1974a; 1974b). During two transects of the Subtropical Convergence, made in November-December 1972, pellucida was found between 44° and 45°S, 55° and 56°W and between 41° and 44°S, 26° and 27°W. The few available records of pellucida from the Indian Ocean show an extension ofthe geographic range into fringing southern subtropical waters, a condition also found in the Pacific but not in the Atlantic. This may be explained, in part, by the fact that the convergence, the mixed waters of which are preferred by the S8 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.1, 1985 species, is less defined and its position more variable in the Indian and Pacific Oceans (Knox, 1970; Krey, 1973) largely reflecting a stronger influence of seasonal winds. Adaption to the more fluctuating conditions in these regions may permit the species to better tolerate and colonize the waters adjoining the convergence. Eleven larvae and juveniles from the collections of the South African Museum were taken between August and November in an area between 31° and 39°S, 27° and 45°E, in the region of the convergence and north into the Agulhus Current. Additionally, the DISCOVERYtook a mature female in May at 32°S, 4loE. To the east, single captures of larvae were made in September by the ANToN BRUUNat 32°S, 600E and in December by the Monsoon Expedition at 34OS,73°E. The collections made by the ELTANINlargely using the 3 m IKMT, during Cruises 4, 5, 6, 16, 21, 23, 24, 25, 26, 35 and 46 fairly well define the limits of distribution of pellucida in the southeastern sector of the Indian Ocean and in the Pacific. On Cruise 35 made between August and October in the southeastern Indian Ocean and south of Australia, 36 hauls taken south of the convergence were negative while 3 (33%) of 9 hauls taken between 38° and 46°S, 117° and l300E in the region of the convergence were positive, capturing 6 larvae of the species. On Cruise 46, working in November mostly southwest of Australia, the ELTANINcaptured the species (3 larvae in separate hauls) only between 35° and 39°S, 114° and l15°E. In the collections of the Australian Museum taken in a series of hauls using the 20 x 15-m Engel MWT in a small area between 33° and 36°S, 150° and l51°E in the subtropical waters of the East Australian Current between October and De- cember, pellucida was the most abundant of the 8 species of cranchiids found. Five (33%) of the 15 hauls examined made over bottom depths adequate for the vertical range of the species were successful, catching 14 specimens induding larvae to subadults. None of the 4 hauls taken over shallower depths were successful. Warm-core eddies of the East Australian Current (Brandt, 1983) may affect the distribution of pellucida in the western Tasman Sea. The ELTANINfound pellucida in greater abundance on Cruise 26 while working in December in the south Tasman Sea where the convergence is a well defined feature which seasonally fluctuates in position and degree of development (Knox, 1970). In the area between 39° and 48°S, and about 160° and 162°E the species was concentrated between 42° and 46°S in the area of the convergence, where 7 (88%) of8 hauls were successful, capturing 32 specimens. Only 1 (14%) of7 hauls made to the north was successful, catching 2 specimens. To the south, 2 (50%) of 4 hauls were successful, capturing 10 specimens. Both larvae and juveniles were caught in all areas. The species appears to be scarce in the area to the south of New Zealand where the water depths over the broad expanse of the Campbell Plateau are too shallow for its vertical range. The southernmost record from this general region is a probable expatriate, an early juvenile, taken at about 53°S, l69°E and found in the collections of the National Museum, Wellington. The area off the east coast of New Zealand is one of fluctuating currents, upwellings and convergence where the water depths in the area of the convergence off South Island are largely less than the vertical range of pellucida. Here the convergence is well defined (Knox, 1970) and seasonally fluctuates between about 42° and 46°S. During the year, the convergence is located partly over the east- west oriented Chatham Rise where depths are approximately 400 to 500 m. In a study ofthe mesopelagic fauna in association with the convergence lying over the rise, Robertson and Roberts (1978) report (under the name Taonius richardsom) the capture of 10 young specimens of pellucida in hauls taken in the Hikurangi Trench to the north ofthe convergence and the lack of captures in the convergence voss: REVISION OF TEUTHOWENIA 59 over the Chatham Rise or in the Bounty Trough to the south. The authors conclude that the findings might suggest an exclusive affinity of the species with subtropical waters. I believe that the findings rather point out the limited nature of the study and the atypicalness of this portion of the convergence. The ELTANINmade hauls in this same area on Cruise 23 in April/May working along a diagonal transect from the southeast. The species was not commonly encountered. Out of the 3 (25%) of 12 hauls that made captures, however, 2 were in the area of the convergence-I haul at 44°S, 174°W caught a mature female of 184 mm GL and the second haul took 2 juveniles from off the eastern slope of the Chatham Rise. The third successful haul captured a juvenile in the Hikurangi Trench to the north. Nearby waters in the Cook Strait and vicinity are the localities for the 3 juvenile and subadult paratypes of Megalocranchia richardsoni (=T. pellucida) and 3 juveniles taken by the GALATHEA. The species extends into the fringing subtropical waters to the north and north- east of New Zealand, areas of mixed and fluctuating currents. Here are the localities for the juvenile holotype of M. richardsoni and the larval holotype of the synonymous M. pardus. The latter, taken on a beach in the Kermadec Islands at about 29OS, 178°W is the northernmost record of pellucida in the Pacific. There is little known on the position and the degree of development of the convergence in the central and eastern sectors of the South Pacific. Collections of cephalopods are meager from these waters which are considerably less productive than the waters of Australia and New Zealand to about 1700W. The species was scarcely encountered in the central sector in the austral midwinter on the July! August Cruise 24 of the ELTANINduring which 47 hauls were made along a north- south zigzag transect in the general area of the convergence between 36° and 46°S, 150° and 124°W. None ofthe 16 hauls taken west of 1400W between 40° and 46°S were successful in capturing the species. Of the 31 hauls from east of 1400W, the 5 (16%) which captured a total of 7 early larvae were all made between 38° and 42°S, 135° and 125°W. Information on the distribution of pellucida in the southeastern Pacific is fur- nished by the ELTANINand by stomach contents of the lancetfishes Alepisaurus and tunas longlined during 2 cruises of the SHOYOMARU.On SHOYOMARUCruise 13, made in the early austral summer months of November and December, fish were caught at 19 stations located within an area west of the Peru Current between 5° and 41°S, 133° and 86°W. All 8 stations north of 310S were negative for the species, but 8 (73%) of the 11 stations occupied between 32° and 41° were positive. The stomachs contained 21 late larvae and early juveniles, all in remarkedly good condition. Twenty of the above specimens came from 7 (100%) of the 7 stations from the area between 32° and 41°S, 124° and lOloW. The hydrographic and biological data from this cruise (Forsbergh and Broenkow, 1965) show that the successful captures were all confined to the area of transition or convergence between the cool surface Subantarctic Water of low salinity to the south and the warm Eastern South Pacific Central Water of high salinity to the north. Across the region of con vergence between 41°and 32°S, at 90-9 5°W, from south to north, the temperature changed from 11° to 19°C and the salinity from 34.2 to 35.20/00. The productivity and plankton abundance was moderately high east of 124°W in the area of the captures, while west of 124°W where no captures were made, the productivity and plankton abundance were low. In the preceding austral summer, the SHOYOMARU on Cruise 12 working in January between 13° and 37°S, 109° and 900W encountered pellucida in the same latitudes. Of the 14 longline stations occupied, the 10 from north of 320S were negative for the species, while 2 (50%) of the 4 stations made to the south of 32°S were positive. The Peru Current, which originates from the northward deflection of part of 60 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.1, 1985

the subantarctic West Wind Drift as it encounters the land mass of South America, is a wide, slow-moving current characterized by bands of northward and southward flow (Wooster and Reid, 1963; Craddock and Mead, 1970). Craddock and Mead, working with the midwater fishes and hydrographic data collected by the ANTON BRUUN on 2 transects of the current at 34°S, found a distinct faunal boundary at about 800W. The midwater fish fauna east of 800W was dominated by species found associated with, or to the south of, the Subtropical Convergence. Teuthowenia pellucida is only known from this eastern portion of the Peru Current. The ELTANINcaptured the species in 3 (30%) of the 10 hauls made between 35° and 42°S, 72° and 78°W, while in route south from Valparaiso to the Antarctic on Cruises 4, 5, 6, 9 and 15. The western portion ofthe current was only sampled on Cruise 21 during which pellucida was not taken in any of the 6 hauls made in the area sampled between 33° and 35°S, 80° and 86°W. Though it is very doubtful that the distributional range of pellucida is continuous between the Pacific and Atlantic around South America, single captures oflarvae were made by the ELTANIN and the WILLIAMSCORESBYfrom off the southern coast of Chile and off Cape Hom. In summary, the findings show that though Teuthowenia pellucida is distributed circumglobally primarily in the mixed waters of the Southern Subtropical Con- vergence, its occurrence may be affected locally by surface winds and currents, and the depths of the bottom. The species in all of its life-history stages which inhabit the water column from the surface to over 2,000 m appears to remain within and below the convergence as it fluctuates under the influence of surface winds and currents. Over its geographic range, the species varies in abundance from sector to sector depending on productivity. Gibbs (1968) first recognized that the Southern Subtropical Convergence had a peculiar fauna which constituted "a distinct zoogeographic region encircling the Southern Ocean." Subsequently, Krefft (1974; 1976), Hulley (1981), and others have recognized a growing number of midwater fishes as members of this convergence fauna.

Vertical Distribution. - Information concerning the vertical distribution of Teu- thowenia pellucida is summarized in Figures 19 and 20. Unfortunately, no collections made with closing nets were available. The species was taken in the Atlantic by the WALTHERHERWIGover a temperature range in the vertical column of 19°-2.8°C (100-:::::2,000 m; Lenz, 1975). The combined data suggest that larvae under 10 mm ML may be found from the surface to midwater depths of about 600 m. Young to about 70 mm GL have been taken in the upper 200 m during both day and night. The inadequacy of the data to clearly show the time of ontogenetic descent of the young is partly due to the broad size range (about 53-95 mm GL) over which metamorphosis from the stalked- to the sessile-eye stage occurs in pellucida. Juveniles and early subadults to 120 mm GL have been taken in nets fishing in the daytime at 700-800 m. During night hours, juveniles and subadults appear to move upwards in the water column where they were caught in the upper 300 m by the WALTHERHERWIG,ELTANIN,KAPALAand other research vessels. The largest animal captured at night in the shallow waters was a subadult female of 140 mm GL taken by the WALTHERHERWIGwhile fishing at 200-212 m at 400S, 44°W. A part of the subadult population appears to extend to the surface where the squid may be fed upon by certain marine birds (Imber, 1978). Both sexes, which mature over the same approximate size range of 141-201 mm GL, appear to mature and subsequently remain primarily in the midwater depths below about 1,500 m to possibly 2,400 m or greater. Along the latitudinal voss: REVISION OF TEUTHOWENlA 61

200 'O Number of specimens t 0

400 -- I mmature specimens - Near-malure females = Malure females and = males 600 --- 800 -

E 1000 ...., - .c 0. 1200 Q) o

f400

1600

1800 =

2000

2200 -

2400 (J 20 40 60 80 \00 120 140 160 180 200 Gladius length (mm)

Figure 19. Teuthowenia pel/ucida. Vertical distribution with respect to stage of maturity. All specimens of this species were collected with nonclosing nets. See text for explanation of question marks used in this and the subsequent graph.

transect of the Atlantic range of the species in the waters of the Southern Sub- tropical Convergence made by the WALTHER HERWIG in 1971, near-mature and mature animals were taken in 5 out of 8 hauls (63%) that fished below 1,500 m and in none of the 29 hauls that fished between 100 and 1,015 m. All of the hauls were made at night except for 1 deep and 1 shallow haul. Of the 11 hauls taken along the combined 2 longitudinal transects through the Atlantic distributional range made during the 1975/1976 cruise, the WALTHER HERWIG II captured near- mature and mature animals in 3 (100%) out of3 hauls fishing between 1,800 and 2,400 m, and in none of the 8 remaining hauls which fished between about 100 and 800 m. The additional captures of a near-mature female of 179 mm GL and a mature male of201 mm GL to the south of the convergence at 48°S, 400W (WH 62 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

200

==l!!I!II!t

400 Number of [10 specimens [0

600 Night Twd Ighl Oay 800 - •..... 1000 •.....E ..c 1200 -Q. 0) 0 1400 =

1600

1800

? ? 2000

2200 -

2400 o 20 40 60 80 100 120 140 160 180 200 Gladius length (mml Figure 20. Teuthowenia pellucida. Vertical distribution with respect to time of day.

101) were made while fishing at 2,000 m. The only known non-Atlantic captures of mature animals are those of a l83-mm ML female made by the DISCOVERY while fishing during midday at about 2,000 m at 32°S, 4loE in the western Indian Ocean and a l84-mm GL female made by the ELTANIN while fishing at night at 1,813-2,274 m in the western Pacific at 46OS, l700W. All of the above catches of near-mature and mature animals were made over bottom depths in excess of about 3,800 m. The lack of captures of mature females in the shallow depths and the presence of sperm reservoirs in the mantles of several of the mature females suggests that mating and spawning in pellucida occurs in the deep midwaters. The captures shown with a question mark on the graphs are those made by the WALTHER HERWIG on the 1971 cruise during which hauls that fished at about 2,000 m were also fished horizontally at 2 intermediate depths, 1,000 and 500 m, in order to cool the trawl winch during net retrieval. This greatly increased voss: REVISION OF TEUTHOWENIA 63

the likelihood that the immature animals captured in these deep hauls are actually from the shallower depths. After 1973, retrieval of the net was done without interruption (Krefft, 1976). Geographic Variation. -A substantial amount of the morphological variation that occurs in Teuthowenia pellucida is manifested in the early and middevelopmental stages and is largely related to the timing of metamorphosis from the larval to the juvenile stage. The broad distributional range over which the young are found encompasses a variety of environmental conditions: mixed, cold and warm waters and areas of high, moderate and low productivity. There appears to be no cor- relation between size at metamorphosis and any single environmental feature. The larval stage terminates at about 53 to 70 mm GL throughout the range of distribution regardless of temperature and productivity, except in four known areas: (1) the warm subtropical, moderately productive waters in the western South Pacific north of New Zealand, between about 31° and 37°S, 177° and 178°E; (2) the cold, highly productive waters of the Benguela Current between about 29° and 35OS, 13° and 17°E; (3) the mixed, moderately productive waters of the Subtropical Convergence in the eastern South Pacific between about 32° and 38°S, 90° and 124°W; and (4) the warm, highly productive waters in the area of the Agulhus Current between about 35° and 39°S, 28° and 34°E. In these four areas the larval stage terminates between 75 and 95 mm GL. As a result, the proportional size of the eyes and the enlarged suckers on arms III, and the length of the arms of the young are usually less at a given size than are found in the young from the nonexceptional areas. These differences, however, are submerged with growth and are not found in the subadults and the adults, where known, from these localities. Other characters vary differently over the range of the species. For example, the exact pairs of suckers on arms III that show maximum enlargement in juveniles and adults is usually typical of the population from an area and appears to be correlated with temperature. Sucker pairs 9-10 or 10-11 are usually the largest in areas of colder waters while pairs 11-12 or 12-13 are usually the largest in areas of warmer waters. Another character that is often shared by the animals from the same area, but appears to be unrelated to anyone feature of the environment, is the number of teeth found on the sucker rings on the ends of the arms. A count of 12-16, for example, is typical of the specimens from the East Australian Current, while 5-9 is commonly found in specimens from the Peru Current. The number of teeth is most variable in specimens from the convergence in the Atlantic. No examples of clinal variation were found. Though Teuthowenia pellucida has polytypic young, little substantial morphological differentiation is found in the subadults and adults in spite of evidence of localized, breeding populations. Remarks. - The type-specimen of Desmoteuthis pellucida, together with several other of Chun's types, originally deposited in the Leipzig Museum are now re- deposited in the Zoological Museum, Humboldt University, Berlin. My examination of the specimen along with the type-specimens of the junior synonyms Megalocranchia pardus, Anomalocranchia impennis, Megalocranchia richardsoni (paratypes VUZ 57, VUZ 110) and Megalocranchia megalops australis showed them all to be in specifically identifiable condition. Though not included in the original descriptions, the species-specific multipoint tubercle on the funnel-mantle cartilages can be clearly seen in the types of impennis and richardsoni. In the case of pellucida, a low, relatively stout knob remains on the single intact cartilage; while in pardus, only a single-point tubercle is developed on each cartilage. The characteristic four rows of suckers on the tentacular stalks not described by Robson 64 BULLETIN OF MARINE SCIENCE. VOL. 36, NO. 1,1985 for impennis are clearly in evidence in the type-specimen. Dell's original illus- tration of the holotype and my examination oftwo of the paratypes of richardsoni show that the specimens are in a contracted condition. Measurements of the broken gladii of the paratypes indicate the specimens are at least 12-15% larger than the original measurements. This discrepancy partly explains the differences in relative size of the eyes and shape of the mantle at a given mantle length that Dell found in his material. The remaining differences and those found among the other synonymous species can be attributed to the incompleteness of the original descriptions and the above described environmental effects on developmental morphology found to occur in the diverse type-localities. Chun (1910) observed the type-specimen of pellucida alive and remarked on the transparency of the animal and the metallic sheen of the envelope of the digestive gland. Additionally, he noted a transparent slime layer covering the mantle and concluded from histological sections that the entire ectoderm was involved in the production of the slime rather than special gland cells. The "slimy" condition may be similar to that found in Megalocranchia fisheri in which the very thick "dermis" makes the animal very slippery (R. E. Young, pers. comm.). Chun also described and illustrated the histology of the ocular photophores. On preservation, he recorded nearly a 40% contraction in the length of the tentacles. Roper (unpubl. pers. journaF) also recorded observations on a living animal, an 81-mm GLjuvenile collected on the March/April 1971 cruise ofthe WALTHER HERWIG,which he observed to hold its tentacles downward with the clubs curved upward. The animal ejected violently when disturbed, and spun away making 3- 4 complete revolutions, spinning particularly from the tail region. The juvenile was also seen to vigorously hold and bite the intruding forceps.

GENERALDISCUSSION Comparison of the Species The following comparison is of adults and subadults; larvae and juveniles are discussed in the subsequent section. Though the three species of Teuthowenia are similar in general appearance, a close study reveals a variety of species-specific features, the majority of which are related to maturity. Selected characters for the three species are compared in Table 14. Within the genus, the maximum size attained varies among the species, and, at least in one case, varies markedly between the sexes. In pellucida, the smallest species, both females and males mature over the same approximate size range of about 141-201 mm GL, while in megalops maximum size is sexually dimorphic; the males mature at about 182-244 mm GL, about 30 to possibly as much as 50% shorter than the females. The markedly larger size at maturity of the female of megalops may be correlated with the greater productivity of the northern waters inhabited by the species and may permit an increase in egg production per individual. Thus I would expect that the female of mega lops would produce sig- nificantly more eggs than the smaller maturing female of pellucida. No mature specimens are known for maculata, but if such developmental features as the late appearance of the modifications on the ends of arms I and II in the male and the overall robust growth displayed by the species are taken as indications, and the high productivity of the waters inhabited in the eastern tropical Atlantic is considered, then maculata may be expected to exceed the size of mega lops at maturity. There are no indications in the species, however, as to the possible presence or absence of maximum size differences between the sexes. The multiple tubercles, which usually fuse basally to form a multipoint tubercle voss: REVISION OF TEUTHOWEN/A 65

Table 14. Comparison of subadults and adults of the three species of Teuthowenia

megalops* maculalat pellucida Size at maturity 9 >350 mmGL ? 141-201 mm GL & 182-244 mm GL ? 141-201 mm GL Number of tubercles, I (rarely 0) 2-4 (occasionally I or funnel-mantle 5) cartilage ALF 9 and &, subadult III> II> I> IV III> II> IV= I III>II>IVE!!I 5, adult I>III>II>IVor ? III>II>I>IV III> 1=11> IV ASI, eIII 2.0-2.8 (2.4) 3.2 2.0-2.3 (2.2) 9 Modifications at maturity Brachial end-organs 6-9% AL (maturing) ? 13-19% AL Glandular processes present ? absent on arms Glandlike organs on ? ? present dorsum of mantle & Arm modifications at maturity Suckers, basal and present ? present medial, modified Arms I elongate; greatly in- 25-27 pairs of suckers increased robustness; creased robustness; prior to elongate 11-18 pairs of suck- midportion modi- modified end; bare ers prior to elongate fied; end not modi- tip modified end; bare, fied; increased suck- whiplike tip er dentition Arms II 15-19 pairs of suckers 25-27 pairs of suckers increased robustness; prior to elongate, prior to elongate, 11-20 pairs of suck- modified end; end modified end; end ers prior to elongate, with 2 rows special- with 2-4 rows of modified end; end ized suckers; in- specialized suckers; with 2-4 rows spe- creased sucker den- bare tip cialized suckers; bare tition whiplike tip Arms III ends whiplike, in- ? increased robustness creased sucker dentition Arms IV ends whiplike; in- ? increased robustness creased sucker dentition MaSTC 19-24 22-26 26-32 SpLI 17-26 (20) ? 8-9 (9) LRL Q 5.4-7.0 (maturing) ? 4.4-5.0 (near mature, mature) & 4.0, 4.1 (mature) ? 4.0, 4.5 (mature) GTILI 24-29 (26) 23 20-22 (21) • No mature or ncar~mature females known. t "l~ macu/ata represented only by one early subadult male. in the large animal, found on the funnel-mantle cartilage distinguishes pellucida from the other two species which characteristically have but a single-point tubercle. Because the fins extend posteriorly to accompany the elongation of the posterior end of the gladius, the fins are longer in megalops, which has a proportionally 66 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.1, 1985

longer gladius tail, than in pellucida, which has an approximately 20% shorter gladius tail. The accelerated growth of the posterior end of the gladius observed in the early and middevelopmental stages of maculata suggests that fin length in the genus may reach its maximum in this species. Such characters as funnel organ, eye size, ocular photophores and the pattern of buccal membrane attachments display little variation among the species, except during growth. The degree of enlargement of the suckers on the approximate third quarters of arms III, and to a lesser extent arms II, varies among the species. The enlargement is greatest in macula/a, followed by megalops, and is least in pellucida. As in all cranchiids (N. Voss, 1980), the arms in Teuthowenia exhibit a variety of dimorphic sexual features, the majority of which are male related. Most of the arm modifications found in the male are similar in general form but may vary in details in the different species. For example, the suckers on the basal and midportions of all of the arms have similar swollen, fleshy collars and rings with reduced apertures in both species where adults are known. The elongate, modified ends of arms I and II in macu/ata and pellucida, and arms II in mega lops have the same type of specialized suckers and show a similar reduction ofthe protective membranes. Variations occur, however, in the number of rows of suckers on the ends, 2 in mega lops, 2-4 in maculata and pellucida; the development of distinctive bare, whiplike tips in pellucida; and the number of pairs of normal suckers found on the arms prior to the modifications, 11-20 in pellucida, 15-19 in megalops and 25-27 in maculata. The overall similarity of form suggests that the use of the modified ends in megalops and maculata may be similar to the presumed use in pellucida, i.e., stroking the inner mantle walls and organs of the female during courtship and copulation. The occurrence of increased robustness of the arms at maturity varies between pellucida and mega lops and suggests differences in the functions ofthe particular arms in mating. The increased robustness of all of the arms, especially arms IV, in pellucida suggests that arms IV and the other arms, to a lesser extent, are used to hold the female. Arms I and II appear to perform a dual function of holding and caressing. The strikingly different modifications of arm I and the increased dentition of the normal suckers on all of the arms in mature males of mega lops correlates with the increased size of the female over the male in that species. The greatly enlarged arms I and the increased sucker dentition are of advantage to the male in copulation with the larger probably less acquiescent female. The absence of the end modification on arms I suggests that the arms are probably primarily used for holding rather than the dual function of holding and internal caressing that is presumed for pellucida, and may also be for maculata. Two of the maturity-related features displayed by the female occur on the arms. One feature, the brachial end-organ, occurs on all of the arms in megalops. In pellucid a, the only other species in which adult females are available, the end- organ occurs on arms IV, the only arms intact in available specimens. Because the occurrence and position of the end-organ are usually constant within genera of cranchiids, and the fact that the end-organ is found on arms IV only in those species that have the organ on all of the arms (N. Voss, 1980), it is presumed that the brachial end-organ occurs on the ends of all of the arms in all species of Teuthowenia. The end-organ varies in proportional size between megalops and pellucida. It is small in the former species, measuring 6-9% of the arm length, and large in the latter species, measuring 13-19% of the arm length. Since the number of normal suckers on the arms prior to the end-organ is affected by the size at which maturity occurs, the number of sucker pairs is larger, 28-43, in the voss: REVISION OF TEUTHOWENIA 67 larger maturing mega lops, than in the smaller species pellucida, in which the sucker pairs number 17-21. The second female modification affecting the arms, the glandular processes found scattered between the largest suckers on all of the arms, may be specific for megalops. The processes are not found in pellucida. Other female changes that occur in the genus, take place so close to maturity that they are known only for pellucida, the one species where mature females are known. The overall deterioration of the body observed in pellucida probably occurs in all of the species as it is found to occur in all cranchiids in which mature females are known and to be common in non-cranchiid teuthoids (R. Young, 1973; Arnold and Williams-Arnold, 1977; Kristensen, 1981b; and others). There is variation, however, as to the extent and timing of the deterioration. The resulting reduction in the density of the tissue may be related to the increased metabolic demands of the maturing female, and may also serve as a buoyancy mechanism to counter the markedly increased weight of the eggs and genital organs. The glandlike organs, for which a pheromone secretory function is speculated, that develop in pellucida along the anterior, middorsal line of the mantle, may either be specific or be shared with one or both of the other species. Their occurrence has not been reported in any other cephalopod. The mature female, however, is not known for many oceanic species, particularly from the midwaters, and it is possible that some will be found to have these organs. The use of pheromones to attract and stimulate the male during mating is not known for cephalopods but was suggested by Fields (1965) based on his observations of mating in Loligo opalescens and the observations of Tompsett (1939) on Sepia. The tentacles and clubs of the three species are similar with the exception of the dentition of the club suckers. The rings of the large suckers bear the largest number of teeth in pellucida, 26-32, and progressively fewer in maculata, 22- 26, and megalops, 19-24. A detailed study was not done on many of the internal structures and organ systems, and probably intrageneric morphological differences do occur. No notable differences, however, are evident in the gross morphology of the digestive and reproductive systems. A few internal features were selected for comparative studies. The spermatophores, while similar in internal structure, exhibit specific differences in proportional length and size of the internal parts. The length differences are substantial. The spermatophore of megalops is moderately large, about 20% of the gladius length, in contrast to the small spermatophore of pellucida which measures about 9% of the gladius length. Less conspicuous variations occur in the proportional size of the internal parts. While the sperm mass is similarly long in both species, occupying about 50% of the spermatophore length, the spermatophore of megalops has a shorter cement body, about 24% of the spermatophore length compared with 37% for pellucida, and a longer ejaculatory apparatus, about 19% of the spermatophore length compared with 13% for pellucida. The gills are similar in length, but the number of lamellae in the outer demibranch varies somewhat among the species, tending to be higher in mega lops (23- 26) and maculata (23-24) than in pellucida (18-22). Though the mandibles are similar in major features, subtle specific differences do occur. Because changes accompany growth, only the mandibles of megalops and pellucida. the species in which adults are available, are compared. Comparing the lower mandibles, the wing fold is moderately low and only slightly obscures the jaw angle in megalops, while in pellucida it is more developed and clearly obscures the jaw angle. Additionally, the hood tends to sit higher above the crest in mega lops than in pellucida. The degree of development of the broad fold on the lateral wall tends to vary in both species. The rostral lengths at maturity are 68 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985 similar for males of megalops and both sexes of pellucida but are greater for the markedly larger maturing females of megalops. Rostral lengths in pellucida tend to be proportionally greater, however, at a given size than in mega lops. Addi- tionally, females of pellucida tend to have greater rostral lengths at a given size than males of the species. No distinctive differences are found in the radulas, with the exception of the noticeably stouter bases of the rachidian and first lateral teeth in pellucida. Variations of the typical form of the "Teuthowenia" gladius are found among the species. The greater elongation ofthe posterior end of the gladius in mega lops and maculata results in distinctly longer vanes, 50% or greater of the gladius length, than are found in pellucida in which the vanes measure about 36-40% of the gladius length. The delicate, thin lanceola of mega lops is distinctive, con- trasting with the sturdier form displayed by the other two species. Though the free rachius of pellucida tends to be the stoutest among the species, the posterior end of the gladius is markedly stoutest in maculata. The latter species, additionally displays the strongest develoment of the dorsal keel.

Growth and Development of Young A comparison of selected features of the larvae and of certain changes that occur with growth in the three species of Teuthowenia are shown in Table 15 and Figures 21-24. A comparison of the young reveals obvious differences in chromatophore patterns and patterns of development. In fresh or well-preserved specimens of the different species, larvae as small as 5-8 mm ML (and possibly smaller) and juveniles can be distinguished on the basis of species-specific variations of the basic "Teuthowenia" chromatophore pattern. Although the patterns on the head, funnel, arms and tentacles are similar, distinct differences occur among the species in the size, number and spacing of the chromatophores on the mantle. The chromatophores are largest, fewest in number and most widely spaced in megalops; followed by pellucida in which they are of medium size and number, and set in a moderately dense pattern; and, finally, by maculata in which the chromatophores are very small, numerous and densely arranged. The different species display additional, less obvious morphological differences, some of which result from differences of size at metamorphosis. Variations in the size at metamorphosis are environmentally induced and may occur both inter- and intraspecifically. The size is smallest, between about 55 and 60 mm GL, in maculata which inhabits warm tropical waters; it is most variable, between about 53 and 95 mm GL, in pellucida over its wide circumglobal range in subtropical convergent and fringing waters; and it is generally largest, between about 75 and 95 mm GL, in megalops which inhabits subpolar and temperate waters. A comparison of the intraspecific variations in the size at metamorphosis as its occurs in megalops and pellucida reveals physiological differences between the species. In megalops, the larval stage terminates at a larger size in the cold waters of the subarctic region in the western Atlantic than in the region of relatively warm temperate waters in the eastern Atlantic. The variations appear to be temperature related. On the other hand, the size at metamorphosis in pellucida is similar over most of its distributional range regardless of temperature and productivity, except in four known, largely border areas where metamorphosis occurs at a larger size. These latter areas encompass a variety of environmental conditions. The evidence indicates that pellucida has a wider physiological tolerance than mega lops that enables it to adapt to a variety oflocal environments. In the case of both species, the areas where the larval stage terminates at the larger size may have marginal voss: REVISION OF TEUTflOWENIA 69

Table IS. Comparison of the larvae of the three species of Teuthowenia

megalops maculata pellucida Chromatophore pattern widely spaced; dense; approx. moderately spaced; on mantle (55-60 mm GL) approx. 16/5 mm2 approx. 8/5 mm2 4/5 mm2 Size at which eyes become sessile 75-95 mm GL 55-60 mm GL 53-95 mm GL 5-24-mm ML size group Number of tubercles, funnel-mantle cartilage o o 0-1 ALl,I1I 4-10 (6) 12-18 (IS) 3-14 (10) ASI, eIlI 0.9-1.3 (1.1) 1.2-1.6 (1.4) 0.9-1.1 (1.0) GiLl 13-16 (IS) 22-29 (26) 16-18 (17) GTILl 3-7 (5) 6-8 (7) 3-9 (7) 25-44-mm ML size group Number of tubercles, funnel-mantle cartilage 0-1 o 1-3 (occasionally 0) ALl,I1I 9-14 (12) 9-22 (16) 8-12 (10) ASI, eIlI 0.7-0.9 (0.8) 1.0-2.0 (1.5) 0.6-0.9 (0.8) GiLl 10-18 (13) 14-21 (18) 11-14 (13) GTiLl 8-12 (10) 11-17 (14) 5-9 (8) 45-64-mm GL size group Number of tubercles, 2-4 (occasionally funnel-mantle cartilage I (rarely 0) o (occasionally I) I or 5) ALl,I1I 11-16 (14) 16-23 (19) 11-17 (14) ASI, eIlI 0.8-1.2 (l.0) 1.3-2.1 (1.7) 0.8-1.3 (1.1) GiLl 10-11 (II) 13-18 (IS) 10-12 (11) GTILI 10-12 (II) 13-15 (14) 10-12 (II)

conditions for early development to which the species is less adapted. The eye stalks may function as a buoyancy mechanism (J. Young, 1970; Clarke et at., 1979); their retention for a longer period would assist the animal to remain longer, and grow larger, in the more productive shallow waters prior to ontogenetic descent. The time at which the tubercles on the funnel-mantle fusion cartilages first appear varies among the species. A single tubercle appears early in pellucida at about 10-20 mm ML followed by additional tubercles at about 25-30 mm ML. The appearance of the single tubercle typical for megalops and maculata is delayed to the midlarval stage, between about 30-60 mm ML, in mega lops, and to the late larval or early juvenile stage in maculata. The eyes show a similar pattern of growth. They are proportionately larger in the early larva to about 18-24 mm ML of all species than in the larva in the immediately subsequent midlarval stage of development. Eye growth in the late larva displays constraint. A maximum size of 11% of the mantle length is found in the final, stalked-eye stage of all three species. The growth of the arms in the early larva is strongly affected by the environment. Arm lengths at a given mantle length are smallest in megalops, vary widely in pellucida and are greatest in maculata. During later larval development, arm lengths tend to become similar in mega lops and pellucida, but in maculata the disproportionate growth, particularly of arms III, continues throughout larval development. Regardless of the variations in arm growth, however, the arm length 70 BULLETlN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

16 •

15 o ()

14 o 00 o 00 o o

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7 0 100 200 G L (mm) Figure 21. Changes in diameter of eye with growth in the three species of Teuthowenia. Circles denote females; squares, males; triangles, individuals of undetermined sex. Open symbols denote immature; half-open symbols, near-mature (in mega[ops, individuals in early maturing stage; see text); solid symbols, mature individuals. voss: REVISION OF TEUTHOWENIA 71

50 •

40 • •• WI • • WI WlO () () 30 () () ~ 0 0 () WI ()

0 () 00 20 0 O~ 0 0 0 0 0 EO~O o 00 10 00 1:;1:; Cb r:&,1:;1:;1:; mega lops

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I:; 10 I:; {J.M{J. l>~ t:L:, pellucida {J. r' 00 100 200 300 400 G L (mm) Figure 22. Changes in length of arms I with growth in the three species of Teuthowenia. For explanation of symbols, see legend to Figure 21. formula, III> II> IV> I, remains the same in the three species. Additional species differences are found in the morphology and growth pattern of the suckers. The arm suckers in larvae of maculata greater than about 10 mm ML are usually noticeably larger and have wider apertures than in the other species. The disproportionate enlargement of the suckers on arms III, and to a lesser extent on arms II, is greatest in maculata. Larvae of maculata larger than about 40 mm ML may have enlarged suckers as much as twice the size in mega lops and pellucida. The 72 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985 pattern of sucker enlargement in megalops is not as markedly disproportionate as it is in maculata and, to a lesser degree, in pellucida. The size of the gills varies among the species. A conflict between the greater proportional lengths of the gills reported for larvae of mega lops by Dilly and Nixon (1976), and the lower values found in the present study and shown in Table 15 requires additional study. Despite the differences, however, the gills in maculata clearly tend to be larger than in the other two species. The larger gill size correlates with the lower oxygen content of the tropical eastern Atlantic where maculata lives. Similar relationships between gill size and oxygen concentration have been reported for some midwater fishes (Ebeling and Weed, 1963; and others) and the bathypelagic squid Bathyteuthis (Roper, 1969). The proportional length of the gladius tail is approximately the same in larvae of all three species up to the mantle length of about 20 mm. Subsequently, in maculata, there is an acceleration in the growth ofthe posterior end of the gladius so that by about 30-40 mm ML, the gladius tail is usually noticeably longer than in its congeners. Additionally, the vanes of the gladius are broader and thicker in cross section in maculata. In general, the larva of maculata is usually more advanced and robust at a given mantle length (with the possible exception of larva smaller than about 7 mm ML) than the larvae of its congeners. The metamorphosis oflarva to juvenile involves in all of the species a similar pattern of accelerated growth, and ecological and physiological changes. Some of the intraspecific differences that are the results of differences in developmental rates that occurred in the larval stage, such as variations in arm lengths and some of the variations in arm sucker enlargements, are gradually submerged over the period of juvenile growth. Subsequent to metamorphosis, the eye abruptly enlarges in all the species to attain similar, near-maximum proportional size in the early juvenile. The enlargement of the near-hemispherical eye is accompanied by a similar pattern of increased fragmentation of the ocular photophores which serves to increase the diffusion of bioluminescence, thereby presumably affording better camouflage for the animal in its newly acquired habitat in the greater depths (R. Young, 1975b). The greater proportional lengths of arms III and size of the enlarged arm suckers that are exhibited by maculata in the late larval stage are maintained during juvenile development. The timing of the initial appearance of the male modifications on the ends of arms I and II in maculata and pellucida, and arms II in megalops varies among the species. It closely follows metamorphosis in megalops and pellucida, but is delayed to the early, midjuvenile stage in maculata. Con- siderable intraspecific as well as interspecific variation occurs. In juveniles of 125- 130 mm GL, 8-16% of arms II are modified in megalops, and 6-22% in pellucida in contrast to 3 and 5% modified in the two available specimens of maculata (123 and 143 mm GL) from the same approximate size range. Early elongation of the posterior end of the gladius subsequent to metamorphosis submerges the interspecific larval differences and results in the proportional length ofthe gladius tail in the early juvenile being similar and greater in mega lops and maculata than in pellucida. In the two species in which adults are known, megalops and pellucida, the gladius tail attains the adult (male adult in the case of megalops) proportional length in the early juvenile stage. The posterior end of the gladius in the juvenile of maculata, as in the late larval stage, is distinguished by its overall robustness and greater development of the dorsal keel. In summary, the species of Teuthowenia have differentiated physiologically, voss: REVISION OF TEUTHOWENIA 73

() ~. () 40 • () ~ () ~ [];j .. ~ 0 () 0 30 •• 8 000 CD 0 o~ 0 00 20 COO0 CO 0 6 0 10 6 66 6 6 6 & megalops

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Figure 23. Changes in length of arms III with growth in the three species of Teuthowenia. For explanation of symbols, see legend to Figure 21. 74 BULLETIN OF MARINE SCIENCE, VOL. 36, NO. I, 1985

() 2.5 ~ () # o ° 0 •

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Figure 24. Changes in diameter oflargest enlarged sucker on arms III with growth in the three species of Teuthowenia. For explanation of symbols, see legend to Figure 21.

developmentally and behaviorally as well as morphologically. They display a high rate of evolution in male and female secondary sexual characters.

Biogeography The discrete, ecologically distinct, distributional patterns of the three species of Teuthowenia reflect the influence of a number of biological and physical factors. One of the most important biological factors, apparently, is primary productivity, which seems to playa markedly restrictive role in mega lops and maculata. The former species is confined to the highly productive Atlantic subarctic and to the highly productive areas of the North Atlantic temperate region. The latter species is restricted to the areas of year-round, high productivity in the eastern tropical Atlantic. T. maculata is observed to be scarce in the Gulf of Guinea where high productivity is seasonal, due to the seasonal occurrence of the equatorial front. The absence may be explained by the fact that maculata, typical of the genus, does not migrate horizontally and its life span is longer than the period over which the effect of the seasonal phenomenon persists. Productivity plays a less restrictive voss: REVISION OF TEUTHOWENIA 75

role in the distribution of the third species pellucid a which inhabits the transitional waters of the Southern Subtropical Convergence. Over the circumglobal range in the convergence, where the species is found in areas of varying environmental conditions, productivity has a strong localizing effect. As a result, pellucida displays a higher degree of provincialism, at least in the young, than the other species. Patchy occurrence in the range of all of the species relates to productivity, with greatest abundance always associated with areas of greatest productivity. While productivity correlates with the limits of distribution of a species in some sectors of its range, major and lesser-defined hydrographic fronts mark the boundaries of distribution for megalops and maculata in other sectors. In the case of pellucida, a convergence serves as a habitat, and its boundaries essentially describe the range of the species. All of the species undergo ontogenetic descent from the epipelagic waters where they develop to mesopelagic and upper bathypelagic depths of about 900 m to in excess of 2,500 m where the near-mature animals remain to mate and spawn. There is evidence, especially in the case of pellucida, that even after descent to the greater depths and the animal has ceased to move upward at night in order to feed, the near-mature and mature animals somehow retain their association with the same shallow waters in which they developed. This phenomenon may have a localizing effect within the range of a species. Additionally, occurrence within range is affected by the animal's requirement for bottom depths sufficient to accommodate its life history. Though the geographic ranges of the species cannot be satisfactorily delineated by particular isotherms or isohalines, temperature, and probably to a lesser extent, salinity, nevertheless correlate with the distributions of the different species. Each species occupies different ranges of temperature and salinity. The controlling effect is probably primarily exerted on the larval stage which is spent in the near-surface waters. Juveniles and subadults become increasingly tolerant to greater differences of conditions as they ontogenetically enlarge their vertical ranges in the water column. Presumably expatriated adults can mature and spawn in the deep midwaters where conditions are more uniform from area to area but subsequent development of the larvae in the foreign, near-surface waters is unlikely. There is evidence that the thermocline may affect vertical distribution patterns in the genus. Data, though meager, suggest that in maculata early development to about 20 mm ML is confined to the surface 100 m, while in megalops and pellucida it occurs in the upper 200 m. The shallow depths at which early development occurs in maculata may be related to the shallow depth of the thermocline in the tropical waters inhabited by the species. Oxygen concentration does not critically limit the distribution of Teuthowenia; one species, maculata, even inhabits the areas oflowest concentrations, 1.5-<0.5 mVliter, found in the Atlantic (Bubnov, 1966). Warm- and cold-water pockets and rings may have an effect, albeit temporary, on the occurrence ofa species within and outside of its normal range. Warm-core intrusions from the Gulf Stream may have an excluding effect on the occurrence and abundance of mega lops within its range in the region of the Slope Water (Lu and Roper, 1979), while to the south in the area of Bermuda, cold-water rings that have separated off from the Gulf Stream may serve to expatriate the species (Lea, 1978). In the subantarctic region, expatriated adults of pellucida have been captured below a warm-water pocket found to the south of the range of the species in the subtropical convergence. Geographic isolation of the three species appears to be complete. However, scattered, expatriated larvae of mega lops found in near-surface waters to the south 76 BULLETIN OF MARINE SCIENCE, VOL. 36, NO.1, 1985

ofthe southern limit of the species in the eastern Atlantic, and of pellucida taken in the Benguela Current north of its range in the subtropical convergence suggest that there may be overlaps of expatriated larvae at the northern and southern ends of the distribution of maculata in the eastern tropical Atlantic. Distributions of the species of Teuthowenia are not closely related to particular water masses but rather are defined by a combination of biological and physical environmental factors primarily associated with the surface waters. The distributions largely conform to and are well described in terms of the zoogeographic scheme of faunal provinces, regions, and distributional patterns proposed by Backus and Craddock (1977) and Backus et al. (1977) for the mesopelagic myctophid fishes in the Atlantic, and predicted by them to be "generally applicable to pelagic plants and animals." The provinces and regions were defined using major and lesser-defined hydrographic features and they serve as the basis for describing the distributional patterns.

Relationships with Other Taoniins

N. VOSS (1980) grouped Teuthowenia together with Egea and Megalocranchia, and this collection of three taoniin genera was called the Megalocranchia group by N. Voss and R. Voss (1983). Shared, derived attributes that support an hy- pothesis of monophyly for the Megalocranchia group include the possession of elongate, terminal-lateral fins and, in mature females, of brachial end-organs. Within the group, Egea and Megalocranchia are proposed to have shared a common ancestor more recently than either genus did with Teuthowenia-character states that distinguish Egea and Megalocranchia from Teuthowenia include possession of a funnel valve, a funnel organ with two lateral flaps and no median element and subtriangular funnel-mantle fusion cartilages (N. Voss and R. Voss, 1983). Teuthowenia is further distinguished from the pair as well as from all other cranchiids by the unique possession offour rows of carpal suckers on the tentacular stalks in contrast to the typical two rows. Since mature females are not known for two of the species of Teuthowenia, the glandlike organs that occur on the dorsum of the mantle in mature females of T. pellucida cannot be regarded as a distinctive generic character. The organs have not been found in the eight other cranchiid genera (Megalocranchia, Taonius, Galiteuthis, Sandalops, Helicocran- chia, Bathothauma, Leachia, Liocranchia) in which mature females are known. The Megalocranchia group is proposed to have shared a common ancestor more recently with the Taonius group (comprised of Taonius, Galiteuthis and Mesonychoteuthis) than either did with members of a third, less cohesive assem- blage oftaoniin genera, the Sandalops group (consisting of Sandalops, Liguriella, Helicocranchia and Bathothauma). The Megalocranchia and Taonius groups are united by the shared, derived absence of a conus and the presence of a pseudo conus on the gladius of the larva and adult; elongate, terminal or terminal-lateral fins; and one large plus one or two small, non-contiguous, complex ocular photophores. The reconstruction oftaoniin relationships advanced above appears better supported than alternative arrangements, but new characters to corroborate or refute these hypotheses would be of considerable interest. Unfortunately, the information that would be necessary to apply novel characters as critical tests is not yet available. For example, additional aspects of gladius morphology not used in previous phylogenetic analyses, the shapes of the free rachis and the vanes, show close similarities between Teuthowenia, Egea and Taonius. The gladii of these three genera have a long, slender free rachis and vanes that form an expanded, spindle-shaped lanceola anterior to the pseudoconus. Similar conditions are also voss: REVISION OF TEUTHOWENIA 77 found in the juvenile but not the adult of Mesonychoteuthis. Megalocranchia and Galiteuthis, on the other hand, have a moderately short, broader free rachis and long, narrow vanes. Dissimilar details of structure and development suggest that the conditions in the latter two genera are independently derived. Megalocranchia exhibits the most specialized gladius in which the lateral margins of the rachis, for most of its length, thicken to form a hollow, rodlike ventral keel, and nearly the entire length of the vanes infold to form the pseudoconus. These patterns of gladial resemblance mayor may not be conformable with existing phylogenetic hypotheses, however, the outcome of the test depending on which of the several states described above is primitive and which derived. A broad survey of this character in outgroup taxa (N. Voss and R. Voss, 1983) is required. Because of the relatively high degree of inter- and intrageneric variation in morphological features of the cranchiid beak (Clarke, in prep. I), that structure may be of little value in determining relationships in the family. Preliminary results of research on other organ systems, however, are more promising. The morphology of spermatophores, statocysts and photosensitive vesicles vary among cranchiids (S. Hess, MS; J. Young, 1984; R. Young, 1978); these investigations together with continuing monographic studies of additional cranchiid genera may confidently be expected to further clarify remaining problems and uncertainties in cranchiid phylogenetics.

ACKNOWLEDGMENTS

I thank the curators and staffs of the various institutions for the loans of types and other specimens. G. L. Voss and R. E. Young critically read the draft manuscript, and R. S. Voss, the section on generic relationships; their comments and suggestions are sincerely appreciated. The illustrations were done by C. S. McSweeny, the histological sections by F. Mucha, and the microphotographs by S. C. Hess; I acknowledge their contributions. This study was supported by National Science Foundation grants DEB-7713945 and DEB-8105193. It is a scientific contribution from the Rosenstiel School of Marine and Atmospheric Science, University of Miami.

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DATEACCEPTED: May 30, 1984.

ADDRESS:Rosenstiel School of Marine and Atmospheric Science, University of Miami, 4600 Rick- enbacker Causeway, Florida 33149-1098.

ApPENDIX Other Material Examined

Teuthowenia megalops.-ALBATROSS: I ?s, 60 mm ML, Sta. 2034, 39°27'N, 69°56'W, 2,460 m, 17 VII 1883, YPM 12468; I 't, 153 mm GL, Sta. 2235, 39°12'N, 72°04'W, 0-1,294 m, J3 IX 1884, USNM 39969.-ANTON DOHRN: I d, 71 mm ML, Sta. 364, 42°05'N, 23°30'W, 155 m, 2 V 1979, ZMH; I 't, 54 mm ML, 2 dd, 75, 58 mm ML, Sta. 382, 44°56'N, 17°18'W, 0:::350m, 4 V 1979, ZMH; 2 't't, 296, 75 mm GL, 2 dd, 61, 59 mm ML, Sta. 389, 45°51'N, J3°42'W, 0:::2,000m, 5 V 1979, USNM 816150; I 't, 83 mm ML, I d, 56 mm ML, Sta. 390, 45°54'N, 13°30'W, 205 m, 5 V 1979, ZMH; 2 't't, 267, 77 mm GL, Sta. 391, 45°55'N, 13°27'W, 350 m, 5 V 1979, ZMH; 3 't't, 311-78 mm GL, I d, 54 mm ML, Sta. 398, 47°42'N, 09°08'W, 2,000 m, 6 V 1979, USNM 816151.-A. T. CAMERON: I d, 147 mm GL, Sta. 62-431, 43°30'N, 59°48'W, 596-841 m, 22 XI 1962, SJBS; I d, 92 mm GL, Sta. 149-2, 47°59'N, 44°52'W, 150-0 m, 6 VIII 1968, SJBS; 4 ?s, 77-53 mm ML, Sta. 149-5, 47°59'N, 44°52'W, 50-0 m, 6 VIII 1968, SJBS; I 't, 88 mm GL, I ?s, 42 mm ML, Sta. 347, 47°56'N, 44°46'W, 125-400 m, 4 VIII 1968, SJBS.-ATLANTIS: I d, 82 mm GL, I ?s, 69 mm ML, Sta. RHB-602, 39°41 'N, 71°IO'W, 57 m, 22 VII 1954, USNM 816 I 52.-BELOGORSK 7903: 6 ?s, 25-15 mm ML, EMT 31-1- 5, 39°44'N, 56°59'W, 50 m, I IV 1979, FOH; I ?s, 29 mm ML, EMT 41-2, 40017'N, 55°58'W, 100 m,6 IV 1979, FOH; I ?s, 22 mm ML, EMT 41-3, 40017'N, 55°58'W, 200 m, 6 IV 1979, FOH; I ?s, 39 mm ML, EMT 43-5, 38°50'N, 54°16'W, 500 m, 7 IV 1979, FOH; 5 ?s, 34-20 mm ML, EMT 46- 2, 40013'N, 54"07'W, 100 m, 9 IV 1979, FOH; 3 ?s, 27-14 mm ML, EMT 46-3, 40013'N, 54"07'W, 200 m, 9 IV 1979, FOH; 2 ?s, 21, 18 mm ML, EMT 46-5, 40013'N, 54°07'W, 500 m, 9 IV 1979, FOH; I d, 30 mm ML, EMT 47-3, 40058'N, 55"OO'W, 200 m, 9 IV 1979, FOH; I ?s, 27 mm ML, EMT 48-3, 41°39'N, 55°46'W, 200 m, 10 IV 1979, FOH; I ?s, 20 mm ML, EMT 54-I, 41°31'N, 54"04'W, 50 m, 13 IV 1979, FOH; I ?s, 37 mm ML, EMT 54-4, 41°31'N, 54°04'W, 300 m, J3 IV 1979, FOH; 2 ?s, 16-11 mm ML, IB-34-I(S), 41°51'N, 59°23'W, 50 m, 3 IV 1979, FOH.-BELOGORSK 7905: I 't, 72 mm GL, EMT 74-01, 42°II'N, 64°28'W, 50 m, 12 V 1979, FOH; I 't, 52 mm ML, I d, 63 mm ML, EMT 74-03, 42°II'N, 64°28'W, 200 m, 12 V 1979, FOH; I d, 46 mm ML, EMT 80- 3, 42°27'N, 62°37'W, 200 m, 15 V 1979, FOH; I 't, 76 mm GL, EMT 81-2, 42°41'N, 62°41'W, 100 m, 16 V 1979, FOH; I 't, 44 mm ML, 2 dd, 55, 47 mm ML, EMT 81-3, 42°40'N, 62°40'W, 200 m, 16 V 1979, FOH; I d, 64 mm ML, I ?s, 32 mm ML, EMT 84-2, 42°46'N, 61°71'W, 100 m, 17 V 1979, FOH; I ?s, 41 mm ML, EMT 84-3, 42°46'N, 61°11'W, 200 m, 17 V 1979, FOH; I d, 60 mm ML, EMT 86-2, 40039'N, 59°35'W, 100 m, 18 V 1979, FOH; I 't, 79 mm GL, EMT 86-3, 40039'N, 59°35'W, 200 m, 18 V 1979, FOH; I d, 59 mm ML, 1 ?s, 35 mm ML, EMT 87-3, 41°06'N, 59°42'W, voss: REVISIONOF TEUTHOWENIA 83

200 m, 19 V 1979, FOH; 2 aa, 65, 60 mm ML, EMT 89-3, 42°47'N, 60003'W, 200 m, 20 V 1979, FOH; I a, 61 mm ML, EMT 90-3, 42°39'N, 60001'W, 200 m, 20 V 1979, FOH; 2 aa, 67, 64 mm ML, EMT 91-3, 43°17'N, 60olO'W, 200 m, 21 V 1979, FOH.-BRANDAL: 2 \?!t, 265,230 mm ML, 2 aa, 190, 125 mm GL, Haul 3, 50030'N, 49°30'W, 1,000 m, 12 VII 1968, SJBS; 1 ~, 240 mm GL, 3 aa, 220-197 mm GL, Haul 5, 50030'N, 49°30'W, 1,050 m, 13 VII 1968, SJBS; I ~, 254 mm GL, Haul 6, 5O-30'N, 49°30'W, 1,050 m, 13 VII 1968, SJBS; 2 ~~, 310, 230 mm GL, 2 aa, 155, 146 mm ML, Haul 7, 49000'N, 48°30'W, 1,100 m, 14 VII 1968, SJBS; I a, 234 mm GL, Haul 8, 49°00'N, 48°30'W, 1,150 m, 14 VII 1968, SJBS; I ~, 311 mm GL, 3 aa, 200-131 mm ML, Haul 9, 49°00'N, 45°00'W, 990 m, 16 VII 1968, SJBS; I a, 145 mm GL, Haul 10, 49000'N, 45OW, 1,000 m, 16 VII 1968, SJBS; I ~, 309 mm GL, I a, 182 mm GL, I ?s, 43 mm ML, Haul II, 47°30'N, 43°00'W, 1,000 m, 17 VII 1968, SJBS; I ~, 210 mm GL, I a, 184 mm GL, Haul 13, 46'00oN, 44°30'W, 1,000 m, 18 VII 1968, SJBS; I a, 180 mm ML, Haul 14, 46°00'N, 44°30'W, 1,000 m, 18 VII 1968, SJBS; 2 ~~, 335, 270 mm GL, 4 aa, 212-103 mm GL, 1 ?s, 88 mm ML, Haul 15, 45°30'N, 47°30'W, 1,000 m, 19 VII 1968, SJBS; I ~, 200 mm ML, Haul 16, 45°30'N, 47°30'W, 1,000 m, 19 VII 1968, SJBS; 4 ~~, 352-218 mm GL, 2 <3d,184,85 mm GL, Haull7, 42°30'N, 50000'W, 1,000 m, 24 VII 1968, UMML 31.1845; I a, 144 mm ML, 2 ?8, 62, mm ML, fragments, Haul 19, 42°30'N, 51005'W, 50 m, 24 VII 1968, SJBS.- CRAWFORD:1 ~, 57 mm GL, 30014'N, 69°17'W, stomach of Alepisaurus sp., 24 IV 1961, UMML 31.362.-DANA: 1 ?s, 19 mm ML, Sta. 1349 II, 36°16'N, 74°33'W, 200 m, 21 V 1922, ZMC; I ?s, 43 mm ML, Sta. 1349 V, 36°16'N, 74°33'W, 50 m, 21 V 1922, ZMC; I ?s, 16 mm ML, Sta. 1349 VII, 17 m, 21 V 1922, ZMC; 1 ~, 96 mm GL, Sta. 1386 I, 45°15'N, 08°43'W, 500 m, 24 VI 1922, ZMC; I a, 136 mm GL, Sta. 1387 III, 46°28'N, 0800I'W, 2,000 m, 25 VI 1922, ZMC; 1 ~, 149 mm GL, Sta. 1387 IV, 1,500 m, 25 VI 1922, ZMC; I a, 79 mm GL, Sta. 1388 IV, 47°00'N, 06°33'W, 33 m, 26 VI 1922, ZMC; I a, 79 mm GL, Sta. 4158 I, 46°28'N, 0800 I'W, 83 m, 17 VI 1930, ZMC; I a, 93 mm GL, I ?s, 49 mm ML, Sta. 4158 III, 46°28'N, 08°01'W, 50 m, 17 IV 1930, ZMC; 1 ~, 105 mm GL, Sta. 4158 IV, 46°28'N, 0800I'W, 33 m, 17 VI 1930, ZMC; I a, 94 mm GL, Sta. 4158 XIV, 46°28'N, 08°01'W, 2,000 m, 18 VI 1930, ZMC; I ?s, 50 mm ML, Sta. 4158 XV, 46°28'N, 0800I'W, 1,500 m, 18 VI 1930, ZMC; 1 ?s, 46 mm ML, Sta. 4169 III, 47°53'N, 14°29'W, 100 m, 28 V 1931, ZMC; 1 ?s, 48 mm ML, Sta. 4169 V, 47°53'N, 14°29'W, 33 m, 28 V 1931, ZMC; I ?s, 37 mm ML, Sta. 4202 I, 47°36'N, 28°39'W, 100 m, 29 VI 1931, ZMC; I ?s, 39 mm ML, Sta. 4202 II, 47°36'N, 28°39'W, 67 m, 29 VI 1931, ZMC; 1 ~, 78 mm GL, Sta. 4205 II, 51°48'N, 30030'W, 200 m, I VII .1931, ZMC; 4 ?s, 40-7 mm ML, Sta. 4205 III, 51°48'N, 30030'W, 100 m, 1 VII 1931, ZMC; 13 ?s, 52-11 mm ML, Sta. 4205 IV, 51°48'N, 30030'W, 67 m, 1 VII 1931, ZMC; 3 ?s, 34-19 mm ML, Sta. 4205 V, 51°48'N, 30030'W, 33 m, 1 VII 1931, ZMC; 2 ?s, 55, 22 mm ML, Sta. 4205 VI, 51°48'N, 30030'W, 17 m, 1 VII 1931, ZMC; 1 a, 67 mm GL, Sta. 4206 I, 53°38'N, 29°41'W, 500 m, I VII 1931, ZMC; 8 ?s, 32-12 mm ML, Sta. 4206 IV, 53°38'N, 29°41'W, 33 m, I VII 1931, ZMC; 6 ?s, 52-13 mm ML, Sta. 4370 I, 59°52'N, 05°50'W, 50 m, 11 VI 1932, ZMC; 7 ?s, 48-11 mm ML, Sta. 4370 III, 59°52'N, 05°50'W, 22 m, II VI 1932, ZMC; I ?s, 15 mm ML, Sta. 4402 II, 62°23'N, 16005'W, 100 m, 25 VI 1932, ZMC; I ?s, 20 mm ML, Sta. 4402 IV, 62°23'N, 16°05'W, 17 m, 25 VI 1932, ZMC; I ?s, 36 mm ML, Sta. 4402 V, 62°23'N, 16°05'W, 2,000 m, 25 VI 1932, ZMC; 3 ?s, 77-13 mm ML, Sta. 4402 VII, 1,000 m, 25 VI 1932, ZMC; 3 ?s, 7-5 mm ML, Sta. 5083, 62°45'N, 1600I'W, 500 m, 7 V 1934, ZMC; 2 ?s, 7, 4 mm ML, Sta. 5083, 62°45'N, 16°01'W, 1,000 m, 7 V 1934, ZMC.- DELAWARE:1 a, 244 mm GL, 36°21'N, 74°21'W, stomach of Alepisaurusferox, 24 IV 1960, USNM 729989.-GERONIMO: 1 ~, 75 mm GL, 8 ?s, 53-34 mm ML, Sta. 1-16, 36°42'N, 74°28'W, 0-238 m, 17 V 1963, USNM 729980; 1 ~, 59 mm ML, 2 aa, 49, 41 mm ML, Sta. 1-17, 36°32'N, 74°27'W, 0- :::::128m, 20 V 1963, USNM 729976; 4 ~~, 61-52 mm ML, Sta.I-20, 36°37'N, 74°24'W, 0-:::::128 m, 20 V 1963, USNM 729979; 3 aa, 50-46 mm ML, Sta. 1-21, 36°40'N, 74°26'W, 0-300 m, 21 V 1963, USNM 729981; 1 ~, 59 mm GL, Sta. 1-23, 36°48'N, 74°28'W, 0-183 m, 21 V 1963, USNM 729982; 1 ~, 51 mm GL, 1 a, 60 mm GL, 2 ?s, 47, 34 mm ML, Sta. 1-24, 37005'N, 74°31'W, 0-420 m, 22 V 1963, USNM 729977; 2 ~~, 44, 35 mm ML, 2 aa, 51,44 mm ML, Sta. 1-26, 37°45'N, 74°26'W, 0- 130 m, 22 V 1963, USNM 729983; 3 ~~, 59-51 mm GL, 7 ?s, 47-40 mm ML, Sta. 1-27, 37°25'N, 74°29'W, 0-194 m, 22 V 1963, USNM 729978.-KNORR: I a, 130 mm GL, JEC 76-39, 38°49'N, 72°24'W, Deepwater Dumpsite 106,0-750 m, 5 IX 1976, USNM 816153.-0CEAN ACRE: 1 ii, 33 mm ML, Sta. 6-14C, 32°20'N, 63°51'W, 0-58 m, 28 IV 1969, USNM 727349; 1 ?s, 23 mm ML, Sta. 6-20N, 31°57'N, 63°46'W, 0-550 m, 29 IV 1969, USNM 727350; 1 ii, 43 mm ML, Sta. 6-24N, 32°13'N, 63°40'W, 0-750 m, 30 IV 1969, USNM 727348; 1 ?s, 8 mm ML, Sta. 13-2M, 31°50'N, 63°52'W, 0-40 m, 23 II 1972, USNM 728514.-PAWNEE III: 1 ?s, 33 mm ML, Sta. 59, 32°19'N, 64°33'W, 1,230 m, 21 IV 1927, YPM 12458.-PRINCESSE ALICE:2 ail, 141+, 136+ mm ML, Sta. 3318, 46°15'N, lOo11'W, :::::732m, 19 V 1911, MOM.-RoCKAWAY: 1 ii, 200 mm GL, Sta. 52, Wilmington Canyon, 0-2,654 m, 8 XII 1967, USNM 816154.- THOR: I a, 63 mm GL, 1 ?s, 52 mm GL, 46°30'N, 07000'W, 2,000 m, 3 IX 1906, ZMC.-WALTHER HERWIG:2 aa, 208,199 mm GL, Sta. 678/73, 65009'N, 32°50'W, 900-1,100 m, 19 IX 1973, USNM 816155; I ii, 215 mm GL, Sta. 695/ 73, 55°43'N, 25°53'W, 2,600 m, 22 IX 1973, ZMH; 1 ~, 191 mm GL, Sta.705/73, 50008'N, 19°46'W, 2,550 m, 25 IX 1973, ZMH; 1 a, 102 mm GL, Sta.706/73, 50007'N, 19°45'W, 490-500 m, 25 IX 1973, ZMH; 1 ~, 187 mm GL, Sta. 712/73, 47"04'N, 16°50'W, 85-100 m, 26 IX 1973, ZMH; 2 ~~, 84 BULLETINOFMARINESCIENCE,VOL.36, NO.1. 1985

223, 154 mm GL, 3 dd, 189-143 mm GL, Sta. 714/73, 46°00'N, 15°49'W, 2,650 m, 27 IX 1973, USNM 816156.-1 ?s, 63± mm ML, SR 1175, 51°38'N, 11056'W, ""732 m, 19 V 1911, BMNH 1912.3.19.1. Teuthowenia maculata.-ATLANTIS II: 2 ?s, 27, 19+ mm ML, Sta. RHB 2036, 21°46'N, 18°01'W, 45-55 m, II XI 1970, USNM 816157; I d, 52 mm GL, Sta. RHB 2047, 19°13'N, 18°17'W, 580-620 m, 12 XI 1970, USNM 816158; I Q, 70 mm GL, Sta. RHB 2061, 16°20'N, 21°33'W, 115-125 m, 15 XI 1970, USNM 816159.-DANA: 1 ?s, 7 mm ML, Sta. 3998 II, 07"34'S, 08°48'W, 200 m, I III 1930, ZMC; I ?s, 47 mm ML, Sta. 3999 III, 03°45'S, 10000'W, 100 m, 2 III 1930, ZMC; 1 ?s, 17 mm ML, Sta. 4003 X, 08°26'N, 15°11'W, 33 m, 9 III 1930, ZMC; 2 ?s, 36, 33 mm ML, Sta. 4004 I, 10°21'N, 17°59'W, 500 m, II III 1930, ZMC; I ?s, 36 mm ML, Sta. 4004 III, 10021'N, 17°59'W, 100 m, II III 1930, ZMC; I Q, 54 mm GL, I ?s, 42 mm ML, Sta. 4005 I, 13°31'N, 18"03'W, 2,000 m, 12 III 1930, ZMC; I d, 64 mm GL, Sta. 4005 II, 13°31'N, 18"03'W, 1,750 m, n III 1930, ZMC; 2 QQ, 55 mm GL, 41 mm ML, Sta. 4005 III, 13°31'N, 18°03'W, 1,500 m, 12 III 1930, ZMC; 2 QQ, 57, 50 mm GL, I d, 53 mm GL, Sta. 4005 IV, 13°31'N, 18"03'W, 1,250 m, 12 III 1930, ZMC; I d, 51 mm GL, Sta. 4005 V, 13°31'N, 18°03'W, 1,000 m, 12 III 1930, ZMC; I ?s, 34 mm ML, Sta. 4005 VII, 13°31'N, 18"03'W, 500 m, I2 III 1930, ZMC; I ?s, 43 mm ML, Sta. 4005 VIII, 13°31'N, 18"03'W, 200 m, 12 III 1930, ZMC; 5 ?s, 27-6 mm ML, Sta. 4005 X, 13°31'N, 18"03'W, 33 m, 12 III 1930, ZMC; 2 ?s, 8, 5 mm ML, Sta. 4006 IV, 15°31'N, 18"05'W, 33 m, 13 III 1930, ZMC; 1 ?s, 42 mm ML, Sta. 4007 I, 18°22'N, 18°14'W, 500 m, 15 III 1930, ZMC; I ?s, 10 mm ML, Sta. 4007 III, 18°22'N, 18°14'W, 100 m, 15 III 1930, ZMC; I d, 55 mm GL, Sta. 4007 IV, 18°22'N, 18°14'W, 33 m, 15 III 1930, ZMC; I d, 56 mm ML, Sta. 4007 VI, 18°22'N, 18°14'W, 2,000 m, 15 III 1930, ZMC; I d, 77 mm GL, Sta. 4007 X, 18°22'N, 18°14'W, 1,000 m, 15 III 1930, ZMC.-DISCOVERY: I ?s, 33 mm ML, Sta. 268, 18°37'S, 10046'E, 0-73 m, 25 VII 1927, lOS; I Q, 45 mm GL, Sta. 269, 15°55'S, 10035'E, 600-700 m, 26 VII 1927, lOS; I ?s, 45 mm ML, Sta. 285, 02°44'S, 00057'W, 125-175 m, 16 VIII 1927, lOS; I ?s, 28 mm ML, Sta. 6662-2, lioN, 200W, 100-200 m, 14 II 1968, MBA; 3 ?s, 20-15 mm ML, Sta. 6662-3, lIoN, 200W, 100 (--0) m, 14 II 1968, MBA; I ?s, 42 mm ML, Sta. 6662-11, lioN, 200W, 510- 590 m, 15 II 1968, MBA; 1 ?s, 53 mm ML, Sta. 6662-15, lIoN, 200W, 600-695 m, 16 II 1968, MBA; I ?s, 48 mm ML, Sta. 6662-23, lIoN, 200W, 320-415 m, 18 II 1968, MBA; 2 ?s, 13,6 mm ML, Sta. 6665-30, lIoN, 200W, 50-53 m, 26 II 1968, MBA; I ?s, 56 mm ML, Sta.7824-22, lIoN, 200W, 610- 700 m, 9 III 1972, MBA; I ?s, 18 mm ML, Sta. 7824-56, lIoN, 200W, 20-60 m, 12 III 1972, MBA; 2 ?s, 16, 10 mm ML, Sta. 7824-73, lIoN, 200W, 0-25 m, 14 III 1972, MBA.-GALATHEA: 2 ?s, 10, 8 mm ML, Sta. 8, 16°58'N, 18°16'W, 133-600 m, 3 XI 1950, ZMC.-WALTHER HERWIG: I d, 143 mm GL, Sta. 498 III/7I, 17°27'N, 22°55'W, 596-610 m, 17 IV 1971, USNM 729962. Teuthowenia pellucida.-ANTON BRUUN: I ?s, 36 mm ML, Sta. 3-157 (7097), 32°II'S, 59°30'E, 0- 275 m, 8 IX 1963, USNM 816161.-DISCOVERY: 4 ?s, 35-20 mm ML, Sta. 101, 34°13'S, 15°49'E, (0-) 1,310-1,410 m, 15 X 1926, lOS; I ?s, 30± mm ML, Sta. 101, 34°13'S, 15°49'E, (0-) 2,480-2,580 m, 14-15 X 1926, lOS; I ?s, 18 mm ML, Sta. 254, 35"04'S, 03°00'E, (0-) 200 m, 21 VI 1927, lOS; I ?s, 22 mm ML, Sta. WS 406, 56°5I'S, 67"03'W, 0-90 m, 24 II 1929, lOS; 113, 103 mm GL, Sta. WS 601, 35°31'S, 74°18'W, (0-) 230 m, 20 V 1931, lOS; I ?s, 32 mm GL, Sta. 968, 42°30'S, 124°52'W, 0 0-86 m, 19 IX 1932, lOS; I Q, 183 mm ML, Sta. 1763, 32"04'S, 40 45'E, (0-) 2,000 m, 5 V 1936, MBA.-ELTANIN: 113,113 mm ML, Sta. 4-99, 5 1043'S, 77°36'W, 1,208-1,219 m, 12 VII 1962, UMML 3I.l848; I Q, 75 mm GL, I ?s, 32 mm ML, Sta. 4-190, 35"05'S, 74°29'W, 2,891 m, 30 VIII 1962, UMML 3I.l849; I ?s, 14 mm ML, Sta. 5-213, 41°IO'S, 75"OI'W, 606 m, 13 IX 1962, USNM 816162; I Q, 70 mm ML, Sta. 6-326, 38°30'S, 74°37'W, 1,691-2,072 m, 25 XI 1962, USNM 816163; I d, 94 mm GL, Sta. 16-1405, 44°18'S, 162"OO'E,200 m, 3-4 II 1965, USNM 816164; 113,98 mm GL, Sta. 16-1409, 49"OO'S, 162°00'E, 350 m, 2 VI 1965, USNM 816165; 2 ?s, 21,15 mm ML, Sta. 21-222, 35"05'S, 91°59'W, 225 m, 3 XII 1965, USNM 816166; I Q, 184 mm GL, Sta. 23-1697, 46°22'S, 0 170 14'W, 1,813-2,274 m, 19-20 V 1966, UMML 3I.l850; I Q, 92 mm ML, 113, 110 mm ML, Sta. 23-1704, 43°45'S, 174°19'W, 725-800 m, 22 V 1966, USNM 816167; 113,75 mm GL, Sta. 23-1710, 41°45'S, I 78"05'W, 750-900 m, 25 V 1966, USNM 816168; I ?s, 5 mm ML, Sta. 24-1764, 38"08'S, 134°41'W, 460 m, 9 VIII 1966, USNM 816169; I ?s, 7 mm ML, Sta. 24-1776, 41°50'S, 1300n'w, 1,250 m, 14 VIII 1966, USNM 816170; I ?s, 6 mm ML, Sta. 24-1777, 41°46'S, 130016'W, 440 m, 14 VIII 1966, USNM 816171; I ?s, 5 mm ML, Sta. 24-1794, 39°46'S, 126°45'W, 160 m, 20 VIII 1966, USNM 816172; 3 ?s, 10-5 mm ML, Sta. 24-1803, 39°36'S, 125°19'W, 600 m, 24 VIII 1966, USNM 816173; I ?s, 39 mm ML, Sta. 25-302, 33°11 'S, 72°40'W, 185 m, 25 IX 1966, USNM 816174; 2 ?s, 120±, 30 mm ML, Sta. 26-1820, 40026'S, 168°30'E, 600-750 m, 2 XII 1966, USNM 816175; 10 ?s, 94-47 mm ML, Sta. 26-1830, 42°08'S, 160"05'E, 700-800 m, 8 XII 1966, USNM 816176; I 13,78 mm GL, 5 ?s, 61-51 mm ML, Sta. 26-1831, 42°19'S, 160"06'E, 30-60 m, 9 XII 1966, UMML 31.1851; 1 ?s, 41 mm ML, Sta. 26-1832, 42°22'S, 160007'E, 115-130 m, 9 XII 1966, USNM 816177; 6 ?s, 57-13 mm ML, Sta. 26-1834, 45°19'S, 160011'E, 700-780 m, II XII 1966, USNM 816178; I 2,82 mm ML, I ?s, 29 mm ML, Sta. 26-1835, 45°27'S, 160017'E, 1,250-1,375 m, 11 XII 1966, USNM 816179; 3 ?s, 63-16 mm ML, Sta. 26-1836, 45°38'S, 160012'E, 2,030-2,181 m, II XII 1966, USNM 816180; 2 22, Ill, 70 mm GL, 113,119 mm GL, I ?s, 47 mm ML, Sta. 26-1838, 45°52'S, voss: REVISIONOF TEUTHOWENIA 85

160"07'E, 190-230 m, 11-12 XII 1966, USNM 816181; 6 ?s, 68-6 mm ML, Sta. 26-1840, 47°20'S, 161°54'E, 25-125 m, 12 XII 1966, USNM 816182; 2 ?s, 37, 12 mm ML, Sta. 26-1841, 47°28'S, 161°52'E, 700-800 m, 12-13 XII 1966, USNM 816183; 2 ?s, 32,18+ mm ML, Sta. 35-2270, 45"05'S, 128"04'E, 250 m, 1 IX 1968, USNM 816184; 2 ?s, 21, 19 mm ML, Sta. 35-2281, 38°59'S, I 17"05'E, 200 m, 12 IX 1968, USNM 816185; 2 ?s, 28, 10 mm ML, Sta. 35-2302, 43°25'S, 129°25'E, 440 m, 2-3 X 1968, USNM 816186; 1 ?s, 5 mm ML, Sta. 46-254, 35°21'S, I 14°59'E, 500 m, 22 XI 1970, USNM 816187; I ?s, 23 mm ML, Sta. 46-261, 35°18'S, 114°58'E, 180 m, 20 XI 1970, USNM 816188; I ?s, 7 mm ML, Sta. 46-273, 38°24'S, 115"06'E, 550-600 m, 24 XI 1970, USNM 816189.-GALATHEA: 1 ?s, 22 mm ML, Sta. 601, 45°51'S, 164°32'E, 3,750 m, 14 I 1952, ZMC; 3 ?s, 88-67 mm GL, Sta. 629, 41°46'S, 175°48'E, 850 m, 24 11952, ZMC.-KAPALA: 200,91, 88± mm GL, 1 ?s, 67 mm GL, Sta. K 77-18-01, 34°10'S, I 52"04'E, 0-660 m, 20 X 1977, AM CI08567; 2 QQ, 83, 81 mm GL, 1o, 94 mm GL, Sta. K 77-18-05, 35°37'S, 150056'E, 0-650 m, 27 X 1977, AM CI08566; I ?s, 61 mm GL, Sta. K 77-19-02, 37004'S, 150020'E, 0-"'150 m, I XI 1977, AM CI09363; 1 Q, 139 mm GL, 1 0, 102 mm ML, Sta. K 77-24-09, 33°20'S, 152°32'E, 0-"'300 m, 14 XII 1977, AM CI08661; 2 QQ, 153, 112 mm GL, 20o, 117,95 mm GL, Sta. K 77-24-10, 33°28'S, 152°33'E, 0-'" 1,000 m, 14 XII 1977, AM CI08662; I Q, 103 mm GL, 1 0, 97 mm ML, Sta. K 77-24-11, 33°53'S, I52°02'E, 0-"'800 m, 14 XII 1977, AM CI08663.-MoNSOON: 1 ?s, 47 mm ML, Sta. 9, 33°38'S, 72°31'E, 1,878 m, 19 XII 1960, SIO.-PRAWN SURVEY,BAYOF PLENTY:I ?s, 72 mm ML, Haul 10, Bay of Plenty, 5 mi N Major Island, 0-366 m, 28 IX 1962, DMNZ.-SEA FISHERIES,SA: I ?s, 4 mm ML, Sta. F2525-44-3, 34°17'S, 17"40'E, surface, 5 VII 1963, SAM; 1 ?s, 22 mm ML, Sta. F253 1-40-4, ca. 34"07'S, 14°09'E, surface, 7 VII 1963, SAM; 1 ?s, 5 mm ML, Sta. A2597-36-3, 33°40'S, 17°15'E, 0-150 m, 18 VIII 1963; SAM; I Q, 85 mm GL, Sta. A31421, 32°40'S, 15°44'E, 0-5 m, 14 X 1964, SAM; I ?s, 12 mm ML, Sta. A4217-WR-l, 29°22'S, 13°09'E, 0-5 m, 7 VII 1966, SAM; 1 Q, 90 mm GL, 3 ?s, 63-55 mm ML, Sta. A4981-WS-3, 35°02'S, 27°51'E, 7 IX 1968, stomach of lancet fish, SAM.-SHOYO MARU: 1 0, 114 mm GL, Sta. 12-F25, 33°17'S, 94°25'W, 27 I 1963, stomach of Thunnus obesus, UMML 31.1852; 1 0, 95 mm GL, 2 ?s, 82, 37 mm ML, Sta. 12-F28, 36°59'S, 90005'W, 30 I 1963, stomach of Alepisaurus sp., UMML 31.1853; I 0, 89 mm GL, 4 ?s, 75-60 mm ML, Sta. 13-F4, 32°51'S, 123°27'W, 7 XI 1963, stomach of Alepisaurus borealis, UMML 31.1854; 4 ?s, 98-65 mm ML, Sta. 13-F5, 35°55'S, 116°53'W, 9 XI 1963, stomach of Alepisaurus sp., UMML 31.1855; 2 QQ, 101,74 mm GL, 2 ?s, 68, 68 mm ML, Sta. 13-F6, 37°11'S, I 14°41'W, 10 XI 1963, stomach of Alepisaurus sp., UMML 31.1856; 1 ?s, 84 mm ML, Sta, 13-F7, 40009'S, 109°38'W, 12 XI 1963, stomach of Alepisaurus sp. or albacore, UMML 31.1857; 2 ?s, 76, 68 mm ML, Sta. 13-F8, 39°12'S, 106°48'W, 13 XI 1963, stomach of Alepisaurus sp., UMML 31.1858; 2 ?s, 77, 76 mm ML, Sta. 13-F9, 38°25'S, 10 I"06'W, 15 XI 1963, stomach of Alepisaurus sp. or albacore, UMML 31.1859; 2 ?s, 104, 88 mm ML, Sta. 13-FIO, 34°48'S, 101°18'W, 17 XI 1963, stomach of Alepisaurus sp., UMML 31.1860; 1 ?s, 79 mm ML, Sta. 13-FI2, 32°13'S, 86°45'W, 21 XI 1963, stomach ofAlepisaurussp., UMML 31.1861.- WALTHERHERWIG:2 QQ, 125, 114 mm GL, 10, 102 mm GL, Sta. 3541/71, 39°18'S, 48°12'W, 100 m, 6 III 1971, USNM 816190; 5 QQ, 201-154 mm GL, 5 00,191-103 mm GL, fragments of21arge mantles, Sta. 354 lI/7l, 39°19'5, 48°02'W, 2,000 m, 6 III 1971, USNM 816191; 1 Q, 81 mm GL, Sta. 35811/71, 39°47'S, 43°38'W, 200-212 m, 7 III 1971, USNM 816192; I Q, 88 mm ML, 1 <3,127 mm ML, Sta. 358 III/71, 39°47'S, 43°30'W, 1,000-1,015 m, 7 III 1971, ZMH; 1 Q, 74 mm ML, Sta. 363 1/71, 40018'S, 39°22'W, 105-110 m, 8 III 1971, ZMH; 3 QQ, 143 mm GL, 128, 112 mm ML, 20o, 138,127 mm GL, Sta. 367/71, 40018'S, 35°07'W, 2,000 m, 9 III 1971, USNM 816193; 2 QQ, 154 mm GL, 87 mm ML, 20o, 161 mm GL, 91 mm ML, Sta. 376/71, 39°55'S, 26°02'W, 2,000 m, II III 1971, USNM 816194; 10,80 mm ML, Sta. 38011/71, 39°53'S, 21°44'W, 260 m, 12 III 1971, ZMH; 10,89 mm ML, Sta. 380 III/71, 39°53'S, 21°33'W, 696-708 m, 12 III 1971, ZMH; 1 Q, 154 mm GL, Sta. 395/71, 36°49'S, 12°17'W, 1,750-2,000 m, 17 III 1971, ZMH; 10,69 mm ML, Sta. 402 III/7I, 40"OI'S, 07°25'W, 800-820 m, 18 III 1971, ZMH; 2 QQ, 147 mm GL, 96 mm ML, 10,166 mm GL, Sta. 406/71, 39°13'S, 03°15'W, 2,000 m, 19 III 1971, USNM 816195; 2 QQ, 177, 154 mm GL, 1o, 141 mm GL, Sta. 417/71, 34°12'S, 16°35'E, 1,550 m, 28 III 1971, USNM 816196; 1 Q, 125 mm GL, Sta. 419 11/71, 34°26'S, 14°43'E, 300-305 m, 28 III 1971, USNM 816197; 13 QQ, 80-43 mm ML, 6 00,66-48 mm ML, 2 ?s, 57, 45 mm ML, Sta. 2/75-76, 36°19'S, 49°35'W, 800 m, 18 XI 1975, USNM 0 816147; 2 QQ, 68, 66 mm ML, Sta. 3/75-76, 37°49'S, 50 06'W, 200-218 m, 18 XI 1975, ZMH; 1o, 50 mm ML, Sta. 4 1/75-76, 40012'S, 50002'W, 620-650 m, 19 XI 1975, ZMH; 2 QQ, 77, 64 mm ML, 50o, 143 mm GL, 73-62 mm ML, Sta. 4 11/75-76, 40020'S, 50"02'W, 2,000-2,350 m, 19 XI 1975, ZMH; I Q, 72 mm ML, 10,58 mm ML, Sta. 5/75-76, ""100 m, 19 XI 1975, ZMH; I Q, 179 mm 0 GL, 10,201 mm GL, Sta. 10111/75-76, 47°45'S, 40 05'W, 2,000 m, 5 I 1976, USNM 816198; I Q, 82 mm GL, 3 00, 118-92 mm GL, Sta. 105/75-76, 41°47'S, 39°58'W, 210-200 m, 7 I 1976, ZMH; I Q, 74 mm ML, Sta. 1061/75-76, 39°39'S, 39°56'W, 800 m, 8 I 1976, ZMH; 2 QQ, 176, 130 mm GL, 10,186 mm GL, Sta. 10611/75-76, 39"08'S, 40000'W, 1,800-1,850 m, 8 I 1976, USNM 816199; I Q, 189 mm GL, I 0,63 mm GL, Sta. 108 11/75-76, 36°18'S, 40"06'W, 2,000-2,400 m, 9 I 1976, USNM 816200.-1 Q, 106+ mm GL, Sta. VUZ 84, 41°47'S, 175°02'E, 0-"'732 m, 19 IV 1957, DMNZ.-I Q, 67 mm GL, olfCampbell Island, N.Z., call. D. Nightingale, DMNZ.-I ?s, 57 mm GL, 33°13'S, I78°24'E, (0-) 641-824 m, 2 VII 1962, DMNZ.

&lt;I&gt;Teuthowenia&lt;/I&gt; (Oegopsida)

<I>Teuthowenia</I> (Oegopsida)