THE OPLOPHORID and PASIPHAEID SHRIMP from OFF the OREGON COAST Redacted for Privacy Abstract Approved (Major Professor)

AN ABSTRACT OF THE THESIS OF

Carl Albert Forss for the Ph. D. in Zoology (Name) (Degree) (Major) 1q1:1) Date thesis is presented l k` -} 1.-(2.t 1 Title THE OPLOPHORID AND PASIPHAEID SHRIMP FROM OFF THE OREGON COAST Redacted for Privacy Abstract approved (Major Professor)

The 13 species of shrimps studied for this thesis were collected off the Oregon coast. The family Oplophoridae is well represented in this area. Five of the seven known genera were identified.

Hymenodora frontalis, H. glacialis, and H. gracilis were described and further differentiating characters were illustrated. Other mem- bers of the family Oplophoridae were illustrated and described as follows: Acanthephyra curtirostris, Meningodora mollis, Notostomus japonicus, Systellaspis braueri and S. cristata. Species of the family Pasiphaeidae, although less well represented, were identified. The presence or absence of the mandibular palp in these two species, Parapasiphaë sulcatifrons and P. cristata, found to be of little systematic value in separating them. Specimens with a carapace length of 7- 10mm lack a mandibular palp; one with a carapace length of 17 -25mm had a two -segmented palp. Pasiphaea pacifica, P. magna, and P. chacei were identified and illustrated. The taxonomy, geographical distribution, color and luminescence are discussed. THE OPLOPHORID AND PASIPHAEID SHRIMP FROM OFF THE OREGON COAST

CARL ALBERT FORSS

A THESIS

submitted to

OREGON STATE UNIVERSITY

in partial fulfillment of the requirements for the degree of

DOCTOR OF PHILOSOPHY

June 1966 APPROVED: Redacted for Privacy

Professor of Zoology In Charge of Major

Redacted for Privacy

Chairman of Zoology Department

Redacted for Privacy

Dean of Graduate School

Date thesis is presented ¿,C.,C-( .u%C ; . 9'612'

Typed by Marcia Ten Eyck ACKNOWLEDGEMENT

The present study has been made possible by a NSF Grant No GB

1588; U. S. Atomic Energy Commission Contracts ÁT(45- 1)1726 and

ÁT(45- 1)1750, administered by Dr. William G. Pearcy, Department of Oceanography, Oregon State University. My sincerest thanks are extended to Dr. Pearcy.

A special word of thanks is tendered Dr. John C. Yaldwyn, of the Australian Museum, Sidney. He most graciously made many essential identifications and added helpful remarks to his identifications of the troublesome species.

I wish to thank Dr. Ivan Pratt, my major Professor of the

Department of Zoology, Oregon State University, for space, facilities and advice. TABLE OF CONTENTS

Page

INTRODUCTION 1

METHODS . . 2

SYSTEMATIC DISCUSSION 3

General Considerations 3 Key to the Oregon Genera of Pasiphaeidae 6 Key to the Oregon Species of Pasiphaea 6

Pasiphaea pacifica 7 Pasiphaea magna 8 Pasiphaea chacei 9 Parapasiphaë sulcatifrons 11 Parapasiphaë cristata 13

Key to the Genera of Oplophorids off Oregon 16 Acanthephyra curtirostris 17

Meningodora mollis . . 19 Notostomus japonicus . 21

Key to the Oregon Species of Hymenodora 23 Hymenodora frontalis .. 23 Hymenodora glacialis . 25 Hymenodora gracilis 27

Key to the Species of Systellaspis 29 Systellaspis braueri 30 Systellaspis cristata 32

GENERAL DISCUSSION AND SUMMARY 34

Taxonomy 34 Geographical Distribution 35 Color of Bathypelagic Caridea 36 Luminescence 37 TABLE OF CONTENTS Continued Page

BIBLIOGRAPHY 38

APPENDIX . . 43 THE OPLOPHORID AND PASIPHAEID SHRIMP FROM OFF THE OREGON COAST

INTRODUCTION

In 1961 the Department of Oceanography, Oregon State University,

undertook a long term ecological study of the Oregon coast. Out of

this study were obtained several unusual bathypelagic Caridea and

Penaeidea taken between 0- 1000m.

The topography and physical characteristics of the near -shore

waters probably are responsible in part, for the distribution of the

shrimps to be discussed.

Vertical migration among decapods has been known for some time

to occur (Murray and Hjort 1912, Chace 1940 and Bainbridge 1961).

Vertical migration also occurs for the shrimps taken off the Oregon

coast. Of the forms considered herein, tows to 200m caught predom- inately a carid shrimp, Pasiphaea pacifica. In tows to 500m P. pacifica was nearly absent, Hymenodora frontalis was taken quite frequently. Even at depths to 1000m almost no P. pacifica were taken, although H. frontalis was common. The few day tows that were taken brought up only a limited number of shrimps, indicating that these species avoid the surface waters during daylight by moving into deeper waters. 2

METHODS

The nektonic organisms were sampled with a six -foot Isaacs -

Kidd midwater trawl (Isaacs and Kidd, 1953). Most of the sampling

was done in the upper 0 -200m depth. The sampling was done at

night except for a few tows taken during daylight hours. The net was

lowered rapidly for tows from 0 -200m and then retrieved at a con-

stant speed of 30m per minute. For tows to 500m the trawl was

fished between 500 and 200m for about 30 minutes, and for tows to

1000m the trawl was fished between 1000 and 500m for one hour.

The net was open at all depths since no opening - closing device was

used.

All the collections were made from the R/V Acona. Records for all collections were made such as geographic position at the start and end of each tow, towing time, speed and environmental conditions.

Samples were preserved aboard ship in 10% formalin. Identification was in the laboratory ashore. Drawings were made with the aid of a

Wild's camera lucida and a Bio- scope. 3

SYSTEMATIC DISCUSSION

GENERAL CONSIDERATIONS:

The Decapoda (Crustacea) is divided into Reptantia and Natantia.

The latter are the shrimps, which characteristically have a large

antennal scale and five pairs of biramous pleopods on the abdomen.

The abdomen is chitinized, with pleura and a complete tail -fan. In

the natantians (swimmers) the antennal scale is generally large and

lamellar; five pairs of characteristically biramous (rarely uniramous)

pleopods present, always well developed and used for swimming; abdomen always macrurous (i. e. symmetrical, well chitinized, with pleura and complete tail -fan). Shrimps and prawns.

The suborder Natantia is divided into three sections: Caridea, Stenopodidea and Penaeidea. These sections usually have been termed tribes. This term, however, cannot be used for them, since during the 14th International Congress of Zoology at Copenhagen, 1953, it was decided that the term tribe is to be used only for units below the subfamily level.

The three sections may be distinguished as follows:

la. Pleurae of second abdominal somite overlapping those

of first and third segments. No chelae on the third

pereiopods. Gills phyllobranchiate . . . . Caridea 4

b. Pleurae of second abdominal somite not overlapping

those of first segment. Third legs with a chela.

2a. Third leg distinctly stronger than the preceding. Males without petasma. Gills trichobranchiate. Stenopodidea

b. Third leg never stronger than the preceding, generally

all chelipeds of equal strength. Males with petasma. Gills dendrobranchiate Penaeidea

The carapace in all cases is measured from its dorsal posterior margin to the posterior margin of the eye orbit.

In order to determine sex in the Caridea it was necessary to observe the inner ramus of the first pleopod in the family Oplo- phoridae. The first pleopod in the males is modified into the appendix masculina, the inner ramus of pleopods II through V are nearly the same in appearance. In the family Pasiphaeidae it was necessary to observe the inner ramus of the second pleopod, which was the loca- tion of the appendix masculina. The inner ramus of all pleopods in the female are similar and their margins edged with long setae.

Order Decapoda Suborder Natantia Section Caridea 5

Superfamily Pasiphaeoida Family Pasiphaeidae

Rostrum a normal forward - directed prolongation of the carapace or absent; if absent, it is represented by an erect postfrontal spine.

Fingers of all four chelae slender with their cutting edges pectinate.

Exopods present on all pereiopods; reduced or absent on second maxilliped. Mandible without a molar process and with or without a mandibular palp.

REMARKS:

The family Pasiphaeidae includes at present seven genera, two of which have been taken off the Oregon coast; Pasiphaea Savigny

(1816) and Parapasiphaë Smith (1884). Pasiphaea is a cosmopolitan genus, the 36 species except P. chacei Yaldwyn having been found in almost all oceans of the world. The above exception is a newly described species. Two of the three species, P. pacifica Rathbun and P. chacei, are bathypelagic, taken in depths to 1000m. P. magna

Faxon has been taken only by otter trawl at depths of greater than

2000m.

Of the seven species of Parapasiphaë, two have been taken off the Oregon coast; P. cristata Smith and P. sulcatifrons Smith. This is the first record of these species occurring in the Pacific Ocean. 6

The only other species of this genus taken in the Pacific Ocean is

P. serrata Rathbun.

KEY TO THE OREGON GENERA OF PASIPHAEIDAE:

1. Rostrum a post - frontal spine; no mandibular palp. Pasiphaea

2. Rostrum a normal forward projection, arising from the

frontal margin of the carapace; mandibular palp present in mature forms Parapasiphaë

Genus Pasiphaea Savigny, 1816

Rostrum represented by a post -frontal projecting spine. Orbits poorly defined. Mandibule without a palp.

KEY TO THE OREGON SPECIES OF PASIPHAEA:

la. Telson forked or notched; carapace arising into a sharp pointed tooth above the anterior gastric region; branchio-

stegal spine not reaching anterior margin of carapace. P pacifica

b. Telson truncate; rostrum projecting beyond anterior margin of carapace; branchiostegal spine reaches anterior margin

of carapace 2a 7

2a. Telson terminating in four pairs of spines; rostrum long

and slender; branchiostegal spine projects well beyond the

anterior margin of the carapace P. chacei

b. Telson terminating in six pairs of spines and a small medial unpaired spine; rostrum heavy, triangular in shape; branchiostegal spine reaches anterior portion of carapace. P magna

Pasiphaea pacifica Rathbun, 1902

Pasiphaea pacifica Rathbun, 1902, p. 905. Pasiphaea pacifica Rathbun, 1904, p. 20, fig. 2.

Pasiphaea pacifica Richardson, 1908, p. 692, fig. 4. Pasiphaea pacifica Richardson and Yaldwyn, 1958, p. 26, fig. 12.

Pasiphaea pacifica McCauley, 1962, p. 867, fig. 1, 2.

CHARACTERISTICS:

Carapace (fig. 1) ridged dorsally throughout its length except on the most posterior portion; lateral gastro - orbital carina (supra - branchial) present; branchiostegal spine supported by a short carina. Eyes large, dark brown. Antennal scale (fig. 2) tipped with a terminal lateral spine. Abdominal segments III through VI are carinated on the median line. Telson (fig. 3) with a deep notch, bordered by spinules, which increase in size and are continued to the posterior 8 outer angle of each lobe; the outer spinule is considerably longer than the others.

Mandible (fig. 4) without a palp; proximal ramus of first maxilla bearing two to three setae and half the length of the distal segment.

The latter with one long setae and several short ones.

DISTRIBUTION:

Taken from Unalaska and the Gulf of Alaska south to the Gulf of

California, Australia, far Eastern Seas of U. S. S. R. Also is known from Cape Natal, South Africa, in 106 to 880m of water. Off the

Oregon coast specimens have been taken to depths of 1000m.

Pasiphaea magna Faxon, 1893

Pasiphaea magna Faxon, 1893, p. 209.

Pasiphaea magna Faxon, 1895, p. 176, pl. 45, fig. 2, 2c.

Pasiphaea magna Rathbun, 1904, p. 18, fig. 1.

CHARACTERISTICS:

Carapace (fig. 5) carinated throughout its length except at its most posterior portion; lateral surface of carapace with a supra - branchial carina and a weak sub -hepatic carina posterior to the branchiostegal spine; branchiostegal spine reaching the frontal margin. Rostrum triangular in shape. Eyes large and heavily pigmented. 9

Antennal scale (fig. 6) tipped with a short, triangular spine.

Abdomen is carinated on abdominal segments II through VI.

Telson (fig. 7) dorsally sulcate and armed with 13 terminal spines.

Mandible (fig. 8) without a palp; proximal ramus of first maxilla less than half the length of the distal ramus and tipped with one seta.

First pereiopod with 4 -6 meral spines; second pereiopod with 10 -16 meral spines.

DISTRIBUTION:

Taken from off Point Arena, California, to the Gulf of Panama at depths of 530 to 1104m. Off Oregon specimens have been taken to depths of 2000m.

Pasiphaea chacei Yaldwyn, 1962

Pasiphaea chacei Yaldwyn, 1962, p. 15, figs. 1 -19.

CHARACTERISTICS:

Carapace (fig. 10) carinated dorsally except on its posterior one- third; suprabranchial carina present; branchiostegal spine origi- nating behind and passing well beyond the anterior margin; anterior frontal margin rounded from orbit to branchiostegal spine, below which the margin runs almost directly backward for a short distance, then turns abruptly downward and joins obliquely the infero- lateral 10

and reaching margin. Rostrum long and slender. Eyes dark brown 11) terminating in a distal to tip of rostrum. Antennal scale (fig. dorsally; relatively long narrow lateral spine. Abdomen noncarinated (fig. 12) truncate the sixth segment is compressed dorsally. Telson pair the longest. and armed with four pairs of spines, the lateral of first Mandible (fig. 13) without a palp; proximal segment with three maxilla half the length of the distal segment and provided outer ramus armed to four long setae and a few shorter ones; the ones. with one long mesial seta and several shorter pereiopod Fourth pereiopod shorter than the third or fifth; first with 7 -26 meral spines. with 1 -13 meral spines; second pereiopod or less show a Immature forms with a carapace length of 16mm carapace length. steady increase of meral spines with increasing of the mature The meral spines on the first and second pereiopods of 20 -22mm. forms reach a maximum number at a carapace length (1962) specimens had The first and second pereiopods of Yaldwyn's

0 -12 and 6 -23 meral spines respectively.

DISTRIBUTION:

taken in meso- This is the second record of this species being California by pelagic waters. The first is recorded off Southern

Yaldwyn (1962). 11

Parapasiphaë Smith, 1884

Rostrum arising from frontal margin. Fourth pereiopod shorter than fifth. Mandible with a two -jointed palp in mature forms.

Paraphasiphaë sulcatifrons Smith, 1884

Parapasiphaë sulcatifrons Smith, 1884, p. 384, pl. 5, fig. 4, pl. 6,

figs. 1-7.

Parapasiphaë sulcatifrons Hansen, 1908, p. 79.

Parapasiphaë sulcatifrons Kemp, 1910, p. 47, pl. 5, figs. 1 -21. Parapasiphaë sulcatifrons Coutiére, 1911, p. 157. Parapasiphaë sulcatifrons Stephensen, 1912, p. 70. Parapasiphaë sulcatifrons Murray and Hjort, 1912, p. 668. Parapasiphaë sulcatifrons Stephensen, 1923, p. 40. Parapasiphaë sulcatifrons Bals s, 1925. Parapasiphaë sulcatifrons Stephensen, 1935, p. 34. Parapasiphaë sulcatifrons Chace, 1940, p. 126, fig. 6. Parapasiphaë sulcatifrons Sivertsen and Holthuis, 1956, p. 30.

CHARACTERISTICS:

Dorsal carina of carapace (fig. 15) unarmed above, the anterior fourth is broadened and provided with a well marked groove; carapace lacking lateral spines; a single lateral carina passes posteriorly 12 from the region of the antennal scale and appears to trifurcate at about the post -hipatic region; well developed sub -marginal carina is present. Rostrum short, unarmed above and below, reaches nearly half the length of the eyes. Eyes (fig. 16) narrower than eye stalks; tubercle small. Antennal scale (fig. 17) with a short, blunt distal lateral spine. Abdomen smooth dorsally; fourth segment slightly compressed dorsally and with a posterior median spine; fifth somite sulcate dorsally. Telson (fig. 18) truncate armed with nine terminal spines; the marginal pair the longest.

Specimens with a carapace length of 25mm have a two- segmented mandibular palp (fig. 19), carapace length of 12mm have a one - segment palp and those with a carapace length of 7 -10mm have no mandibular palp.

Fingers of second chelae not longer than the palm; third pereiopod longer than the fifth; fourth shorter than the fifth.

DISTRIBUTION:

The species is most abundant in the North Atlantic from Green- land and Iceland southward and extends to about 35°S. It has also been reported from off the French Congo, South Africa, and the Southern

Indian Ocean according to Sivertsen and Holthuis (1956). The depths in which the species has been found range from 500- 5400m. This is the first record of this species occurring in the Pacific Ocean. This 13 species off Oregon has been taken in depths of 1000m.

Parapasiphaë cristata Smith, 1884

Parapasiphaë cristata Smith, 1884, p. 388, pl. 5, fig. 3.

Parapasiphaë macrodactyla Chace, 1940, p. 128, fig. 7.

CHARACTERISTICS:

Carapace (fig. 20) carinated throughout its length except on the posterior portion; dorsal carina of the rostrum is armed with one or two teeth above the orbit. Suprabranchial carina extending the full length of the carapace; four elevations with adjacent depressions pass off obliquely from the suprabranchial carina; well developed submarginal carina is present. Eyes narrower than the eye stalks; eye papillae extending beyond tips of eyes. Abdomen dorsally smooth throughout except that the fourth somite has a slight dorsal carina and the posterior margin projects dorsally over the fifth somite in a triangular tooth. Telson (fig. 21) truncate or slightly convexed; ending with nine spines; the lateral pair the longest, nearly twice as long as the others.

Specimens of a carapace length of 7 or 8 mm have no mandibular palp (fig. 22). Those with a carapace length of 10mm have only a bud present for a palp and those with a carapace length of 17mm have a two- segmented palp. Proximal segment of the first maxilla armed 14 with numerous setae and equal in length to the distal segment which is well armed with spines, outer ramus is setose and is provided with a weak carina on its inner margin.

DISTRIBUTION:

This species has been taken off the East Coast of North America,

Bermuda, West Coast of Cuba and from the North Atlantic. The depth at which this species has been taken varies between 1000- 3500m.

This species off Oregon has been taken in depths of 1000m. This is the first record from the Pacific Ocean. 15

Superfamily Oplophoroida Family Oplophoridae

Rostrum variable in shape. Exopods are present on all maxillipeds and pereiopods; no long exopods on the first maxilliped. A triangular terminal segment of the second maxilliped is attached to preceding one by a long oblique joint. Mandibles with a molar process, imperfectly cleft and provided with a palp.

REMARKS:

The family Oplophoridae includes at present seven genera, five of which have been taken off the Oregon coast; Acanthephyra A.

Milne Edwards (1881), Meningodora Smith (1882), Notostomus A.

Milne Edwards (1881), Hymenodora Sars (1887), and Systellaspis

Bate (1888).

Of the 22 described species of Acanthephyra only A. curtirostris which occurs in all oceans has been taken off the Oregon coast. How- ever, three new and undescribed species of Acanthephyra were taken.

All three species of Hymenodora were taken off our coast;

Hymenodora gracilis, H. glacialis and H. frontalis. The latter was the most frequently occurring species. 16

KEY TO THE GENERA OF OPLOPHORIDS OFF OREGON:

la. Sixth abdominal somite carinated 2

b. Sixth abdominal somite not carinated. 4

2a. Lateral surface of carapace lacking any straight carina

from orbit to posterior margin of carapace. . Acanthephyra

b. Lateral surface of carapace with one or more straight

carinae from orbit to posterior margin of carapace. . . 3

3a. Lateral surface of carapace with one straight carina from

. . orbit to posterior margin of carapace. . Meningodora

b. Lateral surface of carapace with more than one straight

carina from orbit to posterior margin of carapace. Notostomus

4a. Eyes small and lightly pigmented; telson terminating in

a truncate tip Hymenodora

b. Eyes very large, larger than eye stalks, and heavily pigmented; telson terminating in a sharp -pointed endpiece laterally armed with spines Systellaspis

Genus Acanthephyra A. Milne Edwards, 1881

Lateral surface of carapace usually lacking any straight carina from orbit to posterior margin of carapace. Abdomen carinate at least on the last four somites. Telson truncate to semi -truncate. 17

Eyes darkly pigmented. Mandible toothed for the entire surface of

its cutting edge.

Acanthephyra curtirostris Wood -Mason, 1891

Acanthephyra curtirostris Wood -Mason and Alcock, 1891, p. 195.

Acanthephyra curtirostris Wood -Mason and Alcock, 1892, p. 364,

fig. 5.

Acanthephyra curtirostris Faxon, 1895, p. 164, pl. 43, fig. 2 -5.

Acanthephyra curtirostris Alcock, 1901, p. 81.

Acanthephyra curtirostris Rathbun, 1904, p. 27.

Acanthephyra curtirostris Kemp, 1906, p. 23.

Acanthephyra curtirostris de Man, 1920, p. 66, pl. 6, fig. 15.

Acanthephyra curtirostris Schmitt, 1921, p. 33.

Acanthephyra curtirostris Balss, 1925, p. 261, fig. 30.

Acanthephyra curtirostris Chace, 1936, p. 26.

Acanthephyra curtirostris Chace, 1937, p. 111.

Acanthephyra curtirostris Chace, 1940, p. 143, fig. 21.

Acanthephyra curtirostris Chace, 1947, p. 17.

CHARACTERISTICS:

Carapace (fig. 24) not dorsally ridged posteriorly; branchiostegal

spine supported by a carina. Rostrum serrated dorsally; with only one ventral tooth. Eyes dark brownish in color and not extending 18 anteriorly to end of lower ventral tooth. Abdomen ridged on segments

II - VI and ending with a median posterior spine on segments III - VI.

Telson (fig. 25) dorsally sulcate and armed with at least 7 to 15 small dorso- lateral spines; with five terminal spines; the lateral pair nearly twice as long as the median three. Antennal scale (fig. 27) ends with a spine.

Mandible (fig. 26) with a strongly serrated molar process; the palp of the first maxilla bears one sharp spine; the proximal segments of the second maxilla are half the length of the two distal segments; second maxilliped with a podobranch.

DISTRIBUTION:

This species has been taken in all three oceans from the East

Coast of Africa to Lower California and Peru. It seems to be more rare in the Atlantic Ocean where it has been taken off British Guinea, the Bahamas and Cuba. The capture of this species off Oregon extended its northern limits. This species off Oregon has been taken in depths of 1000m.

Genus Meningodora Smith, 1882

Lateral surface of carapace with one straight carina from orbit to posterior margin of carapace. Without denticles on the posterior part of the dorsal carina of the carapace. First abdominal segment 19

bears no dorsal carina.

Meningodora mollis Smith, 1882

Meningodora mollis Smith, 1882, p. 74, pl. 11, figs. 8, 9, pl. 12,

figs. 5 -9.

Hymenodora mollis Bate, 1888, p. 841, pl. 136, fig. 5.

Notostomus fragilis Faxon, 1893, p. 207; 1895, p. 170, pl. 44, figs. 2-2b.

Notostomus mollis Balss, 1925, p. 266, fig. 37 (mandible).

Notostomus mollis Chace, 1936, p. 26; 1940, p. 164, fig. 38.

Notostomus mollis Chace, 1940, p. 164, fig. 38.

Meningodora mollis Sivertsen and Holthuis, 1957, p. 12.

CHARACTERISTICS:

Carapace (fig. 28) lacking denticles on the dorsal posterior part; only one lateral longitudinal carina; branchiostegal spine supported by a short ridge. Rostrum dentate only on the dorsal margin and

subequal to anterior margin of eyes. Eyes (fig. 29) black, narrower than eye stalk as seen dorsally; eye stalk bears on its mesial border a small whitish tubercle. Abdominal segments III through VI carinate.

The proximal inner segment of the second maxilla half the length of the two inner distal segments. Endopodite of the first maxilliped

(fig. 30) with two long equal segments. Second maxilliped with a 20 podobranch.

Dactylus of pereiopods I and II ending in small unequal spines.

Antennal scale (fig. 31) with a small subterminal lateral tooth; a mid - dorsal carina; 2. 3 times long as wide.

REMARKS:

The genus Meningodora was erected by Smith (1882) and has since acquired several synonyms. Meningodora was separated from

Notostomus and Hymenodora by Sivertsen and Holthuis (1957) and the original characteristics given it by Smith (1882) were reassigned to this genus.

Only one individual was taken. The margins of the second pleuron

(fig. 28) are asymetrical, the posterior margin protrudes more than the anterior margin. Dorsal -posterior lobes of pleurons IV and V with a minute round bulge.

DISTRIBUTION:

This species has been taken from the Atlantic Ocean (East Coast of United States, Bermuda, Bahamas), off the coast of Brazil, from the Indian Ocean, near the Philippine Islands and off the West Coast of Panama. The Oregon specimen extends the northern range in the Pacific Ocean from Panama. Depth ranges from 900- 3000m. This species off Oregon has been taken in depths of 1000m. 21

Genus Notostomus A. Milne Edwards, 1881

Lateral surface of carapace with more than one straight carina

from orbit to posterior margin of carapace. Denticles present

throughout length of dorsal carina of carapace. All abdominal seg-

ments bear a dorsal carina.

Notostomus japonicus Bate, 1888

Notostomus japonicus Bate, 1888, p. 826, 830, pl. 135, fig. 1.

CHARACTERISTICS:

Carapace (fig. 33) serrated dorsally throughout its length;

upper and lower lateral rostral carinae diverge posteriorly; gastro-

orbital and sub - hepatic carinae connected by post- hepatic carina and nearly parallel; infra - gastro- orbital lying just ventrally to the gastro- orbital; a short, oblique carina extends upward and backward from the middle of the gastro - orbital carina; sub - hepatic carina continuous with branchiostegal spine; sub -marginal carina present ventrally to sub -hepatic. Rostrum greater than half the length of the carapace, serrated dorsally and ventrally. Eyes large and black. Antennal scale (fig. 34) ending in a long lateral spine.

Abdominal carina on all somites dorsally; segments III - VI ending in dorso -posterior spines. Telson (fig. 35) with four or five 22 dorso- lateral spines; with five terminal spines; lateral pair the largest.

Endopodite of first maxilliped (fig. 36) three segmented; second maxilliped with a podobranch. Dactylus of first and second pereiopods ending in two teeth, between which the terminal tooth of the propodus fits.

REMARKS:

This is the first record of N. japonicus taken since its discovery in 1888 (Bate). The original one was taken off the southern part of

Japan at depths of 1030m. Off the Oregon coast this species has been taken to depths of 1000m. My specimen agrees with Bate's

(1888) figure of N. japonicus except that mine has an upper lateral - rostral carina. Dr. Fenner Chace, Jr. of the United States National

Museum has informed me that Bate's specimen had the upper lateral - rostral carina, but that he had failed to put it in his drawing.

DISTRIBUTION:

This species has been taken off the coast of Japan and off the

Oregon coast.

Hymenodora Sars, 1877

No ridge associated with the median lateral line from the orbital 23 region to the hind margin. Abdomen dorsally not carinated on any somite. Telson truncate and laterally armed with spines. Endo - podite of first maxillipeds composed of only two segments. Eyes small, light -brown in color and with a small median tubercle. Incisor process of mandible toothed for the entire length of its cutting edge.

KEY TO THE OREGON SPECIES OF HYMENODORA:

la. Rostrum short, reaching to level of eyes 2

b. Rostrum long, entending beyond level of eyes to distal

joint of antennular peduncle H. frontalis

2a. Podobranch present on second maxilliped; one anterior groove connects the supra and sub -hepatic carinae.

H. gracilis

b. No podobranch present on second maxilliped; a posterior

and an anterior groove connect the supra and sub - hepatic

carinae H. glacialis

Hymenodora frontalis Rathbun, 1902

H. frontalis Rathbun, 1902, p. 904; 1904, p. 28, fig. 8.

CHARACTERISTICS:

Integument thin, rather firm, rugose or covered with fine 24 wrinkles. Dorsal median carina extending almost to the middle of the carapace, and anteriorly into a rostrum extending as far as the distal joint of the antennular penduncle. Dorsal surface of rostrum

(fig. 38) armed with 3 to 6 small spines; ventral surface unarmed.

Orbit deep; a minute spine on the sub - orbital lobe. Branchiostegal spine strong; from it a sharp carina runs back subparallel to the in- ferior margin terminating near the posterior edge. The branchial region is bluntly ridged but bent down at its anterior end and up at its posterior end; a deep groove extends obliquely downward and backward from the orbital sinus nearly to the lower carina.

Eyes reach from one -half to one -third the length of the rostrum; light brown in color; with a slender, blunt spine close to the cornea on the inner margin. Antennal scale lanceolate; tipped with a small spine.

The second maxilliped bears a podobranch; third maxilliped with a well developed exopodite.

Telson (fig. 42) truncate and terminating in two pairs of spines, the outer pair nearly twice as long as the inner pair.

DISTRIBUTION:

This species has been taken by Rathbun (1902, 1904) from the

Bering Sea to off Monterey Bay, California, and Kamchatka; from the Far Eastern Seas of the U. S.S. R. in depths from 166 to 851m. 25

Off the Oregon coast this species has been taken to 1000m in depth.

Hymenodora glacialis (Buchholz), 1874

Pasiphae glacialis Buchholz, 1874, p. 279, pl. 1, fig. 2.

Hymenodora glacialis Smith, 1885, p. 501.

Hymenodora glacialis Smith, 1886, p. 678, pl. 15, figs. 3, 10. Hymenodora glacialis Stephenson, 1923, p. 59.

Hymenodora glacialis Chace, 1947, p. 31. Hymenodora glacialis Sivertsen and Holthuis, 1956, p. 15, fig. 11.

CHARACTERISTICS:

Carapace (fig. 43) extremely soft; anterior and posterior hepatic carinae joined at their lower margin by the obliquely directly sub - hepatic carina and at their upper margins by a bent groove convexed portion of which is directly anteriorly. A small branchiostegal spine and a small suborbital spine tip the suborbital lobe. Rostrum not extending beyond level of eyes, relatively high and convex on its lower margin, upper margin provided with 2 to 4 spines. Eyes brown in pigment; with a whitish papilla. Anterior margin of the second segment of the antennal peduncle forms a round lobe over the antennal scale. Abdominal segments smooth. Telson (fig. 44) dorsally sulcate; provided with five pairs of dorso- lateral spines. Telson ending in three pairs of spines, the lateral pair the shortest. 26

Mandible (fig. 45) toothed along its cutting edge and provided with a strong molar process. A rounded lobe on the posterior portion of the outer ramus of the first maxilla (fig. 46). The three inner lobes of the second maxilla equal in length. The endopod of the first maxilliped (fig. 47) composed of two segments, a very short proximal segment and a long distal one. The second maxilliped

(fig. 48) bears an epipod.

REMARKS:

Distinguishing between Hymenodora gracilis and H. glacialis has been most troublesome to carsinologists until Sivertsen and Holthuis

(1956) made clear the differences between them.

As pointed out by Smith (1886) the important distinction is the presence of a podobranch on the second maxilliped (fig. 53) of H. gracilis and its absence on H. glacialis (fig. 48). This was true for specimens of Sivertsen and Holthuis (1956) and also for the Oregon specimens. The rostrum in all of the Oregon specimens of H. glacialis (fig. 43) is higher and slightly shorter than in H. gracilis and its ventral margin is strongly convex whereas in H. gracilis

(fig. 50) the ventral rostral margin is not convexed but straight.

Another important distinction is the spination on the tip of the telson; in H. glacialis (fig. 44) there are three pairs of spines with the lateral pair the longest. Smith pointed out this difference between 27

H. glacialis and H. gracilis (1886). There are five pairs of dorso- lateral spines in my specimens of H. glacialis and only four pairs of spines in H. gracilis. Chace (1940) mentioned that a few of his specimens of H. gracilis had five or six terminal spines on the telson instead of the usual four. Only a few specimens of both species collected off the Oregon coast had undamaged telsons.

The integument of H. glacialis is thinner and more pliable than is that of H. gracilis.

DISTRIBUTION:

This species as reported by Sivertsen and Holthuis (1956) appears to be nearly restricted to temperate and arctic waters. However, it has been taken from the Gulf of Panama. Chace (1940) did not find this species off Bermuda, but it was taken from off the Bahamas nearly ten degrees further south. Since H. glacialis and H. gracilis have been considered in the past to be synonyms, one cannot trust the distributional records. Off the Oregon coast this species has been taken to 1000m in depth.

Hymenodora gracilis Smith, 1886

Hymenodora gracilis Smith, 1886, p. 681, pl. 12, fig. 6.

Hymenodora gracilis Murray and Hjort, 1912, p. 668.

Hymenodora gracilis Chace, 1940, p. 175, fig. 46. 28

Hymenodora gracilis Chace, 1947, p. 32. Hymenodora gracilis Sivertsen and Holthuis, 1956, p. 16, fig. 11, 13.

CHARACTERISTICS:

Carapace (fig. 50) thin and pliable; the anterior and posterior hepatic carinae are connected at their lower margins only by the carina extending obliquely from the orbits of the eyes. Small antennal and branchiostegal spines. Rostrum not extending beyond tip of eyes; its dorsal margin may have up to four spines or more. Eyes light brown and with a mesial papillae. Anterior margin of the second segment of the antennal peduncle forms a blunt tooth over the antennal scale. Abdomen lacking dorsal carina; the sixth segment 1. 9 times as long as the fifth.

Mandible (fig. 51) toothed its entire cutting surface and with a strong molar process. Endopodite of the first maxilliped (fig. 52) two- segmented, a short basal segment and a long distal segment.

The three inner lobes of the second maxilla equal in length. Second maxilliped (fig. 53) with a podobranch.

End of telson (fig. 54) with two pairs of spines; lateral pair the largest; telson armed with four pairs of dorso- lateral spines.

DISTRIBUTION:

This species is cosmopolitan and is found in all oceans. It has 29 been taken from 500 to 5300 meters in depth and off Oregon during night tows it has been taken near the surface to 1000m.

REMARKS:

A field observation for separating Hymenodora glacilis from

H. glacialis is that in almost all cases the carapace of the formers remains undistorted as result of contact with the net, whereas the carapace of the latter is extremely distorted and in many cases, nearly unrecognizable. The sixth abdominal segment is 2. 0 times as long as the fifth in H. gracilis; in H. glacialis the sixth is 1. 4 times as long as the fifth.

Genus Systellaspis Bate, 1888

Carapace with no straight lateral ridge from anterior margin to posterior margin. Sixth abdominal somite never carinate. Telson laterally armed with spines and terminating in sharply pointed end - piece. Eyes black and large. Antero -lateral margin of first abdominal somite with a tooth. Mandible toothed for its entire cutting edge.

KEY TO THE SPECIES OF SYSTELLASPIS:

la. Rostrum sub -triangular, less than half the length of the carapace; abdomen lacking dorsal carina on any somite.

S. braueri 30

b. Rostrum long and narrow, greater than half the length

of the carapace; abdomen dorsally carinated on fourth

and fifth segments S. cristata

Systellaspis braueri (Bals s), 1914

Acanthephyra braueri Balss, 1914, p. 595. Systellaspis densispina Stephensen, 1923, p. 57, fig. 17. Systellaspis braueri Balss, 1925, p. 245, figs. 17 -20, pl. 21. Systellaspis echinurus Coutière, 1911, p. 158.

Systellaspis braueri Chace, 1940, p. 180, fig. 50.

Systellaspis braueri Chace, 1947, p. 35. Systellaspis braueri Sivertsen and Holthuis, 1956, p. 19.

CHARACTERISTICS:

Carapace (fig. 56) compressed, not dorsally carinate; no lateral ridges running the entire length of the lateral surface of the carapace. Rostrum straight and triangular; about two - fifths as long

as carapace; dorsal edge with 8 -11 teeth, of which 3 -4 are on the

carapace; 1 -3 teeth ventrally. Antennal spine prominent; branchiostegal spine strong, directed outward and forward and situated on a short carina. Eyes large, black and with a large tubercle on the inner side.

Antennal scale (fig. 57) concave dorsally; external margin

I 31 straight; terminal spine joined by the truncate laminar blade.

Third segment of abdomen only with a carina which extends posteriorly as a stout spine. Antero -lateral margin of first abdominal somite toothed; third, fourth and fifth abdominal somites each with a dorsal posterior directed tooth; tooth on the third abdominal segment the largest; a tooth at the lower posterior margin of segment five; postero -lateral margin of somite six with a tooth; sixth abdominal segment twice as long as the fifth. Telson (fig. 58) laterally armed with spines and terminating in a sharply pointed endpiece.

DISTRIBUTION:

This species is cosmopolitan in distribution. Taken from the

Gulf of Guinea, Bay of Bengal, off Los Angeles, California, Azores and Newfoundland. The Oregon specimens extend its northern range.

Chace (1940) off Bermuda found this species in 2100 to 3000m. The smaller specimens were taken in the shallower water and the largest specimens in the deeper. Off Newfoundland and Azores Sivertsen and Holthuis (1956) took specimens in depths of 150 to 1500m. Off Oregon they were taken to depths of 1000m. This species appears frequently in the collections. 32 Systellaspis cristata (Faxon), 1895

Acanthephyra cristata Faxon, 1895, p. 162, pl. 44, figs. la, b. Systellaspis cristata Balss, 1925, p. 244, figs. 14, 15.

CHARACTERISTICS:

Carapace (fig. 62) firm and provided with one interrupted lateral gastro- orbital carina and a continuous submarginal carina. Branchio- stegal spine supported by a short carina; strong antennal spine. Carapace ridged throughout its length and elevated to its greatest height just posterior to the level of the eye orbit. Rostrum longer than half the length of the carapace, armed above and below with spines. Eyes large and black. First abdominal segment with a blunt tooth on its anterior mar-

gin; only abdominal segments three and four ridged dorsally; a small sharp pointed tooth is located on each side of the posterior margins of abdominal segments four and five. Telson (damaged terminally) dorsally sulcate and provided with at least 25 spines.

Mandible (fig. 63) toothed throughout its cutting edge and pro-

vided with a strong ridged molar process. The proximal segment of

the inner ramus of the second maxilla less than half the length of

the two distal segments. Second maxilliped (fig. 64) provided with

a podobranch. The distal portion of the appendix masculina (fig. 65)

of the first pleopod bears small teeth. 33

DISTRIBUTION:

This species occurs in three oceans. It was first taken in the

Gulf of Panama, Pacific Ocean (Faxon, 1895), and it was subsequently captured in the Indian Ocean (Anderson, 1896; Alcock, 1899, 1901; deMan, 1920, Balss, 1925), the Atlantic Ocean (Balss, 1925;

Holthuis, 1951; Springer and Bullis, 1956; Figueira, 1957; Fisher and Go.ldie, 1961). The Oregon specimen is a northern extension of its range and was taken from a depth of 1000m off Oregon.

REMARKS:

Only one specimen had been taken and is in relatively poor condition. Photophores were observed at the base of each pleopod and of the fifth walking leg. They were also distributed over the lateral surface of the carapace. 34

GENERAL DISCUSSION AND SUMMARY

TAXONOMY:

The name of the family to which the largest number of bathy-

pelagic carideans belong has been the cause of considerable confusion;

at various times it has been known as the Oplophoridae, Ephyridae,

Miersiidae and Acanthephyridae. For a detailed account of the evo-

lution of the family name as it is known today, see Chace, 1940, p. 199. He also discussed the proper spelling of the type genus. Systellaspis and Oplophorus are well defined genera and there

is little likelihood of confusing the species of these genera with any others. Systellaspis is easily recognized by the tooth on the anterior margin of the first abdominal segment. The tooth overlaps the posterior margin of the carapace. The telson is laterally armed with

spines and ends in a tapering sharp point. Oplophorus, a genus not discussed herein, is easily recognized by the spines on the outer margin of the antennal scale; the exopods on the third maxilliped and first pair of pereiopods are foliaceous and somewhat ridged. The telson ends in a sharp point. Likewise Acanthephyra causes little trouble.

It is set apart from the other genera by the shape of the incisor part of the mandible and the incomplete or straight ridge running the entire length of the lateral surface of the carapace from orbit to hind margin along the median lateral line. Meningodora is weakly 35 separated from Notostomus in that the former has only one lateral ridge on the carapace; the dorsal ridge of the carapace is lacking teeth on the posterior three- fourths and the abdomen is not dorsally carinate on the first somite. Notostomus has more than one lateral ridge on the carapace, the dorsal margin of the carapace is dentic- ulated nearly throughout its entire length and all abdominal somites are dorsally carinate. Hymenodora is separated most readily by the light brown eyes and a truncate, spinose tipped telson.

The family Pasiphaeidae is characterized by the pectinate chelae and the mandible lacking a molar process. The genera of pasiphaeids have been in the past separated by the presence or absence of a mandibular palp and whether this palp, when present, was one- or two - segmented. It was found that certain species such as Parapasiphaë in the adult condition have a two - segmented palp but in the immature forms have either no mandibular palp or in sub -adult forms may have a one - segmented palp. Also in the genus Sympasiphaea, not included herein, the adults are characterized by a one - segmented palp, but in the immature forms the palp is lacking and in some only a small bud is observed. The genera are in need of revision.

GEOGRAPHICAL DISTRIBUTION:

The portion of the sea through which nets have been drawn is extremely small and our knowledge of oceanic barriers to the 36

dispersal of bathypelagic organisms is correspondingly sparse. From such distributional data as are available at present, it is ob-

vious that the temperature and chemical composition of the sea have

a much greater effect on the dispersal of such species that have any land barriers. Probably then, oceanic currents will be found to de- limit faunistic zones in the ocean depths much as do mountain ranges,

deserts and bodies of water on land. In most cases, it is certainly

misleading to define the limits of pelagic organisms on the basis of

parallels of latitude and longitude.

It is found that of the species taken off the Oregon coast that only Pasiphaea pacifica, P. magna, Notostomus japonicus and Hymenodora frontalis had previously been taken in the Pacific Ocean.

The range of Para.pasiphaë cristata is now extended into the Pacific

Ocean. Four of the species P. magna, P. chacei, Meningodora mollis and Notostomus japonicus have ranges extended further north, either from Panama or lower California to off the Oregon coast. One

species N. japonicus taken only off the Japanese coast is now taken

off the Oregon coast.

COLOR OF BATHYPELAGIC CARIDEA:

Since most of the species had been preserved for some time before identification, precise color definition cannot be given. How- ever, those that were in formalin for not more than one week were of 37 some shade of red. The species of Acanthephyra, Parapasiphaë

sulcatifrons, Hymenodora gracilis, and H. glacialis have bodies entirely scarlet or scarlet-red. In another group of species, which are found relatively near the surface, the scarlet coloring is absent, and they are translucent. This is characteristic in Pasiphaea pacifies+, and P. chacei.

LUMINESCENCE:

Only Systellaspis cristata was observed to have photophores.

It appears that there are two types of light producing organs. The photophore or dermal light organs may possibly function as a species recognition. A second type of luminescence results in the production of a luminous cloud by a structure at the anterior portion of the gill chamber. The latter probably functions as a defensive organ, as was shown when some type of shrimp accidently hit the side of the bathy- sphere in which Beebe (1934, p. 304) was making observations. 38

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15. Faxon, W. 1893. Reports on the dredging operations off the west coast of Central America, etc. VI. Preliminary descriptions of new species of Crustacea. Bulletin of the Museum of Comparative Zoology, Harvard University 24:149 -220.

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38. . 1885. On some new or little known decapod Crustacea, from recent fish commission dredgings off the east coast of the United States. Proceedings of the United States National Museum 7:493 -511. 42

39. Smith, S. I. 1886. Report on the decapod Crustacea of the Albatross dredgings off the east coast of the United States during the summer and autumn of 1884. U. S. Commissioner of Fish and Fisheries. Annual Report, 1885, p. 605 -705.

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41 Stephensen, K. 1912. Report on the Malacostraca collected by the "Tjalfe "- Expedition, under the direction of Ad. S. Jensen, especially at W. Greenland. Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i Kjobenhavn. 64:57-134.

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44. Wood -Mason, J. and A. Alcock. 1891. Natural history notes from H. M. Indian Marine Survey Steamer "Investigator ", Commander R. F. Hoskyn, Commanding. No. 21. Note on the results of the last season's deep -sea dredging. Annales des Sciences Naturelles. Zoologie, series 6, 7:186 -202.

45. . 1892. Natural history notes from H. M. Indian Marine Survey Steamer "Investigator ", Commander R. H. Hoskyn, R. M. , Commanding. Series II, No. 1. On the results of deep -sea dredging during the season 1890 -91. Annales des Sciences Naturelles. Zoologie series 6, 9:358- 370.

46. Yaldwyn, John C. 1962. A new Pasiphaea (Crustacea, Decapoda, Natantia) from Southern California waters. Los Angeles. p. 15 -24. (Allan Hancock Foundation for Scientific Research. Contribution no. 245) Figure 1. Pasiphaea pacifica.

Figure 2. P. pacifica, left antennal scale.

Figure 3. P. pacifica, tip of telson.

Figure 4. P. pacifica, right mandible.

Figure 5. Pasiphaea magna.

Figure 6. P. magna, left antennal scale.

Figure 7. P. magna, tip of telson.

Figure 8. P. magna, right mandible.

Figure 9. P. magna, left first pleopod. 44 Figure 10. Pasiphaea chacei.

Figure 11 P. chacei, left antennal scale.

Figure 12. P. chacei, tip of telson.

Figure 13. P. chacei, right mandible.

Figure 14. P. chacei, left first pleopod

Figure 15. Parapasiphaë sulcatifrons.

Figure 16. P. sulcatifrons, dorsal view of left eye. Figure 17. P. sulcatifrons, left antennal scale.

Figure 18. P. sulcatifrons, tip of telson.

Figure 19. P. sulcatifrons, right mandible. Figure 20. Parapasiphaë cristata.

Figure 21. P. cristata, tip of telson. Figure 22. P. cristata, right mandible.

Figure 23. P. cristata, left antennal scale. 46

1/ .

// IB

mm 17 M 20 prp

23 10

pl, Il1 3 12 Figure 24. Acanthephyra curtirostris.

Figure 25. A. curtirostris, tip of telson.

Figure 26. A. curtirostris, anterior view of left mandible.

Figure 27. A. curtirostris, left antennal scale. Figure 28. Meningodora mollis.

Figure 29. M. mollis, dorsal view of right eye.

Figure 30. M. mollis, left first maxilliped.

Figure 31. M. mollis, left antennas scale.

Figure 32. M. mollis, anterior view of left mandible. 48 Figure 33. Notostomus japonicus.

Figure 34. N. japonicus, left antennal scale.

Figure 35. N. japonicus, tip of telson.

Figure 36. N. japonicus, left first maxilliped.

Figure 37. N. japonicus, anterior view of left mandible.

Figure 38. Hymenodora frontalis.

Figure 39. H. frontalis, right first maxilliped.

Figure 40. H. frontalis, anterior view of right mandible.

Figure 41. H. frontalis, left first pleopod.

Figure 42. H. frontalis, tip of telson. 50 Figure 43. Hymenodora glacialis.

Figure 44. H. glacialis, tip of telson.

Figure 45. H. glacialis, anterior view of right mandible.

Figure 46. H. glacialis, right first maxilla.

Figure 47. H. glacialis, left first maxilliped.

Figure 48. H. glacialis, right second maxilliped.

Figure 49. H. glacialis, left first pleopod.

Figure 50. Hymenodora gracilis.

Figure 51. H. gracilis, anterior view of left mandible.

Figure 52. H. gracilis, right first maxilliped.

Figure 53. H. gracilis, right second maxilliped.

Figure 54. H. gracilis, tip of telson.

Figure 55. H. gracilis, left first pleopod. 52 Figure 56. Systellaspis braueri.

Figure 57. S. braueri, left antennal scale.

Figure 58. S. braueri, tip of telson.

Figure 59. S. braueri, left second maxilliped.

Figure 60. S. braueri, anterior view of left mandible.

Figure 61. S. braueri, left first pleopod.

Figure 62. Systellaspis cristata.

Figure 63. S. cristata, anterior view of right mandible.

Figure 64. S. cristata, right second maxilliped.

Figure 65. S. cristata, right first pleopod. 54