/. LAURENS BARNARD A Review of the Family

Synopiidae (== Tironidae),

f Mainly Distributed in J

the Deep Sea

(Crustacea: Amphipoda)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 124
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INTERNATIONAL WOK YEAR • 1972
SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 124

J. Laurens Barnard A Review of the Family

Synopiidae (=Tironidae),

Mainly Distributed in

the Deep Sea

(Crustacea: Amphipoda)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1972
ABSTRACT

Barnard, J. Laurens. A Review of the Family Synopiidae ( = Tironidae), Mainly

Distributed in the Deep Sea (Crustacea: Amphipoda). Smithsonian Contributions

to Zoology, number 124, 94 pages, 46 figures, 1972.—The family Synopiidae ( =

Tironidae) comprises 69 species in 14 genera, of which 12 new species and 3 new

genera are described herein. Keys and diagnoses to the taxa include all described

species. The new genera, lleraustroe, Latacunga, and Priscosyrrhoe, are elaborated

from species formerly placed in Austrosyrrhoe or from new species and were suggested

by similarity analysis of generic groups arranged in a dendrogram. Three main

evolutionary branches occur in the family, comprising groups typified by SynopiaTiron, Syrrhoe, and Syrrhoites-Bruzelia as based on gnathopodal structure. The

shallow-water Synopia-Tiron group has the most advanced gnathopods but retains

eyes in most members. The Syrrhoe group, containing shallow and deep-water species,

retains eyes in the shallow-water members; but the majority of synopiid species occurs

in the anoculate, deep-sea group typified by Syrrhoites and Bruzelia, having ordinary

gnathopods.

The type-genus, Synopia, is morphologically remote from other genera in the

family, formerly known as Tironidae, but in comparison to related families, it

remains close to other tironids and should not be segregated at familial level.

Synopia and Tiron and possibly Syrrhoe, the only shallow-water tropical genera,

are not directly antecedent to the cold-water Syrrhoites group but may be the

oldest synopiid genera despite the highly evolved gnathopods in Tiron and pelagont

coxae in Synopia and Syrrhoe. Tiron is a benthic genus but Synopia and Syrrhoe

are nektodemersal. Most synopiids are adapted to epibenthic life though many feed

on the benthos. These adaptations make them very successful in the deep sea but not

in the hydrologic turmoil of the shallow sea, where amphipods are usually wholly

nektonic or wholly fossorial except during male swarming.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

Library of Congress Cataloging in Publication Data

Barnard, Jerry Laurens, 1928—

A review of the family Synopiidae (=»Tironidae)

(Smithsonian contributions to zoology, no. 124)

Bibliography: p.

1. Synopiidae. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to

zoology, no. 124.

QL1.S54 no. 124 [QL444.A5] 591'.08s [595'.371] 72-1787

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402 - Price $1.25 (paper cover)
Contents

Page

Introduction 1

Discussion 1

Relationships of the Synopiidae 3

Evaluation of Generic Similarities 6

Relationships According to Least Differences (Six-Character Analysis) 9

Evolution from Oculate Ancestors 10

Enlarged Molar Theory 12

The Direction of Evolution in Characters 13

Synopiidae 13

Key to the Genera of Synopiidae 15

Austrosyrrhoe K. H. Barnard, revised 16

A. crassipes K. H. Barnard 17

A. rinconis J. L. Barnard 17

A. septentrionalis Stephensen 17

A. fimbriatus (Stebbing and Robertson) 17

Bruzelia Boeck 17

Key to the Species of Bruzelia 18

B. ascua J. L. Barnard 18

B. australis Stebbing 18

B. diodon K. H. Barnard 19

B. guayacura, new species 19

B. inlex J. L. Barnard 21

B. pericu, new species 21

B. (?) popolocan, new species 24

B. poton, new species 27

B. tuberculata Sars 29

B. typica Boeck 29

Bruzeliopsis Chevreux 32

Key to the Species of Bruzeliopsis 32

B. alberti Chevreux 32

B. cuspidata (J. L. Barnard) 32

B. turba J. L. Barnard 32

Garosyrrhoe J. L. Barnard 33

Key to the Species of Garosyrrhoe 34

G. bigarra (J. L. Barnard) 34

G. disjuncta J. L. Barnard 34

Ileraustroe, new genus 34

/. ilergetes (J. L. Barnard) 35

/. torpens (J. L. Barnard) 35

Jeddo J. L. Barnard 35

/. simplisyrrhis J. L. Barnard 35

Latacunga, new genus 35

L. latacunga, new species 41

L. comanita, new species 41

Priscosyrrhoe, new genus 44

P. priscis (J. L. Barnard) 44

Pseudotiron Chevreux 44

Key to the Species of Pseudotiron 45

P. bouvieri Chevreux 45

P. coos J. L. Barnard 45

P. golens J. L. Barnard 45

in
SMITHSONIAN CONTRIBUTION TO ZOOLOGY

P. longicaudatus Pirlot **•

P. pervicax J. L. Barnard * 6

Stephobruzelia J. L. Barnard 50

S. dentata (Stephensen) 50

Synopia Dana 50

Key to Seven Species of Synopia 51

Key to Three Species of Synopia 51

S. angustifrons Dana 51

S. caraibica Bovallius 51

S. gracilis Dana 51

5. orientalis Kossmann 51

S. scheeleana Bovallius 51

S. ultramarina Dana - 51

S. variabilis Spandl 52

Syrrhoe Goes 52

Key to the Species of Syrrhoe 52

S. affinis Chevreux 53

S. crenulata Goes 53

S. longifrons Shoemaker 53

S. nodulosa K. H. Barnard 54

S. oluta, new species 54

S. papyracea Stebbing 55

S. psychrophila Monod 56

S. semiserrata Stebbing 60

5. serrima, new species 60

S. tuberculata Dahl 64

Syrrhoites Sars 64

Key to the Species of Syrrhoites 64

S. anaticauda K. H. Barnard 65

S. cohasseta J. L. Barnard 65

S. columbiae, new species 65

S. cu, new species 67

S. dulcis J. L. Barnard 72

S. lorida (J. L. Barnard) 72

S. pacifica Nagata 73

S. pantasma, new species 73

S. pusilla Enequist 76

5. redox J. L. Barnard 76

S. serrata (Sars) 76

S. silex J. L. Barnard 76

S. sorpresa (J. L. Barnard) 76

S. tenella K. H. Barnard 79

S. terceris J. L. Barnard 79

S. trux J. L. Barnard 79

S. walkeri Bonnier 79

S. species A 80

Tiron Liljeborg 33

Key to the Species of Tiron 83

T. altifrons Reid 84

T. antarcticus K. H. Barnard 84

T. australis Stebbing 34

T. biocellata J. L. Barnard 84

T. brevidactylus (Pillai) 35

T. intermedia Reid 35

T. spiniferum (Stimpson) 8g

T. thompsoni Walker 8g

T. tropakis, new species ng

Literature Cited 8 g

Appendix g 2

IndeX

 IIZIIZIIIZZ 93
J. Laurens Barnard A Review of the Family

Synopiidae (=Tironidae),

Mainly Distributed in

the Deep Sea

(Crustacea: Amphipoda)

Introduction

The Family Synopiidae (=Tironidae) is reviewed

and reorganized and a number of new species and

three new genera are described. The materials examined in this study come mainly from deep-sea

collections made by the research vessels Vema of

Lamont Geological Observatory and Eltanin of the

National Science Foundation. Because individuals of

this family are friable and often fragmented, only

those specimens in reasonably good condition have

been studied. Numerous undescribed species remain

to be collected in good condition as evidenced from

a cursory examination of fragments in various

samples. Hopefully this review of the family will

stimulate further collecting and special care in washing mud from bottom samples so as to provide good

materials of unknown species.

This work was supported by NSF GB 3285,

providing the skilled assistance of Mrs. Elaine R.

Hodges, who made the original drawings, later inked

by the writer with materials supplied by the Richard

Rathbun Fund. Dr. Andrew G. Carey, Jr., of Oregon

State University, kindly provided specimens from

his deep-sea transect (see J. L. Barnard, 1971, for

main study). Dr. John S. Garth and Dr. Olga Hartman of the Allan Hancock Foundation kindly provided specimens of Tiron for study.

/. Laurens Barnard, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Discussion

The Synopiidae are, perhaps, the most diverse of

those families primarily confined to deep seas. Only

a few genera, such as Tiron Liljeborg, Synopia Dana,

Syrrhoe Goes, and Garosyrrhoe J. L. Barnard, have

species confined to warm shallow seas; the other

genera, along with many species of Syrrhoe, are

primarily confined to deep cold-water.

KEY TO LEGENDS

Figures are labeled with capital and lower-case

letters, the capitals usually indicating the name of a

structure and the lower case letters indicating a modification such as orientation or specimen considered.

A, antenna; B, upper lip; C, coxa; D, dactyl of a

pereopod; E, epimeron; F, accessory flagellum; G,

gnathopod; H, head; /, inner plate; /, various meanings according to specific figure legend, but if unmentioned meaning epistome; L, palp; M, mandible,

O, outer plate; P, pereopod; Q, mandibular molar; R,

various meanings according to specific legend but if

unmentioned meaning lower lip; S, maxilliped; T,

telson; U, uropod; V, urosome; W, pleon; X, maxilla;

Y, prebuccal mouthparts; Z, mandibular incisor, a,

anterior; d, dorsal; e, various meanings according to

specific legend; i, inner; /, left or lateral; n, meaning

according to specific legend; o, outer or in case of

thoracic appendages meaning opposite view of a rightsided appendage; q, various meanings; r, right; s,

setae removed or missing; t, spine; u, half; v, ventral;

iv, various meanings according to specific legend; y.

various meanings; z, broken appendage; 8, rostrum.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The family therefore has not been of much

concern until recent explorations of the deep sea

have revealed a very diverse assemblage of synopiids.

The number of species now totals 69. Extraordinary

variations in morphology occur among these species.

Many of them are strikingly ornamented with dorsal

teeth and lateral carinae. Unfortunately, many of

the deep-sea collections made in the past have resulted in scarce individuals, badly fragmented or

damaged, and collected in widely scattered basins.

Setae are often removed by sandblasting or siltblasting

during the washing of mud samples. These defects

are slowly being lessened but taxonomists are still

beset with a host of species inadequately described.

Despite imperfect descriptions, one may recognize the

unsatisfactory state of taxonomy in the deep-sea

genera. There are two main problems: (1) the recognition of individual groups of species intragenerically and intergenerically and (2) the difficulty of

determining the extent to which intraspecific variation is a measure of geographic subspeciation. One

must presume that numerous species are yet undescribed but some progress in perfecting the classification can nevertheless be made now.

In the early days of gammaridean taxonomy, even

as late as Stebbing's (1906) monograph, only 2

valid species in Bruzelia and 3 in Syrrhoites were

known and all were from the arctic and boreal

provinces of the northeastern Atlantic Ocean. Both

genera have an enormous, smooth molar dominating

the mandible; the entire mandibular body is also

extraordinarily enlarged and therefore the palp appears relatively weak. The two genera were distinguished by the presence of a cleft in the telson of

Syrrhoites and its absence in Bruzelia. One of the

species of Syrrhoites, S. fimbriatus Stebbing and

Robertson, has since been shifted by J. L. Barnard

(1964) to Austrosyrrhoe but that may be an erroneous move, as the mandibular molar is still poorly

known.

In 1911, Chevreux described Bruzeliopsis from the

abyss of the Bay of Biscay but did not specifically

mention any characters differentiating it from its

obvious relative, Bruzelia, as indicated by the absence

of a cleft in the telson. In retrospect one may infer

that Chevreux considered the absence of serrate

palmar spines on the gnathopods and the strong

disproportion in coxae 3 and 4 to be generic characters.

Unlike the 5 early species of Bruzelia and Syrrhoites,

Bruzeliopsis alberti has a very broad coxa 3 with

a distinct posterodorsal excavation and very small

"comma-shaped" and blunt coxa 4 as in Syrrhoe.

Article 1 of antenna 1 is elongate and distally cuspiform and articles 2 and 3 relatively short in Bruzeliopsis, whereas in the early species of the other

genera this is not true.

K. H. Barnard (1925) described Austrosyrrhoe

from the bathyal of South Africa. The generic

diagnosis distinguished the genus from Syrrhoe but,

without other considerations, essentially gave a picture

of a species of Syrrhoites having a very robust

gnathopod 1. In his further description of the typespecies, however, K. H. Barnard noted that the

mouthparts resembled Syrrhoe and this made possible

many years later a better generic diagnosis in which

the condition of the mandibular molar was specifically included (J. L. Barnard, 1964). Stephensen

(1931) had confirmed the validity of Austrosyrrhoe

by describing a second species with robust gnathopod

1 and mouthparts like Syrrhoe but was understandably hesitant in his classification of it in view of the

paucity of illustrations accompanying the description

of the type-species. Indeed, Gurjanova (1951) shifted

Stephensen's A. (?) septentrionalis to Syrrhoites but

it was returned to its proper place by J. L. Barnard

(1964). By 1956 only 14 species had been described

in the 4 genera.

In 1962, J. L. Barnard described a new species of

Syrrhoites from the abyss off South Africa and suggested that it might represent K. H. Barnard's

Austrosyrrhoe crassipes. The type-specimen of the

latter had been found to be essentially destroyed

through deterioration. J. L. Barnard discussed elongation of the telson in S. torpens and the robust gnathopod 1 as possible generic characters, but even though

he figured the syrrhoe-like mandible, he failed to note

its significance. This error was rectified by J. L.

Barnard (1964) and Syrrhoites torpens, S. fimbriatus,

A. crassipes, A. septentrionalis and a further new

species, A. ilergetes were united as members of

Austrosyrrhoe.

The genus Kindia was erected by J. L. Barnard

(1962a) in the mistaken belief that two species

assigned to it were distinguishable from Syrrhoites in

their simple gnathopods and slightly enlarged fourth

coxae but Bonnier (1896) had already described

Syrrhoites walkeri with these conditions and S.

pusillus Enequist (1950) was known to have nearly
NUMBER 12 4

simple gnathopods. J. L. Barnard (1964) dismissed

these characters as having no significance and synonymized Kindia with Syrrhoites.

The host of new species described in these 4

genera now sheds light on further difficulties. If one

examines characters such as serration or simplicity

of palmar spines on gnathopods, minute differences

in shapes of coxae 3-4, spination of uropods, telsonic

clefts, and elongation and narrowing of telson, one

may detect pairs of species now assigned to Bruzelia

and Syrrhoites which have strong relationships and

other species now assigned to Austrosyrrhoe, which

if given treatment equivalent to that afforded

Syrrhoites and Bruzelia, would be relegated to newly

erected genera. There is now considerable doubt that

J. L. Barnard (1962a and 1964) had anything at all

resembling Austrosyrrhoe crassipes when he described

Syrrhoites torpens and that species presumably must

stand. In pursuing this most crucial matter first I

have discarded any preconception that A. torpens

and A. crassipes have characters in common and relied entirely on K. H. Barnard's statements that

various undescribed characters of A. crassipes are

"like Syrrhoe," or are "as in Syrrhoe crenulata" or

"[resemble] . . . Bruzelia typica."

The families Synopiidae and Tironidae were synonymized as late as 1964 by J. L. Barnard. The

Synopiidae contained only the genus Synopia. That

genus bears the following similarities to various members of Tironidae: coalesced eyes, accessory flagelluni,

conjoint bases of certain flagella, accessory eyes,

galeate or ploughing head, narrow space between

outer lobes of lower lip, pelagont coxae, powerful

abdomen, reduced article 3 of mandibular palp, both

extremes of mandibular molar found in Tironidae,

and evenly extended pereopods 3-5. Despite these

similarities, Synopia stands apart from tironids in the

short telson, foliaceous maxillipeds, vestigial dactyl

of gnathopod 2, and absence of dorsal body ornamentation. The short telson and simple body segments

suggest that Synopia retains more of the primitive

character of a hypothetical early tironid than any

other living group of species. In the similarity analysis

presented below, Synopia stands well apart from

many tironids, though it is connected to the TironPseudotiron group by many similarities, such as the

simplified gnathopods. The latter character itself is

far advanced from the primitive gammaridean and

is one of many complications in constructing a parsimonious systematic scheme in which Synopia would

stand near the base of an elaboration. One would

not expect a phyletic scheme to contain a sequence

passing from subchelate to simple to subchelate

gnathopods. But the subchelate gnathopods of tironids

are in no measure comparable to those of primitive

gammarideans. The emergence of heavily subchelate

gnathopods in evolutionary lines such as corophiideans which passed through many ages of gnathopodal

simplicity also refutes the suggestion that once a

gnathopod has become simple the genetic complex

of the organism has totally lost the capacity to reconstruct a subchelate gnathopod in later evolutionary sequences.

Relationships of the Synopiidae

Synopiids and tironids differ little from the basic

gammaridean (J. L. Barnard, 1969a, compare Figures

1-2 with 31). The main difference is the galeate

head but even this is occasionally so heavily modified

in epibenthic forms as to be unrecognizable. There

are many unusual developments in the synopiids that

are not found in the basic gammaridean but many

of these may also be found in the Gammaridae, the

family with presumed closest relationships to the

hypothetical basic gammaridean. In most synopiids

the telson becomes elongate, thus is considerably

longer than the peduncle of uropod 3, but the typegenus Synopia is the exception and fits the basic

gammaridean in this character. About half of the

species in Synopiidae have the telson uncleft and

most of the genera have the mandibular palp either

grossly enlarged, strongly slenderized, or palp article

3 is very shortened, or, rarely, the palp is lost. About

half of the genera have the mandibular molar extraordinarily enlarged, tumid and fuzzy, in contrast to

the basic gammaridean. Like other gammaridean

families, there is also a progression from subchelate

to simple in the gnathopods, these appendages already

lacking the primitive conditions of sexual dimorphism

and enlargement of gnathopod 2. The coxae of

synopiids also exhibit a great variety of disproportionate changes from the basic condition. Uropod 3 often

has an elongate peduncle.

But none of these modifications is wholly unique

to Synopiidae, and therefore it is sometimes difficult

to identify an amphipod as a member of Synopiidae.

The galeate head, in combination with the presence
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of a macroscopic accessory flagellum, nonfleshy telson,

reasonably proportional articles 5-6 of pereopods 1—2,

well developed molar, elongate telson (with one

generic exception), unbroadened mandibular incisors,

reasonably short article 3 of gnathopod 2, and slender

peduncle of antenna 1 with articles 2 and 3 not

strongly shortened or telescoped into article 1, distinguish synopiids from similar members of Haustoriidae, Oedicerotidae, Phoxocephalidae, Argissidae,

Eusiridae, (?Vitjazianidae), Astyridae, Stilipedidae,

Acanthonotozomatidae, Stegocephalidae and

Lysianassidae.

Synopiidae and Oedicerotidae are very similar

morphologically. The latter family is presently divisible into two sections, the primitive section containing species with a basic head and paired eyes, the

advanced section containing species with a galeate

head like that of Synopiidae, and dorsally coalesced

eyes like oculate species of Synopiidae. But many

species of Oedicerotidae, like Synopiidae, live in the

deep sea, have lost the eyes, and certain species have

lost the galeate features of the head, the rostrum

being severely reduced in size. Primitive oedicerotids

occur mainly in shallow waters of the southern

hemisphere whereas the oculate advanced oedicerotids dominate shallow waters of the northern hemisphere. In either hemisphere the main distribution

occurs in cold-temperate waters.

Synopiids presumably differ from Oedicerotidae in

their gross niche category. The Oedicerotidae have

weakly to strongly fossorial pereopods and therefore

are mainly burrowers in soft bottom. Except in certain species of Tiron and Pseudotiron, synopiids have

elongate, nonfossorial pereopods, powerful pleopods

and various other adaptations to epibenthic behavior.

The wide tendency for synopiids to have modified

coxae 3—4 suggests that even those without modified

coxae are poorly fossorial and that the general behavior of synopiids can be called demersal. The modified coxae are hereafter termed "pelagont." The

function of this adaptation is unknown but it is

associated especially with those species believed to

be demersal as evidenced by the kinds of collecting

devices in which these organisms have been entrapped. The presence of silt grains in the midgut

of various specimens having pelagont coxae suggests

deposit feeding but this is believed to be errant (the

organisms move from place to place). One may therefore visualize an ordinary deep-sea synopiid skimming

above the surface of the bottom in short hops, then

descending to the bottom to feed selectively on

favorable bottom deposits. Since most synopiids

probably do not find shelter in deep burrows they

must therefore have powerful escape mechanisms

from predators. Certain species have heavily protuberant foreheads suggesting that they plow the mud

surface. The shallow-water genus Tiron appears to

be the strongest exception in the Synopiidae to the

ecological picture just painted but it is also very

near to the logical concept of a primitive synopiid

and therefore it may reflect, more than other genera,

the basic nestling behavior of the early gammaridean.

The epibenthic modification of certain species of

Tiron is, however, very striking; species still living,

like Pseudotiron coos, reflect the morphological

transformation so neatly as to confound the generic

differences between Tiron and Pseudotiron, the latter

genus containing fully epibenthic (pelagont) species.

Synopiidae also differ from Oedicerotidae in the

presence of a large accessory flagellum. Presumably

this appendage has some tactile and sensory property

no longer necessary to the relative sedentary

Oedicerotidae. Some oedicerotids, like Synchelidium,

are not always sedentary; males of some species of

Synchelidium are strongly neritic. All Oedicerotidae

maintain the primitively short telson which has advanced from the basic gammaridean state by having

the lobes coalesced and formed into a small oval

plate. In Synopiidae only Synopia has a short telson

and only in Synopia variabilis have the lobes coalesced

to form an ovate plate. Synopia also bears highly

pelagont coxae. If one suggests that Oedicerotidae

and Synopiidae have a common ancestor the primitive member would have a large accessory flagellum,

ordinary coxae, a short cleft telson, nonfossorial

pereopods, and paired eyes. The reduction of accessory flagellum and solidification of telson would then

mark one branch, the Oedicerotidae, and the retention of accessory flagellum and clefted telson would

mark the Synopiidae. Only one species of Synopia

then evolved a solidified telson, 2 species retained

the short and cleft telson, whereas all descendent

synopiids are characterized by an elongate telson

which becomes solidified and even more elongate in

several advanced genera. In Oedicerotidae some of

the primitive members with uncoalesced eyes still live,

but as far as is known, those species with uncoalesced

eyes in Synopiidae are extinct. The Oedicerotidae
NUMBER 1 2 4

developed fossorial appendages whereas the

Synopiidae occupied the neritic and demersal niche

categories. Generally the Synopiidae have retained

more primitive morphological characteristics than

the Oedicerotidae but both have certain parallel developments, such as coalesced eyes and the tendency

to a solid telson. Oedicerotids universally, except for

Metoediceros Schellenberg, have a very long styliform

uropod 3 with an elongate peduncle. The elongation

of the peduncle occurs in a few species of Synopiidae

and does not seem to have any generic value attributable to this development.

The Astyridae also appear to be a product of evolutionary divergence from ancestors similar to those

of oedicerotids and synopiids. Presumably astyrids,

with one genus and 5 species, occupy a niche category

similar to that of synopiids because one or two species

have the plow-shaped heads of certain synopiids, and

the appendages are not visibly fossorial;1 but astyrids

appear to have weaker pleopods than do synopiids

and the accessory flagellum is strongly reduced but

not as much as in oedicerotids. Uropod 3 has a short

peduncle, the telson is short and either cleft or uncleft, the head is ungaleate in primitive astyrids, eyes

are absent and the genus is entirely distributed in the

deep sea. The gnathopods have the appearance of

certain oedicerotid characters; they have article 5

broadly lobed posteriorly and wider than article 6.

Astyra differs qualitatively from synopiids in the

mandible, which bears a heavily and sharply serrate,

subflabellate incisor, a swept, conical molar bearing

setae and in the lower lip, which has a wide channel

between the outer lobes. The mandible has the

descriptive characteristics of that in Synchelidium,

an oedicerotid. The excessively large number of raking spines on the mandible suggests a feeding behavior distinct from the synopiids and more intensely

adapted for raking than in most oedicerotids. Astyra

definitely appears to be another branch of the early

divergence between oedicerotids and synopiids but as

far as is known the shallow-water lines have become

extinct.

The Vitjazianidae, with 2 genera and 3 species,

occur in bathyal and abyssal depths. Their mandibles

have ordinary triturative molars, the telson is short

1

 However the type-species of Astyra has posterior setal

spines on article 6 of pereopods 3-5 suggesting a weak

fossorial adaptation.

and cleft, the peduncle of uropod 3 is short, the

accessory flagellum is long, and the base of the flagellum on antenna 1 is conjoint as in the majority of

synopiids. The lower lip is normal, the raker spines

on the mandible are sparse and gnathopod 1 is always

simple, raptorial, and very similar to that seen in

Tiron and Pseudotiron of the Synopiidae. The one

known species of Vitjaziana has a very elongate

antenna 2 and pereopods 3—5, suggesting that it is

adapted, like Melphidippa, of the Melphidippidae,

to sit upside down on a soft ooze bottom in a cradle

formed of the elongate legs (Enequist, 1950). The

posterior margins of those legs have short setae similar to those mentioned for Astyra and this enforces

the suggestion that Vitjaziana comes in contact with

bottom sediment. The second genus of Vitjazianidae,

Vemana, with two species, has short pereopods like

certain species of Tiron; they are subfossorial.

Gnathopod 2 bears a palm. Neither genus of Vitjazianidae has a galeate head but the distinction from

Synopiidae is otherwise very unclear. The short telson

distinguishes vitjazianids from all but Synopia in the

Synopiidae. The coxae are extremely short in Vitjaziana, like those in the Melphidippidae, but

Vemana has coxae longer than broad while definitely

tending to become discontiguous.

Tracing the phylogeny of synopiids beyond the

previous remarks appears fruitless at this time. Such

activity is fraught with many difficulties. For example, the Pardaliscidae, Stilipedidae and Hyperiopsidae have many adaptations similar to those of

Synopiidae, and the observer could easily develop

schemes that included those families in the early

evolutionary divergence of synopiids and oedicerotids.

The complex of genera variously assigned to such

families as Eusiridae, Pontogeneiidae and Calliopiidae

has not been mentioned and they offer serious obstacles in clarifying relationships. They are characterized, at least in their primitive members, by greatly

reduced accessory flagella (or lost entirely in advanced members), paired eyes, generally short cleft

or uncleft telsons variously modified but often elongate in advanced members, and some of these eusiridpontogeneiid-calliopiid genera appear as good ancestral grades to the Oedicerotidae; some of them

have elongate peduncles on uropod 3, others have

fossorial pereopods, others have an elongate pereopod

5 like that seen in oedicerotids, so that various primitive oedicerotids appear scarcely distinct from certain
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

calliopiids. The reduction in accessory flagellum becomes a key to gammaridean advancement and the

evolutionary distance, in terms of linkages between

the primitive synopiid and the primitive oedicerotid,

may be vast. A clearer statement might be that if

synopiids and oedicerotids did have a direct ancestor,

the descendent with reduced accessory flagellum became the ancestor to a far greater diversity of living

genera than did the ancestor retaining the long

accessory flagellum. The final possibility, and that

with the highest probability in logic is that a variety

of early gammarideans, of the grade of structure now

seen in the Gammaridae, advanced in evolution by

loss or reduction of the accessory flagellum and that

the similarities now seen between advanced oedicerotids and various primitive synopiids are simply

convergences.

One very good example of the possible polyphyly

in the eusirid-pontogeneiid-calliopiid group of genera

is the obvious relationship of Gammarellus in the

Gammaridae with Calliopius in the Calliopiidae. The

two genera have a large number of similarities, yet

Gammarellus bears a long accessory flagellum and is

classified as a member of the Gammaridae while

Calliopius has a reduced or lost accessory flagellum

and forms the type-genus of the Calliopiidae. Neither

Calliopius nor Gammarellus could have been the ancestors to most genera of the pontogeneiids and

eusirids because the latter families retain the primitive cleft of the telson whereas Gammarellus and

Calliopius already bear the advanced and nearly solid

telson. Other ancestors to various pontogeneiids and

eusirids must be sought. Most of the genera other

than Calliopius assigned to the Calliopiidae also have

little relationship to Calliopius and, indeed, pairs of

species now found in different genera, and in apposite

families, should probably be joined together in a

common genus by disregarding the degree of telsonic

clefting. Classification of eusirids, pontogeneiids and

calliopiids is probably the greatest of those problems

facing the gammaridean taxonomist. Since the hypothetical ancestor of synopiids has the same characteristics that one imagines belong to the ancestors of

eusirids (etc.) one must face the high probability

that they did not have a common ancestor except in

the remotest context (they probably had the same

ancestor at some time if gammarideans themselves

are monophyletic).

Evaluation of Generic Similarities

FIGURE 1

Twenty-five characters both of possible generic value

and known for most of the species in the family were

analyzed in a matrix from which dendrograms were

evolved. Each character was assigned a value ( + or

0) for each species and these species were then

condensed into 23 groups, each group of species having 24 or 25 character congruencies. Each group,

provisionally called an incipient genus, was then

compared with every other group so as to produce a

matrix of values composed of the number of characters in common between each possible pair. The

weighted pair group method of dendrogram evolutions was then performed on the matrix to produce

Figure 1. The species groups are labeled with their

generic names only if the type-species of a named

genus is contained within a group, or with both

generic and specific names if no generic name has

been proposed for the group. To save space, the

characters are marked in Table 1.

The ultimate dendrograms are based on high

values, indicating both a success in selecting characters and a fairly tightly related family of incipient

genera. No arbitrary dendogrammatic value can be

selected to define a genus since the species were preclassified into groups and numerous characters have

not been included in the analysis. The analysis is

thus below the standards of true phenetics but ultimately has several valuable results.

Several surprising results came from this study.

Pelagont coxae and enlarged molars did not grossly

influence the digits, whereas the three main branches

of the system, Tiron, Syrrhoe and Syrrhoites are

mainly distinguishable by gnathopodal characters,

the Tiron group by simple gnathopods, the Syrrhoe

group by transverse palms and the Syrrhoites group

by oblique palms. To some extent an overemphasis

occurred in the system because 3 of the 25 character

places had to be allotted to the presence of 1, 2 or

4 spines (or total absence) on the palms and three

ether gnathopodal characters were included. Pelagont

coxae and enlarged molars received only one place

each.

The four groups of Austrosyrrhoe (now called

Austrosyrrhoe 1, 2, Ileraustroe, Priseosyrrhoe), all of

which were believed to have a small molar, fall into
NUMBER 12 4

TABLE 1.—List of generic or limiting characteristics in synopiid genera

The following characters are listed with the genera in which they occur; the alternative to each character occurs in all

genera not listed. Asterisks denote characters used in similarity analysis. Parentheses on genera denote weak conditions.

•Forehead protuberant (ploughing head): {Priscosyrrhoe), Pseudotiron, Synopia, Syrrhoe (part)

Rostrum short and blunt: Bruzelia (part), Bruzeliopsis, Garosyrrhoe, Pseudotiron, Synopia, Tiron (part)

•Ocular lobe sharp: Latacunga, Stephobruzelia, Syrrhoe (part), Tiron (part)

*Eyes present: Garosyrrhoe, Synopia, Syrrhoe (part), Tiron

Accessory eyes present: Synopia, Tiron

•Article 1 of antenna 1 elongate or bearing dorsodistal tooth: Bruzeliopsis, Syrrhoe (part)

•Article 2 of antenna 1 elongate or bearing dorsodistal tooth: Austrosyrrhoe (part), Prucosyrrhoe, Syrrhoe (part)

•Mandibular palp absent; Jeddo

Mandibular molar fully triturative: ?Austrosyrrhoe, Garosyrrhoe, Synopia, Syrrhoe, Tiron

Mandibular molar small and partially triturative: Ileraustroe, Pseudotiron

Mandibular molar small and nontriturative: Priscosyrrhoe

•Mandibular molar large and nontriturative: Bruzelia, Bruzeliopsis, Ileraustroe, Jeddo, Latacunga, Stephobruzelia, Synopia

variabilis, Syrrhoites

•Maxillipeds folicaeous: Synopia

•Coxa 1 tapering distally: Latacunga

•Coxae 3-4 pelagont: Bruzeliopsis, Ileraustroe, Jeddo, (Priscosyrrhoe), Pseudotiron (part), Synopia, Syrrhoe

•Gnathopods distinctly simple (no palmar defining spines): Pseudotiron, Synopia, Tiron

•Gnathopods with transverse palms: Garosyrrhoe, Syrrhoe

•Gnathopodal defining spines serrate: Austrosyrrhoe, Bruzelia, Garosyrrhoe, Ileraustroe, Latacunga, Priscosyrrhoe, Syrrhoe

(part)

•Dactyl of gnathopod 2 vestigial: Synopia

•Uropod 3 greatly exceeding uropods 1-2: Pseudotiron, Synopia, Syrrhoe (part), Tiron (part)

•Peduncle of uropod 3 elongate: Austrosyrrhoe, Bruzelia (part), Bruzeliopsis (part),Stephobruzelia, Syrrhoe (part),Syrrhoites

•Telson short, cleft or not: Synopia

•Telson elongate and cleft at least one-fourth: Austrosyrrhoe, Garosyrrhoe, Jeddo, Latacunga, Pseudotiron, Syrrhoe, Syrrhoites,

Tiron

Bruzeliopsis, Ileraustroe, Priscosyrrhoe, Stephobruzelia Telson elongate and entire or cleft one-fifth: Bruzelia,

Characters of poor generic value:

•Gnathopodal palmar defining spines 4 in number: Stephobruzelia

•Gnathopodal palmar defining spines 2 in number: Austrosyrrhoe (part), Bruzelia (part), Bruzeliopsis, Ileraustroe, Latacunga, Syrrhoe (part), Syrrhoites (part)

•Gnathopodal palmar defining spines one in number: Austrosyrrhoe (part), Bruzelia (part), Garosyrrhoe, Priscosyrrhoe,

Syrrhoe (part), Syrrhoites (part)

•Dactyls of pereopods 3-5 modified or reduced: Tiron (part)

•Article 2 of pereopod 5 widely expanded and ventrally truncate: untenable

•Pereopods 3-5 very short: Tiron (part)

•Pleonites 1-3 dorsally denticulate: Syrrhoe (part), Tiron (part)

the main Syrrhoites branch otherwise containing

genera with an enlarged molar. One might have expected the Austrosyrrhoe group to form a branch of

its own or fall close to the Tiron branch. In Figure

1 the highest similarities among the incipient genera

are indicated by dashed lines in order to show the

strength of connections giving rise to the phenogram.

One of several manipulations was made in testing the

system where Austrosyrrhoe group 3 (Priscosyrrhoe)

was removed entirely from the complex in order to

find where Austrosyrrhoe 1 might then be connected.

(In addition the condition of gnathopods in the

Syrrhoites cohasseta group is changed from simple to

subchelate.) Instead of a wide separation from the

base of the Syrrhoites group, Austrosyrrhoe I moved

into the Syrrhoe branch. To some extent this is a

function of the generalized plan in Garosyrrhoe but it

is also one of the vagaries in dealing with unranked

character combinations in a tightly knit group. Nevertheless the unranked analysis focuses on previously

unconsidered relationships. Despite its nearly simple

gnathopods and pelagont coxae, Jeddo falls into the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

r Ufr

Austrosyrrhoe 1

Priscosyrrhoe

Austrosyrrhoe 2

Syrrhoites serrata

Latacunga

Bruzelia typica

Bruzelia tuberculata

Ileraustroe

Bruzeliopsis

Jeddo

Syrrhoites cohasseta

Stephobruzelia

Syrrhoe crenulata

Syrrhoe affinis

Syrrhoe papyracea

Syrrhoe oluta

Garosyrrhoe

Pseudotiron longicaudatus

Pseudotiron bouvieri

Pseudotiron coas

Tiron

Synopia

Synopia variabilis

FIGURE 1.—Dendrogram of generic affinities in Synopiidae. Dashed lines denote highest

affinities between remote pairs. Twenty-five characters are used and the species fall into 23

generic groups, some of which are invalid from other interpretations.

Syrrhoites branch and remains close to Syrrhoites

cohasseta despite changing one character in S. cohasseta in another test of goodness. The latter group

is remarkably distinct from the type of Syrrhoites

despite changing this one character value.

The most readily apparent case of intermediacy

(or straight-line evolution) lies in the pairing of

Tiron and Pseudotiron coas, then connecting them to

remaining members of Pseudotiron at low level.

Pseudotiron coas distinctly combines characters of

Tiron and typical Pseudotiron. By analogy one might

use this case as a standard for evaluating similar

cases in the other branches except that the Tiron

branch is obviously a group different from the other

branches and the same taxonomic developments cannot be said to occur in the other branches nor to

have value equal to those in the Tiron branch.

Since this universal dilemma of taxonomists, the

uneven evolution of attributes occurs even within a

family or genus or species, the taxonomist must
NUMBER 12 4

gather several alternative methods of analysis and

draw conclusions from all them, in the process making value judgments and ranking characters ultimately. For example, in Figure 1, one might conclude

that the reversal in molar enlargement is a polyphyletic situation since this character (pair) occurs

alternately with small molars in the otherwise closest

pairs and triads of genera (Austrosyrrhoe 2 with

Syrrhoites, Ileraustroe ( = Austrosyrrhoe 4) with 2

groups of Bruzelia). Uncleft telsons are highly significant as generic characters.

Relationships According to Least Differences

(Six-Character Analysis)

FIGURE 2

Six valuable generic attributes occur among the

synopiid genera if one disregards eyes. Some of these

attributes are complex, being composed of two or

more parameters occurring together and counted as

a single character. The six characters are: (1) the

enlargement of coxa 3 ("pelagont"), its quadrate

shape and posterodorsal excavation combined with a

reduced and comma-shaped coxa 4; the alternative

is an ordinary rectangular, unexcavate coxa 3 with or

without an anteroventral point combined with a

reasonably large, adz-shaped coxa 4; (2) the elongation of either article 1 or 2 on antenna 1 and the

prolongation of a dorsodistal tooth easily seen from

lateral view; the alternative is a regularly short

article and the absence of a tooth or the development

of a tooth hidden on the medial edge of article 1

only; (3) telson entire; the alternative, telson cleft at

least one-third of its length; (4) three alternatives of

the gnathopods, either transversely subchelate,

obliquely subchelate, or simple; (5) the presence or

absence of conspicuous palmar spines on the gnathopods, whether one, two or four spines are present

and whether or not one or more of these spines is

serrate; (6) the enlargement of the mandibular molar

and the loss of ridges and teeth; the alternative is

the ordinary gammaridean molar with trituration

surface or remnants of that surface.

The synopiid genera are arranged in Figure 2

according to the least number of differences among

them in terms of the 6 complex characters cited

above. The figure has been constructed parisimoniously on a grid so that the lines connecting genera

represent those only having one difference between

pairs. Any two noncontiguous genera will have at

least as many, if not more differences between them

than between the most closely adjacent pairs. Boxes

are drawn to encompass genera: (1) with eyes; (2)

with pelagont coxae 3-4; (3) with enlarged and

fuzzy molar; (4) with entire telson.

This system is imperfect because so few characters

Garosyrrhoe Tiron Pseudotiron

>•

Syrrhoe

Priscosyrrhoe

Synopia Jeddo

Lj.

Austrosyrrhoe 1

Austrosyrrhoe 2

Ileraustroe Bruzeliopsis

Bruzelia Stephobruzelia

i > •

A Illl

4?._.__ _/[v

Latarunga Syrrhoites

I

it an

FIGURE 2.—Grid of 15 genera or generic groups arranged

so that each genus, as marked by a connecting line, stands

as close as possible to a similar genus having only one difference among the 6 complex characters stated in the text. If

an arrow terminates the line, an evolutionary direction is

suggested but only in a remote sense that the ancestor stands

close to a hypothetical ancestor of the descendent genus.

The descendent genera have specializations considered to be

irreversible. Boxes enclose genera with common characters

in 4 categories marked with symbols, eyes present (upper

left), molar enlarged and fuzzy (bottom right), coxae pelagont (upper right), telson long and entire (middle right).

Symbols near generic names denote conditions of palmar

defining spines on gnathopods; no symbol — spines absent

and gnathopods simple; vertical line-spines simple; vertical

line with flags = spines serrate; flagged line with top horizontal bar=bar palm transverse. Synopia (upper middle) is

the only genus with short telson. Zigzag lines near 4 genera

denote dorsal serrations on pleonites 1—3; certain species

in these genera have lost the serrations.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

are being utilized but the method reveals how the

Syrrhoites-Bruzelia group is generally divorced from

the Austrosyrrhoe group except between Latacunga

and Austrosyrrhoe 2. It shows how all pelagont

genera including Bruzeliopsis have similarities among

themselves and it brings the oculate genera reasonably close to one another. A pattern distinct from

that seen in Figure 1 is therefore apparent and one

obtains a rudimentary phyletic scheme in which the

pelagont coxae assume a direct significance. A series

of stages can be visualized in which the enlarged

molar becomes a character of specialization rather

than primitivity but insoluble difficulties emerge, as

one must explain several reversions of characters,

for example, return of Synopia to shortened telson,

and return of the Bruzelia line to nonpelagont coxae.

Evolution from Oculate Ancestors

FIGURE 2

If one assumes that the loss of eyes in deep-sea

amphipods is irreversible, and because a few littoral

synopiids today bear eyes, one must then assume that

the immediate ancestors of synopiids carried eyes and

lived in shallow seas. Synopiids may have found

especially favorable ground for evolution in deep

water, or the present shallow-water members may be

a fragment of a former diverseness now retracted perhaps in favor of the more abundant Oedicerotidae,

Haustoriidae and Phoxocephalidae, which have similar ecological adaptations. The Oedicerotidae also

have galeate heads but the accessory flagellum is

vestigial, pereopod 5 is greatly extended, the peduncle

of uropod 3 is elongate and the telson is very short

and entire.

The hypothetical ancestor of the synopiids thus

had eyes, a short telson, a normally triturative

mandibular molar, regular anterior coxae, subchelate

gnathopods and was probably a shallow sublittoral

benthic organism. No living synopiid quite fits that

definition, although Synopia, a neritic, if not a pelagic

genus, has the eyes, short telson, and weakly subchelate gnathopods; but the coxae are pelagont, the

ecological position is wrong, and the maxillipeds are

heavily foliaceous and setiferous. The latter mouthparts suggest Synopia is a bottom feeder and therefore

the ecological position alluded to above may be

correct.

Tiron, a sublittoral benthic genus, appears closely

related to Synopia in eyes and accessory eyes, but

like all other synopiids has the telson elongate.

Coxae 3-4- are reasonably basic, the maxillipeds are

basic, but the gnathopods are simple and therefore

highly advanced.

Garosyrrhoe, a 2-species genus from the eastern

Pacific Ocean also occurs sublittorally, though we are

uncertain whether it is actually of neritic provenance

since so few specimens have yet been collected in

bottom grabs, a sign that they were only accidentally

obtained on the bottom. Garosyrrhoe also has eyes,

no accessory eyes, ordinary coxae and maxillipeds,

but the telson is elongate, and the mandibular molar

is poorly triturative. The gnathopods, though small

and elongate, are transversely subchelate and have a

spine similar to the dactyl that gives the appearance

of chelateness to the gnathopods. Garosyrrhoe is the

logical precursor to the final genus bearing eyes,

Syrrhoe. Presumably Syrrhoe is an epibenthic genus

extending from sublittoral to abyssal depths; some

of its members have apparently lost their eyes, although there is some suggestion that certain members

with weak eyes are actually strong vertical migrators.

Syrrhoe differs from Garosyrrhoe in the enlarged and

excavate coxa 3 and small coxa 4 typical of Synopia.

This coxal arrangement also occurs in Jeddo, Bruzeliopsis, Pseudotiron and weakly in certain members

of the Austrosyrrhoe complex (Priscosyrrhoe) so as

to suggest that the stronger coxal condition marks

true pelagonts whereas the weaker condition marks

weakly epibenthic species that may spend time both

on the bottom and in the nekton, perhaps darting

and swimming from one benthic feeding station to

another. Momentarily, I have disregarded the presence of this coxal arrangement as a special mark of

phyletic affinity mainly on the grounds developed in

Figure 1. On the surface it rather appears as a

case of repeated adaptation to nonbenthic existence.

One good exception to this opinion however is the

smaller difference between Syrrhoe and Pseudotiron

than between Syrrhoe and its presumed ancestral line

represented by Garosyrrhoe. Syrrhoe and Pseudotiron

differ mainly in the chelate and simple gnathopods

respectively.

In retrospect, Synopia differs from the hypothetical

ancestor by one main character, advanced coxae,

whereas Tiron differs, like all other synopiids, by the

elongate telson but also by the fully simple gnatho-
NUMBER 12 4 11

pods. Garosyrrhoe differs in the elongate telson and

poorly triturative molar.

Pseudotiron appears to be an advancement from

Syrrhoe in which the eyes have been lost, the gnathopods simplified and frequently a shovel-like or sharknose head has been developed. Since specimens of

Pseudotiron are frequently caught in benthic samplers

and their midguts contain silt grains, one might assume they plow the substrate but also swim epibenthically. In contrast, the chelate gnathopods of

Syrrhoe suggest that, as pelagonts, members of that

genus may grasp some other member of the nekton.

But gut contents of Syrrhoe are unstudied.

The next increment of specialization proceeds via

loss of eyes to Austrosyrrhoe, which is considered to

be closer to the main line of specialization than later

branches characterized by changes in gnathopods.

Austrosyrrhoe is the first anoculate genus with subchelate gnathopods, reasonably ordinary coxae, and a

small molar. Austrosyrrhoe is divided into 2 genera or

subgenera and unfortunately the type-species of

Austrosyrrhoe is not necessarily the least advanced.

One must construct an hypothetical ancestor to this

group that has a deeply cleft telson, 1-2 serrate spines

on the palms of the gnathopods, unmodified antenna

1, apically spinose outer rami of uropods 1—2,

posteriorly unexcavate coxa 3 (basic gammaridean)

and large coxa 4.

Group 2 of Austrosyrrhoe (rinconis and septentrionalis) is closest to this ancestor, differing only in the

slightly reduced comma-shaped coxa 4. Group 1,

composed only of the type-species (crassipes) differs

in 3 characters: loss of one spine on the gnathopods,

the development of a tooth on antenna 1, and

probably the loss of apical spines on the outer rami

of uropods 1—2. Group 3 (Priseosyrrhoe) has an

especially elongate telson in which the lobes are nearly

fused, but otherwise it appears close respectively to

groups 1 and 2, possibly as a separate descendent

group. Group 3 differs from group 1 mainly in the

telson and shares the gnathopodal spine (one), tooth

on antenna 1, naked rami of uropods and coxae 3-4

(presumably). A fourth group is removed to a new

genus, lleraustroe, as developed in the phenogram

(Figure 1). One presumes that certain character

developments are irreversible and proceed in this

direction: cleft to uncleft telson; loss of spines on

uropodal outer rami; posterior excavation on coxa 3;

and loss of one palmar spine on the gnathopods.

However the development of a tooth on antenna 1

may be reversible (later lost) as also may be the

changes noted in coxa 3. If all of these character

alternatives are reversible in evolution, the relationships of species will be very difficult to detect. Because coxa 3 is partially modified in the pelagont condition of Syrrhoe, Synopia and Pseudotiron, the

species of lleraustroe may be more powerful swimmers than groups 1 to 3 of Austrosyrrhoe. Coxal

modifications may also correlate with improved benthic ploughing during food ingestion. The development of an antennal tooth and loss of one palmar

spine on the gnathopods suggests that groups 1 and

3 (Priscosyrrhoe) have improved adaptations to

plowing mud and may spend more time on the

bottom than lleraustroe or Austrosyrrhoe. If the

number of swimming journeys is reduced the function

of grasping something upon alighting would be reduced and the palmar spine might not be required.

The next stage implies a single phyletic change, the

enlargement of the mandibular molar so as to dominate the mandible entirely. Six genera are characterized by a fully enlarged molar and they are considered to have a common ancestor although this

problem has not yet been studied thoroughly and the

assumption is based mainly on the existing parsimony. One can thus better establish a tree of relationships having fewer gross differences among the

branches for the six genera than if one aggregates the

differences among all of the six genera to distinguish

them from the various other synopiids. For example,

Jeddo differs by 3 generic characters from Syrrhoe, 2

from Pseudotiron; Bruzeliopsis differs in 3 characters

from Priscosyrrhoe and Bruzelia differs from either

lleraustroe or Priscosyrrhoe by 2 characters each.

Except for the closeness between Jeddo and Pseudotiron, none of the 6 genera with an enlarged molar

has fewer connections outside the 6 than with its

immediate relative within the 6, a linkage suggesting

parsimony within the 6 genera. I strongly suggest,

however, that this question be examined from time to

time as more species are discovered. For example,

members of Austrosyrrhoe may be on the direct and

independently assorted ancestral lines to various

species of Bruzeliopsis, Bruzelia, Latacunga and

Syrrhoites, but in each case this might require independent enlargement of the molar. See next section

on "Enlarged Molar Theory."

Syrrhoites appears to be the most primitive member
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of the group with an enlarged molar, as it bears

reasonably normal coxae, simple spines on the

gnathopods and a cleft telson. Latacunga is more

advanced in the tapering of coxa 1 but also has serrate spines on the gnathopods like Austrosyrrhoe.

Latacunga differs from Austrosyrrhoe in the enlarged mandibular molar and tapering coxa 1 and

may be an independent evolutionary line direct from

Austrosyrrhoe. Jeddo is very advanced in having the

syrrhoe-like pelagont coxae, no mandibular palp and

simple gnathopods without spines, but otherwise it

seems closest to Syrrhoites because of the cleft telson.

The next major single step is the fusion of the

telsonic lobes. Again this may be confuted as a single

precursorial event by the knowledge that it has

occurred twice in the Austrosyrrhoe complex

(Priscosyrrhoe and Ileraustroe). Thus Bruzelia, with

an entire telson, resembles Latucunga more than it

does Syrrhoites mainly because of the serrate gnathopodal spines; it therefore also resembles certain members of Priscosyrrhoe and Ileraustroe, though the

latter two have only one serrate spine on the palms of

the gnathopods and Bruzelia has 2. Bruzeliopsis resembles Priscosyrrhoe and group 1 of Austrosyrrhoe

in the tooth of antenna 1, but in having the coxae fit

those of Ileraustroe, Bruzeliopsis is even closer to that

group than to the others because of the uncleft telson.

The solution to these relationships is presently difficult; in any case the enlargement of the molar would

have to occur independently more times than the

development of pelagic coxae.

Enlarged Molar Theory

This section pursues an earlier discussion of (1) the

multiple evolution of molars and (2) the possibility

that the enlarged molar preceded the small molar. I

am not convinced that these two theses have any

great merit but a few hazy precedents are available

to support them. Since amphipod taxonomists find it

necessary to think about evolutionary simplifications,

in the sense that many families of amphipods recurrently demonstrate an evolutionary course from

complex to simple morphology, this discussion is

deemed valuable for future reference as more species

are discovered. J. L. Barnard (1969a) has discussed

briefly the process of simplification found in gammarideans, from the basic gammaridean with complex, sexually dimorphic gnathopods, to the majority

of families with monomorphic gnathopods, from the

presence to the absence of an accessory flagellum,

and many other examples including cephalization and

pygidization.

The hypothetical progress (Figure 2), from

synopiid ancestors having enlarged molars to those

descendents having normally triturative molars of the

basic gammaridean, implies that a basic synopiid

developed the enlarged molar first while retaining the

short telson so that Synopia variabilis (the aberrant

Synopia) with its advanced coxae, and bulbous

molar, can be divided off immediately. Thereafter

all synopiids except two species of Synopia descended

from a stage in which the telson became elongate.

If one uses the precedent of molarial reversal one

could also use telsonic reversal and derive Synopia

from the Garosyrrhoe line. Improbabilities already

increase in this scheme.

A certain logic nevertheless prevails from the next

hypothetical stage onward because one presupposes

that the fully blown synopiid is represented here by

the large molar, elongate telson and serrate spines

on the gnathopods. Syrrhoites stands near this stage

but no shallow-water oculate genus is anywhere near

it and this alone is a serious fault; however the

complexity of spinal representations proceeds more

smoothly through this scheme as one generally goes

from complex to simple.

Under most circumstances this scheme would seem

ludicrous if the very close relationship between the 4

groups of Austrosyrrhoe (including Ileraustroe and

Priscosyrrhoe) and 4 genera of the enlarged molar

group did not occur (Figure 2, four genera lower

right) In this simplified scheme Bruzeliopsis and

Ileraustroe have only one distinction, the spines on

the gnathopods. Bruzelia and Priscosyrrhoe also have

only 2 distinctions, the molar and article 1 of antenna

1. Only 2 distinctions occur between Latacunga and

Austrosyrrhoe 1 or between Latacunga and Austrosyrrhoe 2. This tight assortment of generic differences

involving telsonic clefts, spinal serrations, tooth presence or tooth placement on article 1 of antenna 1

and presence or absence of apical spines on the outer

rami of uropod 2 suggests a high degree of polytypy.

That the molarial difference between the main groups

above and below has been selected for the major

division would have no value to a pheneticist. The

especially close relationship in coxae between Bruzeliopsis and Ileraustroe is a good showpiece of the
NUMBE R 12 4 13

morphologist unable to decide whether coxae signify

closer relationships than do molars. If change in

molars occurred in a single phyletic funnel then

Bruzeliopsis and Ileraustroe evolved in parallel. But if

advanced coxae are made the base of a single evolutionary group, then, in the whole scheme, enlarged

molars must have evolved at least twice.

The Direction of Evolution in Characters

FIGURES 1-2

The patterns developed in Figures 1—2 demonstrate

how complicated, if not impossibly so, are the relationships among the synopiids. Ultimately the phyletics must consider the absurdity of numerous reversals

in evolutionary direction. As obvious evolutionary

trends can be seen I suggest in these patterns that

the following ultimate conditions in character developments are irreversible: (1) uncleft telson; (2)

simple gnathopods; (3) simple palmar spines (except

once)2; (4) loss of palmar spines entirely; (5)

blindness; (6) pelagont coxae; (7) loss of mandibular

palp (Jeddo).

But I am unable to suggest any logic as to whether:

(1) the enlarged fuzzy mandibular molar, once developed, can revert to a normal molar or at least

to the small molar of Austrosyrrhoe and Priscosyrrhoe bearing weak indications of triturative surface; (2) transversely subchelate gnathopods are

reversible towards subchelate and simple conditions;

(3) elongate telsons can revert to shortened telsons

as seen in Synopia.

If one accepts the development of the enlarged

fuzzy molar as a single evolutionary event confined to

one ancestor of the 6 bruzelian genera, then one is

confronted with the development of pelagont coxae

in at least 3 distinct evolutionary lines and the development of an uncleft telson several times. If, in

Figure 2, the pelagont coxae are considered as

descendent from one ancestor one is forced to conclude that enlarged molars, simple gnathopods and

simple palmar spines developed twice, and uncleft

telsons developed several places independently.

The connecting lines in Figure 2 suggest evolutionary directions only in the remotest context. The

1

 Meaning that complex spines developed from simple

spines once within the family.

arrows do suggest that the descendent genus could

not have been the ancestor in any pair, as specialized

characters occur in the genera marked as descendents.

For example, Stephobruzelia is more highly specialized than Bruzelia in palmar spines and though

Bruzelia is not necessarily the living ancestor of

Stephobruzelia, it stands in a pool of taxa which may

have been ancestral to Stephobruzelia. Syrrhoites is

more specialized than Latacunga in the palmar

spines if we accept the thesis that serrate palmar

spines were an early development lost in later evolution, but Latacunga cannot be a direct ancestor of

Syrrhoites because coxa 1 is specialized. Latacunga

therefore stands in a pool of taxa ancestral to

Syrrhoites.

Synopiidae

DIAGNOSIS.—Gammaridean with galeate head or

with plough-shaped protuberance on forehead; accessory flagellum of antenna 1 large, multiarticulate;

upper lip fleshy, ventrally truncate, rounded or

weakly incised, usually with small marginal hairs;

mandibles with 3-articulate palp (3 exceptions), molar

present and never amalgamated with spine row, latter

often vestigial; lower lip with well developed mandibular lobes, inner lobes present and separate from

each other, no extraordinarily wide space occurring

between outer lobes; maxillae 1—2 well developed

but setation variable, palp of maxilla 1 biarticulate;

inner and outer plates and palp of maxilliped well

developed, palp usually 4-articulate, rarely 3; coxae

1 and 2 large and unhidden by posterior coxae, except

coxa 1 rarely narrowed; gnathopod 1 present, 7-

articulate; gnathopods not sexually dimorphic, gnathopod 2 not enlarged; pereopods 3—5 basic, pereopod

5 not grossly longer than 4 as in Oedicerotidae;

uropods 1—3 present, all strongly biramous; telson

present.

VARIABLES (Table 1).—Head generally galeate, almost as long as pereonites 1—3 together (except in

Tiron), with a rostrum like that of a Roman helmet,

but many species have an extremely protuberant forehead presumably used for ploughing mud, the rostrum then being reduced in prominence; lateral

cephalic lobes broad, rounded or with sharp corner;

eyes present or absent, when present composed of

numerous ommatidia in two main eyes tightly appressed, if not coalesced dorsally; occasional genera
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

like Tiron and Synopia have an accessory eye on each

side below main eye, composed of one or more

ommatidia, appressed or segregated; eyes are present

only in Tiron, Syrrhoe, Synopia, Garosyrrhoe and

possibly some species of Pseudotiron.

Antennae generally elongate; peduncular articles of

antenna 1 untelescoped, of variable length, article 3

long or short, occasionally either article 1 or 2 also becoming elongate and bearing apicodorsal tooth or not;

most genera with apicomedial tooth on article 1 (this

tooth hidden from lateral view and not otherwise

considered taxonomically herein, in contrast to apicodorsal tooth); accessory flagellum large, generally

at least 2-articulate if not further articulate, basal

article often elongate in males (conjoint) ; basal part

of primary flagellum usually conjoint in males and

bearing brushes of setules or setae (or Paesthetascs);

articles 4—5 of antenna 2 elongate but especially so

in many males, dorsal margins often with short brushes

of setules, flagellum also elongate in many males.

MANDIBLES.—In Tiron, Synopia and Pseudotiron

the molar is columnar and triturative as in the basic

gammaridean, the incisor projects and is weakly

toothed; the lacinia mobilis occurs on both mandibles

(right and left) and also is toothed, the spine row

contains a normal number of spines (example, 8). In

Synopia variabilis, however, the spines are lost; in

Syrrhoe and Garosyrrhoe the molar becomes less

triturative, the spines fewer and the incisors and

lacinia mobili poorly toothed. In Priscocyrrhoe and

Austrosyrrhoe the molar is nontriturative, if not

minutely fuzzy. In the remaining genera such as

Bruzelia, Bruzeliopsis, Jeddo, Syrrhoites, Stephobruzelia, Latacunga and lleraustroe, the molar is enlarged, fuzzy and nontriturative; the lacinia mobilis,

spines and incisor become less distinct from one

another, although they remain distinct in lleraustroe

while almost fully amalgamated in Bruzelia; remnants

remain in Jeddo and certain species of Bruzelia. In

the latter genus the mandible is extremely bulky

and the molar dominates the mandible entirely; the

incisor is untoothed or nearly so in Bruzelia,

Syrrhoites and Latacunga. The mandibular palp is

absent in Jeddo and two species of Tiron but otherwise present and 3-articulate, article 3 being very

short in all but Bruzeliopsis and apparently always

shorter than article 1; the palp is heavily dominated

by the mandibular body in those genera with extremely enlarged molar.

LOWER LIP.—The main variable appears to be the

apical division of the outer lobes in all species of

Syrrhoites and certain species of Bruzelia.

MAXILLA 1.—Generally the inner plate is richly

furnished with marginal setae but one species of

Tiron lacks setae on the inner plate; the outer plate

is basic and the palp 2-articulate.

MAXILLA 2.—The inner plate usually has a long,

medial, submarginal row of setae except in one species

of Tiron.

MAXILLIPEDS.—Synopia has a foliaceous niaxilliped

with 3-articulate palp in one species or article 4 extremely reduced in the others.

COXAE.—Coxa 1 is generally broad except in Latacunga; the ordinary condition of coxa 3 is rectangular

but in many species the anteroventral margin either extends to a point or the coxa becomes widely expanded

apically, the posterior margin becoming deeply excavate and in the ultimate condition known as

"pelagont," the coxa becomes a shield similar to coxa

4 in many other gammaridean families; coxa 4 in

synopiids is ordinarily adz-shaped, never becomes an

enlarged shield as in other gammarideans except in a

small way in Garosyrrhoe; in the ultimate pelagont

condition coxa 4 is reduced in size, is dominated by

coxa 3, fits the posterodorsal margin of coxa 3 and

is usually comma-shaped. The pelagont condition

usually follows generic lines but tendencies towards

its development can be seen in species of nonpelagont

genera. For example Pseudotiron coas falls between

the ordinarily nonpelagont Tiron and the typical fully

pelagont members of Pseudotiron.

GNATHOPODS.—Article 6 is slender, simple, and

elongate in Tiron and Pseudotiron, but not as elongate as article 5; in Synopia article 6 varies from

slightly to strongly tumid, from simplicity to apparent

subchelation but no spines define the heavily setose

palm; in the remaining genera article 6 is considered

as subchelate because a palm defined by spines is

present, although in Jeddo and some species of

Syrrhoites the hand is very slender and appears simple

except for the presence of the defining spine(s); in

Syrrhoe and Garosyrrhoe the palm is nearly transverse

and the defining spine so enlarged as to form a

chela; in various genera the defining spine(s) are

either simple or serrate; the simple spines generally

have a setal trigger; the main evolutionary lines of

synopiids appear to be defined by simple, obliquely

palmate and transversely palmate gnathopods. In
NUMBER 1 2 4 15

Synopia the dactyl of gnathopod 2 becomes vestigial.

PEREOPODS 1—2.—Few gross taxonomic features

occur on these appendages; variation in setation or

spination of article 5 is especially apparent and

certain species have article 6 slightly inflated and

heavily setose, tending towards prehensibility.

PEREOPODS 3—5.—The main variables concern

quantitative distinctions in length relative to body

size in different species or genera, in the degree of

expansion and ventral truncation of article 2 on

pereopod 5 (especially) and in Tiron the alternatives

between reasonably ordinary dactyls and the kind

herein called stubby, in which the dactyl becomes

very shortened, almost gnarled, though it may then

bear sharp setiferous extensions.

UROPODS 1—2.—The outer rami are slightly to

strongly shorter than the inner rami and the latter

may become excessively elongate and broad. The

posteriorwards extension of these uropods is generally

a function of the length of uropod 3 or the length

of urosomite 3, although together they may be shortened or uropod 1 alone may be shortened; in these

cases they do not extend posteriorly as far as uropod

3. The outer rami particularly vary from group to

group, especially in Austrosyrrhoe, Ileraustroe and

Priscosyrrhoe in the presence or absence of apical

spines; this may prove to have generic significance.

UROPOD 3.—The peduncle may be very short and

the rami elongate, and broadly lanceolate, or the

peduncle is often elongate and the rami become more

strongly styliform, resembling the typical condition of

oedicerotids. The degree of posteriorwards extension

of uropod 3 depends on the elongation of its parts

and the elongation of urosomite 3 or instead the shortKey to the Genera of Synopiidae

1. Both telson and peduncle of uropod 3 very short and subequal in length, mandibular palp

extremely stout Synopia

Telson elongate, exceeding peduncle of uropod 3 even when peduncle elongate, mandibular

palp not extremely stout 2

2. Gnathopods simple, sixth articles elongate 3

One or both gnathopods subchelate, sixth articles short 4

3. Rostrum with sharp apex, eyes present, uropod 1 reaching apex of uropod 2 Tiron

Rostrum with blunt apex, eyes absent, uropod 1 failing to reach apex of uropod

2 Pseudotiron

4. Gnathopodal palms transverse or nearly so, defined by enlarged spine(s) 5

Gnathopodal palms oblique and defined by enlarged spine(s) 6

5. Coxae 3-4 pelagont Syrrhoe

Coxae 3-4 not pelagont Garosyrrhoe

6. Mandibular molar of medium size, occasionally smooth and minutely fuzzy but usually

triturative and generally columnar or subcolumnar, body of mandible stout but not

extraordinarily bulky or subglobular, palp relatively strong 8

Mandibular molar very large and smooth, minutely setulose or fuzzy, molar completely

dominating mandible, body of mandible bulky, subglobular, together with molar often

dwarfing palp (exceptions in Ileraustroe and Jeddo) 8

7. Telson very long and narrow, entire Priscosyrrhoe

Telson of medium length, broad, cleft about halfway Austrosyrrhoe

8. Coxae 3-4 pelagont 9

Coxae 3-4 not pelagont 11

9. Telson deeply cleft, mandibular palp absent Jeddo

Telson cleft one-fourth or less, mandibular palp present 10

10. Antenna 1 with apicodorsal tooth on article 1, telson entire or apical cleft

vestigial Bruzeliopsis

Antenna 1 lacking apicodorsal tooth on article 1, telson cleft about one-fifth. Ileraustroe

11. Telson cleft one-third or more 12

Telson entire 13

12. Palms of gnathopods bearing 1-2 serrate spines, coxa 1 narrow and tapering

distally Latacunga

Palms of gnathopods bearing 1-2 simple spines, coxa 1 not tapering Syrrhoites

13. Palms of gnathopods bearing 1-2 serrate spines Bruzelia

Palms of gnathopods bearing about 4 simple spines Stephobruzelia
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ening of uropods 1-2. The outer ramus of uropod 3

may be 1- or 2-articulate, that second article may

be long or short and the rami may be moderately

setose, only setulose, or spinose. Males may or may

not have increased setosity.

TELSON.—The telson is highly distinct among the

various groups of genera. It is very short in Synopia,

not exceeding the apex of the peduncle on uropod 3;

it is as broad as long and either cleft or entire and

trifoliate. All remaining genera have a moderately to

fully elongate telson, longer than broad, and exceeding the peduncle of uropod 3 even if that peduncle

is elongate. In Syrrhoites the telson is broad and

strongly cleft, in Bruzelia it is broad and entire; in

Ileraustroe it is very elongate, narrow and cleft onefifth; and finally in Priscosyrrhoe it is like that in

Ileraustroe but uncleft. These represent the present

extent of variations found in the family.

BODY SEGMENTS.—Pereonites 1-3 or 1-4 are generally very short, whereas pleonites 1—3 are very long

and massive; some kind of dorsal ornamentation

occurs in all synopiids except Synopia and extremely

weak posterodorsal teeth occur in at least one species

of that genus. Dorsal teeth are most strongly developed in Bruzelia and Syrrhoites. Double teeth in

tandem occur on some posterior segments of certain

species of Bruzelia; teeth may or may not occur on

urosomites; if they do occur they are usually larger

on pleonite 5 in males than in females but this is a

point needing additional study. Few species have lateral bulges or teeth on pleonites and pereonites; occasionally coxae also have bulges or ridges and teeth.

Posterodorsal serrations occurring transversely are

developed on certain species of Tiron, Pseudotiron,

Syrrhoe and Garosyrrhoe. Epimera vary widely in

terms of teeth, serrations, bulges and quadrate corners. The elongation of urosomite 3 is especially

developed in Pseudotiron.

Austrosyrrhoe K. H. Barnard, revised

Austrosyrrhoe K. H. Barnard, 1925: 354.—J. L. Barnard,

1964: 26

DIAGNOSIS.—Forehead not protuberant, lateral

cephalic lobe not sharp; eyes absent; mandible with

palp, molar of medium size and not dominating

mandible, weakly to strongly triturative; articles 1-2

of antenna 1 either basic or article 2 elongate and

bearing apical tooth; coxa 1 ordinary; coxae 3-4 not

pelagont or weakly so; gnathopods typically subchelate, palms oblique and bearing at least one serrate

spine, occasionally gnathopod 2 with obsolescent

transverse palm; dactyl of gnathopod 2 usually

normal; pereopods 3—5 elongate, dactyls elongate,

article 2 of pereopod 5 weakly or strongly expanded,

rounded or truncate ventrally; pleonites 1-3 not

denticulate; uropod 3 not grossly exceeding uropods

1-2, peduncle elongate; telson of medium length and

cleft halfway.

TYPE-SPECIES.—A. crassipes K. H. Barnard, 1925.

REMARKS.—This genus remains confounded because of uncertainties in the type-species, the unique

type specimen of which has deteriorated (letter from

the late K. H. Barnard and my examination of it in

Capetown, November 1968). Austrosyrrhoe remains

a catchall of species with presumed unenlarged molar

distinct from that of Ileraustroe and Bruzelia, bears

a deeply cleft telson and oblique gnathopodal palms

with one enlarged serrate spine. Species of Austrosyrrhoe and Priscosyrrhoe are of vital interest in attempting to determine whether the small mandibular

molar is an evolutionary apex with origins in the

Bruzelia group or whether the Austrosyrrhoe molar

represents the condition primordial to the later evolution of the Syrrhoites-Bruzelia complex. The members of Austrosyrrhoe are the only species in

Synopiidae with distinctly oblique palms that lie outside the Bruzelia complex; the Tiron group has simple

gnathopods, the Syrrhoe group has transverse palms

and Synopia is problematical.

Two groups of species occur in Austrosyrrthoe. Two

groups, now placed in Ileraustroe and Priscosyrrhoe

were removed after similarity analysis (Figure 1).

Some of the species now placed in Austrosyrrhoe

were formerly in Syrrhoites. Because the precise shape

of coxa 3 and armaments on uropods 1—2 are unknown in the type-species, it seems unwise to divide

the two groups generically; many more species of

deep-sea synopiids undoubtedly remain to be

described.

DIAGNOSES OF SUBGENERIC GROUPS IN Austrosyrrhoe.—

1. Article 2 of antenna 1 bearing apical tooth; coxa 3 not

adequately known; coxa 4 apparently large and adzshaped; gnathopods with one palmar spine; outer

rami of uropods 1-2 not definitely known but presumed from the description to have simple outer rami

but much room for doubt is possible in light of other

similarities to group II A. crassipes

2. Article 2 of antenna 1 simple; coxa 3 subquadrate and

scarcely excavate posteriorly, coxa 4 medium and
NUMBER 12 4 17

strongly adz-shaped; gnathopods with 2 palmar spines

(male of A. fimbriatus aberrant and female not

definitely known in this regard) ; outer rami of

uropods 1-2 apically spinose.

A. rinconis, septentrionalis, ?fimbriatus

Austrosyrrhoe crassipes K. H. Barnard

Austrosyrrhoe crassipes K. H. Barnard, 1925: 354-355, pi.

34, fig. 11.

DIAGNOSIS.—Rostrum large (but sharpness unknown) ; article 2 of antenna 1 with apical tooth;

coxa 3 with acute anteroventral corner, (posterior

margin unknown) ; coxa 4 large, thick, (presumably

adz-shaped) ; gnathopod 2 with palm; article 2 of

pereopod 5 scarcely expanded, rounded posteroventrally; (outer rami of uropods 1-2 not definitely

known and originally compared with Syrrhoe

crenulata, in which outer rami are simple) ; telson

short and broad (presumably in the context of this

genus), cleft nearly halfway; (epimeron 3 unknown) ;

pleonites 1-4 each with small dorsal tooth.

Off Cape Point, South Africa, 700 fms.

Austrosyrrhoe rinconis J. L. Barnard

Austrosyrrhoe rinconis J. L. Barnard, 1967: 160-162, fig. 79.

DIAGNOSIS.—Rostrum thick and sharp; article 2 of

antenna 1 simple; coxa 3 with acute anteroventral

corner, not excavate posteriorly; coxa 4 large, thick,

adz-shaped; gnathopod 2 with palm; article 2 of

pereopod 5 moderately expanded, rounded posteroventrally; outer rami of uropods 1-2 spinose apically;

telson short and broad, cleft nearly halfway; epimeron

3 with slightly enlarged blunt posteroventral tooth;

pleonites 1-3 and 5 each with dorsal tooth.

Hand of gnathopod 1 narrow; article 2 of antenna

1 longer than article 1.

REMARKS.—The three species of group 2 are all

closely similar to one another; the appended diagnostic sentence above helps distinguish them.

Baja California, 1095-1205 m.

Austrosyrrhoe septentrionalis Stephensen

Austrosyrrhoe (?) septentrionalis Stephensen, 1931: 259-

260, fig. 74

DIAGNOSIS.—Rostrum medium, with slightly

blunted apex; article 2 of antenna 1 simple; coxa 3

with slightly acute anteroventral corner, posterior

margin weakly excavate; coxa 4 large, thick, adzshaped ; gnathopod 2 with palm; article 2 of pereopod

5 expanded, subtruncate posteroventrally; outer rami

of uropods 1-2 spinose apically; telson short and

broad, cleft halfway; epimeron 3 with slightly enlarged, sharp tooth; pleonites 1—2 each with dorsal

tooth.

Hand of gnathopod 1 very stout, article 2 of antenna 1 shorter than article 1.

Arctic North Atlantic, 885-2702 m.

Austrosyrrhoe fimbriatus (Stebbing and Robertson)

Syrrhoe fimbriatus Stebbing and Robertson, 1891: 34-36,

pi. 5 B.

Syrrhoites fimbriatus.—Stebbing, 1906: 280-281.

DIAGNOSIS.—Rostrum large and sharp; article 2 of

antenna 1 simple; coxa 3 slightly attenuate but softly

rounded anteroventrally, not excavate posteriorly;

coxa 4 large, thick, adz-shaped; female gnathopod 2

with normal oblique palm, in male hand linear, rectangular, palm transverse; article 2 of pereopod 5

moderately expanded, rounded posteroventrally;

outer rami of uropods 1—2 spinose apically; telson

short and broad, cleft halfway or more; epimeron 3

with slightly enlarged, sharp posteroventral tooth;

pleonites 1-2 and 5 each with dorsal tooth.

Hand of gnathopod 1 slender; article 2 of antenna

1 shorter than article 1.

Firth of Clyde; Irish Sea (Calf of Man), depths

unknown.

Bruzelia Boeck

Bruzelia Boeck, 1871: 149.—Stebbing, 1906: 274.

DIAGNOSIS.—Forehead weakly protuberant in type

but not in others, lateral cephalic lobe not sharp; eyes

absent; mandible with palp, molar greatly enlarged,

not triturative, fuzzy; mouthparts basic; articles 1-2

of antenna 1 basic; coxa 1 ordinary; coxae 3-4 not

pelagont, coxa 3 not strongly expanded distally except

for acute anteroventral cusp and not posteriorly excavate, coxa 4 variable, excavate posteriorly (rarely

weak), and slightly smaller or slightly larger than

coxa 3; gnathopods typically subchelate, palms

oblique, defined by 1-2 serrate spines, second spine if

present occasionally simple; dactyl of gnathopod 2

normal; pereopods 3-5 weakly to strongly elongate,

dactyls weakly elongate, article 2 of pereopod 5

typically rounded posteroventrally but truncate in
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species of Bruzelia

1. Pereonites 2-7 with distinct dorsal teeth 2

Only a few posterior segments of pereon with distinct teeth or teeth absent 4

2. Pereon and pleon with lateral teeth below dorsal teeth B. ascua

Pereon sometimes with lateral bulges just above coxae but subsidiary teeth otherwise

absent 3

3a. Article 1 of antenna 1 with small cusp; pleonites 1-2 with lateral carina; dorsal teeth

large and sharp B. australis

b. Article 1 of antenna 1 lacking cusp; no lateral carina on pleonites 1-2; dorsal teeth

large and sharp B. guayacura

c. Article 1 of antenna 1 lacking cusp; pleonites 1-2 with lateral carina; dorsal teeth

forming low crests B. tuberculata

4. Pereonite 7 with small dorsal tooth 5

Pereonite 7 lacking dorsal tooth 6

5. Palm of gnathopods 1-2 bearing one spine, pleonites 2 and 5 with dorsal tooth .... B. poton

Palm of gnathopods 1-2 bearing 2 spines, pleonites 2 and 5 smooth B. diodon

6. Coxa 4 comma-shaped; article 2 of pereopod 5 forming large, apically expanded and

truncate trapezoid B. popolocan

Coxa 4 adz-shaped; article 2 of pereopod 5 ovatorectangular 7

7. Pleonites 1-3 with weak dorsal rugosities, no teeth; article 1 of antenna 1 strongly

hatchet-shaped; outer rami of uropods 1-2 apically spinose B. pericu

One or more of pleonites 1-3 with dorsal tooth; article 1 of antenna 1 scarcely hatchetshaped; outer rami of uropods 1-2 apically simple 8

8. Only pleonite 3 with large dorsal tooth B. typica

Pleonites 1-2 and 5 with small dorsal tooth B. inlex

B. tuberculata and B. popolocan; pleonites 1-3 not

denticulate; uropod 3 not exceeding uropods 1-2,

peduncle typically short but elongate in B. tuberculata

and B. popolocan; telson elongate, entire.

TYPE-SPECIES.—B. typica Boeck (1871).

REMARKS.—The diagnoses do not properly account

for the presence or absence of a dorsal tooth on

pleonite 5 as a sexual difference. Bruzelia typica is

known to have this present in males and absent in

females; because in other species based only on females this tooth is present, it may not be a consistent

sexual difference.

Unusual characters are appended as separate sentences in the diagnoses.

Bruzelia ascua J. L. Barnard

Bruzelia ascua J. L. Barnard, 1966: 94, figs. 45,46; 1967:

162.

DIAGNOSIS.—Gnathopod 1 with 2 palmar spines,

gnathopod 2 with 1 palmar spine; coxa 3 with sharp

and attenuate anteroventral corner; coxa 4 much

shorter than coxa 3 and with posteroventral margin

excavate; coxae 5-6 with posterolateral tooth (unusual) ; article 2 of pereopods 3-5 with flat posterior

margin bearing 2-3 enlarged serrations; epimera 2—3

each with long, thin posteroventral tooth; pereonites

2-7 and pleonites 1-3 with large dorsal tooth; pleonites 4-5 with small dorsal tooth; pereonites 1-7 and

pleonites 1—3 with dorsolateral tooth; pereonites 1-7

with fully lateral tooth; pleonites 1-3 with second,

lower, dorsolateral tooth.

California, 1687-1720 m.

Bruzelia australis Stebbing

Bruzelia australis Stebbing, 1910: 590-592, pi. 50.

DIAGNOSIS.—Gnathopods 1-2 with 2 palmar

spines; coxa 3 acute anteroventrally; coxa 4 as long

as coxa 3, adz-shaped, posteroventral margin straight;

article 2 of pereopods 3—5 ovatorectangular and with

6 + enlarged posterior serrations; epimeron 2 with

small posteroventral tooth; epimeron 3 with medium

sharp posteroventral tooth; pereonites 1-7 and

pleonites 1-2 with dorsal tooth; pleonites 1-2 with

lateral carina.

Article 1 of antenna 1 with small cusp.

Off eastern Australia, 54-59 fms.
NUMBER 12 4 19

Bruzelia diodon K. H. Barnard

Bruzelia diodon K. H. Barnard, 1916: 168-169.—J. L.

Barnard, 1962a: 54, figures 44-45.

DIAGNOSIS.—Gnathopods 1—2 with 2 palmar

spines; coxa 3 acute anteroventrally; coxa 4 as long

as coxa 3, thick, adz-shaped, posteroventral margin

straight or slightly concave; article 2 of pereopods 3—5

ovatorectangular, posterior serrations small; epimeron

2 posteroventrally quadrate and slightly produced;

epimeron 3 with medium sharp tooth; pereonite 7

and pleonite 1 with dorsal tooth.

Off South Africa, 1189-2972 m.

Bruzelia guayacura, new species

FIGURES 3, 4

DIAGNOSIS.—Gnathopods 1—2 with 2 palmar

spines; coxa 3 acute anteroventrally; coxa 4 as long

as coxa 3, adz-shaped, posteroventral margin straight

or slightly concave; article 2 of pereopods 3—5 rectangular, posterior serrations of medium size;

epimeron 2 with large sharp posteroventral tooth;

epimeron 3 with large blunt posteroventral tooth;

pereonites 2—7 with medium-sized, blunt dorsal tooth

and with lateral bulge just above coxae; pleonites

1-2 also with blunt dorsal tooth; pleonite 5 with

small sharp dorsal tooth.

DESCRIPTION.—Rostrum of medium length, slender, straight, slightly exceeding lateral cephalic lobes,

latter broadly adz-shaped; article 1 of antenna 1

weakly hatchet-shaped, bluntly produced ventrodistally, article 2 subequal to 1 in length, moderately

thick, article 3 shorter than 2; coxa 1 expanded

distally, anterodistal corner blunt; anteroventral

corner of coxa 3 strongly produced and acute; coxa

4 scarcely smaller than coxa 3, ventral margin

slightly oblique, posterodistal lobe short, broad,

distally blunt, coxae 5-6 each with ventrolateral

ridge; article 2 of pereopods 3—5 slender, anterior

and posterior margins parallel except pereopod 5 with

slight distad expansion, posterior margins sparsely and

strongly serrate, remaining articles slender, posteroventral lobe of article 2 on pereopod 5 truncate

ventrally; palms of gnathopods denned by 2 large

spines, one serrate; posterior margin of article 6 of

pereopods 1-2 with several short setae but no

pectinae; body with strong dorsal ornamentation,

pereonite 1 with weak dorsal crest, following segments

with increasingly larger crests ending in blunt posterodorsal tooth, teeth becoming more acute

posteriorly, pleonites 1-2 with sharp dorsal tooth,

pleonite 3 with dorsal crest but no teeth, pleonite 4

dorsally smooth, 5 with small, sharp tooth; no segment with dorsolateral ridges or teeth; pereonites 2—7

with blunt, ridgelike ventrolateral tooth; lateral

projection of pleonite 4 above insertion of uropod I

weak, softly rounded; pleonal epimeron 1 forming

trapezoid with softly rounded corners, with distinct

posteroventral corner bearing strong, sharp tooth,

epimeron 3 with slightly convex posterior margin and

slightly upturned, slender posteroventral tooth; uropods apically damaged, peduncles of uropods 1—2 with

distolateral cusps; rami of uropod 1 and outer ramus

of uropod 2 apparently with weak, nearly fused distal

spine; telson very broad and apically acute.

HOLOTYPE.—USNM 136594, female, 3.2 nun.

TYPE-LOCALITY.—Eltanin 50, off southern Peru,

16°12'S, 74°41'W, 2599-2858 in, 15 June 1962.

MATERIAL.—8 specimens from the type-locality.

RELATIONSHIP.—The affinities of this species lie

with Bruzelia australis Stebbing (1910) from Wata

Mooli, Australia, 54—59 fins. There are no strong

characters distinguishing the 2 species and if abyssal

Peru were not so far geographically and bathymetrically from sublittoral Australia I should have considered I had B. australis in hand. The two species

resemble each other in dorsal ornamentation, coxae,

gnathopods, pereopods, uropods, telson and mouthparts and generally in antennae. Bruzelia guayacura

differs from B. australis, however, in the following

proportions: (1) the ventrodistally rounded (not

slightly cuspidate) apex of article 1 on antenna 1;

(2) the absence of a "point directed over the head"

on pereonite 1; (3) the absence of a lateral carina

on pleonites 1—2. The first 2 characters may be subject to different interpretations by different students

and the third character therefore is that of qualitative

distinction between the 2 species. Stebbing mentions

the ventrolateral carina of the pereon occurring in

B. australis but he does not mention the carinae on

coxae 5 and 6 and this may be another character

of discrimination between the two species. Stebbing

draws the posteroventral lobe on article 2 of pereopod

5 as a sharp cusp above a concave ventral margin

but this may be an erroneous interpretation as that
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Dl

FIGURE 3.—Bruzelia guayacura, new species, holotype, female, 3.2 mm, Eltanin 50;

g=juvenile, 2.9 mm.
NUMBER 12 4 21

FIGURE 4.—Bruzelia guayacura, new species, holotype, female, 3.2 mm, Eltanin 50.

lobe in B. guayacura, unless visually separated from

the underlying article 4, has the same appearance.

The posterior serrations on pereopods 3—5 of B.

australis appear to be more irregular and tightly

packed than they are in B. guayacura.

The tooth on pleonal epimeron 2 of B. guayacura

is larger in the holotype than it is on B. australis but

juveniles of B. guayacura have a much smaller tooth

(see Figure 3) and shallower posterior sinuosity above

the tooth.

I have not illustrated the accessory flagellum of

B. guayacura on the lateral view of the specimen; it

is 2-articulate, slightly longer than article 1 of the

primary flagellum and article 2 is very small. Bruzelia

australis was described as having a uniarticulate

accessory flagellum.

The dorsal teeth of B. guayacura are slightly

weaker than those of B. australis but this characterization often varies in Amphipoda phenotypically. Small

specimens (2.0-2.5 mm) have a slightly better defined

dorsal carina of urosomite 4 than does the adult

holotype.

Maxillipeds and maxillae are like those of Bruzelia

pericu, new species, in contrast to those of B. typica

(see figures herein).

Peru, 2599-2858 m.

Bruzelia inlex J. L. Barnard

Bruzelia inlex J. L. Barnard, 1967: 162-164, figure 80.

DIAGNOSIS.—Gnathopods 1-2 with 2 palmar

spines; coxa 3 with sharp and attenuate anteroventral

corner; coxa 4 as long as coxa 3, thick, adz-shaped,

posteroventral margin rounded; article 2 of pereopods

3-4 rectangular, of pereopod 5 ovatorectangular,

posterior serrations small; epimeron 2 with small posteroventral tooth; epimeron 3 with medium sharp

posteroventral tooth; only pleonites 1—2 and 5 with

small dorsal tooth.

Baja California, 1748-2398 m.

Bruzelia pericu, new species

FIGURES 5, 6

DIAGNOSIS.—Gnathopods 1-2 with 2 palmar

spines; coxa 3 anteroventrally acute; coxa 4 as long

as coxa 3, weakly adz-shaped, with elongate posteroventral lobe, posteroventral margin straight; article 2

of pereopods 3-5 ovatorectangular, posterior serrations small; epimeron 2 with small sharp posteroventral tooth; epimeron 3 with medium sharp posteroventral tooth; pleonites 1—3 with weak dorsal rugosities, no teeth; pleonite 5 with dorsal tooth.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

w

FIGURE 5.—Bruzelia pericu, new species, holotype, female, 4.3 mm, Vema 15-20; y = damaged

version of head.
NUMBER 12 4 23

Article 1 of antenna 1 strongly hatchet-shaped.

DESCRIPTION.—Rostrum short, stout, beaklike,

downturned, not exceeding lateral cephalic lobes,

latter beveled dorsally, anterior margin long, broadly

convex; article 1 of antenna 1 strongly hatchet-shaped,

article 2 subequal to 1 in length, but thin, attached to

article 1 subdistally, article 3 shorter than 2; coxa 1

distally expanded, anterodistal comer blunt; anteroventral corner of coxa 3 strongly produced and acute;

coxa 4 scarcely smaller than 3, ventral margin

oG2

FIGURE 6.—Bruzelia pericu, new species, holotype, female, 4.3 mm, Vema 15-20.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

straight, posterodistal lobe large, thin, blunt distally;

article 2 of pereopods 3—5 slender, anterior and

posterior edges parallel, posterior margin sparsely and

weakly notched; no other articles especially broad except article 4 of pereopod 5 with strong proximal

tumidity, posteroventral lobe of article 2 rounded

ventrally; palms of gnathopods defined by 2 large

spines, one serrate and one simple; posterior margin

of article 6 of pereopods 1—2 with numerous long

setal spines; body almost without dorsal ornamentation, pleonites 1—3 with very weak dorsal rugosities,

pleonite 3 with weak posterodorsal bilateral humps,

pleonite 5 with thin (broken) posterodorsal tooth;

posterolateral margin of pleonite 4 with small mammilliform cusp above insertion of uropod 1; pleonal

epimeron 1 broadly convex posteriorly, lacking distinct posteroventral corner, epimeron 2 with strong

posterior sinuosity below bulge, posteroventral corner

with medium-sized tooth, epimeron 3 with straight

posterior margin and slightly upturned tooth of

medium size at posteroventral corner; uropods badly

damaged but similar to those shown by Sars (1895)

for Bruzelia tuberculata Sars, except outer ramus of

uropod 1 about three fourths as long as inner, distolateral corners of peduncles of uropods 1—2 with

small, petiolate cusp, apices of outer rami of uropods

1—2 and inner ramus of uropod 1 with 2-3 mediumsized spines; telson typical of genus in overall shape

but slightly stouter than normal.

HOLOTYPE.—AMNH 14,401, female, 4.3 mm.

Unique.

TYPE-LOCALITY.—Vema 15-20, Caribbean Panama,

09°46.5' N, 79°37.5' W, 825-860 m, 10 November

1958.

RELATIONSHIP.—Apparently all species of Bruzelia

have a slight tendency to develop a hatchet-shape on

article 1 of antenna 1 but this species is the most

strongly developed so far reported. Bruzelia pericu

bears strong resemblance to B. tuberculata Sars

(1895: pi. 139, figure 2) because of the weakly developed if almost nonexistent dorsal body ornaments

but B. pericu differs from B. tuberculata in the development of a dorsal tooth on pleonite 5 in the

female (a character generally restricted to males), in

the stronger anteroventral tooth of coxa 3, the nonexcavate ventral margin of coxa 4, the slenderness

of article 2 on pereopods 3-5, the absence of sharp

protrusions on the midposterior margins of pleonal

epimera 1—2 and in the presence of distal spines on

the outer rami of uropods 1-2 and inner ramus of

uropod 1.

Bruzelia diodon K. H. Barnard (1916, and see J. L.

Barnard, 1962a) is very similar to B. pericu because of

resemblance in coxae, pereopods 3-5 and pleonal

epimera but B. diodon has 2 very distinct dorsal

teeth, on pereonite 7 and pleonite 1.

Bruzelia inlex J. L. Barnard (1967) has close affinities withh B. pericu but has distinct dorsal teeth

on pleonites 1-2, no distal spines on the rami of

uropods 1—2 and a relatively unmodified article 1 of

antenna 1.

Caribbean, 825-860 in.

Bruzelia (?) popolocan, new species

FIGURES 7, 8

DIAGNOSIS.—Gnathopods 1-2 with 2 palmar

spines; coxa 3 with anteroventral corner rounded;

coxa 4 as long as coxa 3, comma-shaped but large,

scarcely excavate posteriorly; article 2 of pereopods

3-4 ovate, of pereopod 5 very broadly expanded and

subtruncate ventrally, posterior serrations small;

epimeron 2 with middle posterior hump, margin then

sweeping concavely to quadrate posteroventral comer; epimeron 3 with small, thick sharp posteroventral

tooth; pleonites 1-2 with small dorsal tooth.

Article 1 of antenna 1 with sharp tooth; peduncle

of uropod 3 elongate.

NOMENCLATURE.—This species does not belong to

Bruzeliopsis because coxa 3 is not excavate and article

1 of antenna 1 is not elongate; it does not belong to

Latacunga because coxa 1 is normally expanded; and

it is excluded from Syrrhoites because one palmar

spine on gnathopods 1-2 is serrate.

The relationship to Bruzeliopsis however appears

also in the expanded article 2 of pereopod 5 but the

palp of the mandible is excessively large and article 3

exceptionally short, while the rostrum is thinner and

straighter than in Bruzeliopsis.

DESCRIPTION.—Rostrum of medium length, thin,

acute, down turned, beaklike; lateral cephalic lobes

very broad, scarcely extended forward, almost

smoothly convex anteriorly; article 1 of antenna 1

scarcely adz-shaped, with reverted medial cusp, article

2 as long but thinner than article 1, attached normally

to article 1, article 3 shorter than 2; coxa 1 rectangular, not distally expanded, anterodistal corner blunt;
NUMBER 12 4 25

FIGURE 7.—Brutelia popolocan, new species, holotype, juvenile, 2.7 mm, Vema 15-55.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8.—Bruzelia popolocan, new species, holotype, juvenile, 2.7 mm, Vema 15—55.
NUMBER 12 4 27

anteroventral corner of coxa 3 blunt and rounded;

coxa 4 slightly smaller than coxa 3 and scarcely lobed

posterodistally; article 2 of pereopods 3-5 increasingly

broadened, that of pereopods 3-4 ovate, that of pereopod 5 extremely broad, trapezoidal, expanding distally and truncate ventrally; notches on posterior

margins of medium density; articles 3-7 of pereopods

3-5 slender; palms of gnathopods defined by 2 large

spines, one serrate and one simple; posterior margin

of article 6 of pereopods 1-2 with 5-6 slender setae,

no comb, weak comb on anterior margin; body almost

without dorsal ornamentation, only pleonites 1-2 each

bearing one small posterodorsal tooth; posterolateral

margin of pleonite 4 with no tooth above insertion of

uropod 1; pleonal epimeron 1 convexly tapering ventrally, posteroventral comer of epimeron 2 with quadrate corner scarcely projecting, epimeron 3 with medium-sized posteroventral tooth, slightly sinuous

posterior margin and no serrations; both rami of uropods 1-2 and outer ramus of uropod 2 with apical

spines, rami of uropod 2 not as strongly disproportionate as in most species of Bruzelia, distolateral

corners of peduncles scarcely toothed, outer ramus of

uropod 3 clearly 2-articulate; telson not as strongly

broadened as and longer than that of most species of

Bruzelia.

HOLOTYPE.—AMNH 14,403, juvenile, 2.7 mm.

Unique.

TYPE-LOCALITY.—Vema 15-55, off Pacific Nicaragua, 12°45' N, 88°38' W, 3777-3950 m, 11 November 1958.

RELATIONSHIP.—This species has affinities with

Bruzelia inlex J. L. Barnard (1967) from the Cedros

Trench of Baja California because of the reduction

of the dorsal cuspidation to small teeth on pleonites

1-2 but differs from that species in the absence of a

distinct tooth on pleonal epimeron 2, the presence of

a reverted cusp on antenna 1, the absence of a tooth

on pleonite 5, the stouter gnathopods, the very weak

rostrum and the very much broader expansion of article 2 of pereopods 4—5, and the presence of distal

spines on some of the rami of uropods 1—2.

The present unique specimen of B. popolocan, new

species, is about the same size as B. poton, new species,

and they resemble each other in many ways but one

would have to presume they are opposite sexes of the

same species in order to unite them. Bruzelia popolocan differs from B. poton in the very thin rostrum,

the presence of 2 and not one locking spines on the

gnathopods, the presence of a strong cusp on article

1 of antenna 1, the absence of dorsal teeth on pereonite 7 and pleonite 5, and the basally broad but distally narrow pereopods 3—5.

Bruzelia popolocan differs from B. pericu J. L.

Barnard by characters of the same magnitude as those

stated for B. poton and in addition by the normally

developed article 1 of antenna 1 which is hatchetshaped in B. pericu.

The shapes of coxa 4 and article 2 on pereopod 5

are unusual in this genus. The tooth on article 1 of

antenna 1 is shared with B. australis.

Pacific Nicaragua, 3777-3950 m.

Bruzelia poton, new species

FIGURE 9

DIAGNOSIS.—Gnathopods 1-2 with one palmar

spine; coxa 3 acute anteroventrally; coxa 4 slightly

shorter than coxa 3, weakly adz-shaped, posteroventral

margin straight; article 2 of pereopods 3-5 ovatorectangular, posterior serrations small; epimeron 2

with tiny sharp posteroventral tooth; epimeron 3 with

medium thick sharp posteroventral tooth; pereonite

7 with small dorsal tooth; pleonites 1—2 and 5 with

small dorsal tooth.

DESCRIPTION.—Rostrum of medium length, stout,

beaklike, downturned, scarcely exceeding lateral cephalic lobes, latter rounded anterodorsally, anterior

margin long, scarcely convex; article 1 of antenna 1

weakly hatchet-shaped, ventrodistally and bluntly produced, article 2 subequal to 1 in length, but thin,

attached distally to article 1, article 3 shorter than 2;

coxa 1 distally expanded, anterodistal corner blunt;

anteroventral corner of coxa 3 strongly produced and

acute; coxa 4 scarcely smaller than 3, ventral margin

slightly oblique, posterodistal lobe of medium length,

broad, distally blunt; article 2 of pereopods 3-5

slender, anterior and posterior margins parallel, posterior margins sparsely and strongly serrate, of remaining articles only article 5 especially stout, posteroventral lobe of pereopod 5 rounded ventrally;

palms of gnathopods defined by one large serrate

spine; posterior margin of article 6 of pereopods 1—2

with one long seta and numerous minute pectinae;

body with weak dorsal ornamentation, pereonite 7 and

pleonites 1-2 each with weak posterodorsal tooth,

pleonite 5 with weak dorsal tooth; posterolateral mar-
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Gl

FIGURE 9.—Bruzelia poton, new species, holotype, juvenile, 2.9 mm, Eltanin 350.
NUMBER 12 4 29

gin of pleonite 4 with medium-sized tuberous cusp

above insertion of uropod 1; pleonal epimeron 1

trapezoidal, with softly rounded corners, with distinct

posteroventral comer, epimeron 2 with weak sinuosity

below bulge, posteroventral corner with minute, sharp

tooth, epimeron 3 with slightly convex posterior margin and slightly upturned tooth of medium size at

posteroventral corner; uropods apically damaged,

peduncles with distolateral cusps, rami of uropod 1

with 1 (? or more) distal spines, outer ramus of uropod 2 possibly with spine (damaged); telson typical

of genus, of medium width.

HOLOTYPE.—USNM 136596, juvenile, 2.9 mm.

Unique.

TYPE-LOCALITY.—Eltanin 350, east of Tierra del

Fuego, 55°03'S, 58°57'W, 2452 m, 4 December 1962.

RELATIONSHIP.—This species bears affinities with

Bruzelia pericu, new species, B. diodon K. H. Barnard

(1916, and see J. L. Barnard, 1962a) and B. inlex

J. L. Barnard (1967). It differs from all of them in

having a dorsal tooth on all three segments, pereonite

7 and pleonites 1-2, whereas the other species have a

lesser number of teeth on those segments. Bruzelia

pericu is distinguished markedly by the hatchetshaped article 1 of antenna 1. The weakness of the

dorsal tooth on pleonite 2 of B. poton and the congruence of almost all other characters suggests a very

close affinity to B. diodon. Bruzelia poton has only

one defining spine on the gnathopodal palms

whereas B. diodon has 2 but this may reflect only age

differences. The developmental morphology of Bruzelia is poorly known and many of the above-mentioned character differences used to distinguish the

species may simply reflect different stages of maturity

in individuals of a single species.

All of the species of Bruzelia, Syrrhoites and Austrosyrrhoe studied in this project have a marginal armament of small setal scales on the anterior edge of

article 6 of pereopod 2 but not on pereopod 1. Bruzelia poton, unlike the others, also has these scales on

article 5 of pereopod 2.

Article 5 of pereopods 3-5 is especially stout in the

new species and may be a tumidity characteristic of

juvenility, rather than a character of specific importance.

Maxillae and maxillipeds similar to those of B.

pericu but much less setose and spinose, possibly because of smaller size of holotype.

Off Tierra del Fuego, 2452 m.

Bruzelia tuberculata Sars

Bruzelia tuberculata Sars, 1895:397-398, pi. 139, fig. 2.—

Stebbing, 1906:275.—Stephensen, 1931:252; 1938:232.—

Gurjanova, 1951:589, fig. 395.—J. L. Barnard, 1962b:

73; 1966:94.

DIAGNOSIS.—Gnathopods 1-2 with 2 palmar

spines; coxa 3 with weakly attenuate and slightly blunt

anteroventral corner; coxa 4 as long as coxa 3, thick,

adz-shaped, posteroventral margin excavate; article 2

of pereopod 3 ovatorectangular, of pereopods 4-5

irregularly ovate, posterior serrations small; epimeron

2 with small sharp posteroventral tooth; epimeron 3

with medium tooth bearing ventral serrations; pereonites 3-7 with weak dorsal crest; pleonites 1—3 with

enlarged dorsal crest; pleonites 1-2 with lateral carina.

Peduncle of uropod 3 elongate.

Arctic-North Atlantic, 82-309 fins; California,

121-565 m.

Bruzelia typica Boeck

FIGURES 10, 11

Bruzelia typica Boeck, 1871:150; 1876:478-480, pi. 10,

fig. 3.—Sars, 1895:395-397, 696, pi. 138, 139, fig. 1.—

Norman, 1895:488.—Stebbing, 1906: 274-275.—Stephensen, 1926:87; 1928:229-230, fig. 46 (12-14); 1929:122,

fig. 29, 195.—Oldevig, 1933:143.—Stephensen, 1938:

232.—Gurjanova, 1951:588-589, fig. 394.—Oldevig,

1959:62.—J. L. Barnard, 1964:29.

DIAGNOSIS.—Gnathopods 1—2 with 2 spines; coxa

3 with sharp, attenuate anteroventral corner; coxa 4

as long as coxa 3, thick, adz-shaped, posteroventral

margin rounded; article 2 of pereopods 3-5 ovatorectangular, posteriorly smooth; epimeron 2 with

small, sharp posteroventral tooth; epimeron 3 with

medium thick posteroventral tooth; in female, only

pleonite 3 with sharp dorsal tooth; male pleonite 5

also with tooth.

MATERIAL.—Vema 14-58 (4).

REMARKS.—These specimens are approximately 4.0

mm in length and differ in minor ways from the

figures of Sars (1895), but none of these small variations has ever been used as a specific character in

this genus. Most of the extremely sharp corners of the

coxae and pleonite 3 are blunt in these specimens.

The posterolateral corners of pereonites 6 and 7 project more strongly, are blunter and have stronger
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURE 10.—Bruzetia typica Boeck, male, 3.9 mm, Vema 14—58; g=female, 4.0
NUMBER 1 2 4 31

rH<l "~ oG2s

FIGURE 11.—Bruzelia typica Boeck, male, 3.9 mm, Vema 14—58; g=female, 4.0 mm.

lateral eminences than those figured by Sars. The posterior edge of pleonal epimeron 3 is slightly more

convex and the second articles of pereopods 3—4 are

more slender than in northern B. typica.

The lower lip has strong medial notches on the

outer lobes; on one specimen the notches are even

deeper than shown for Syrrhoites serrata Sars (1895,

pi. 137) but on the other three they are rather shallow; perhaps they vary according to differential effects of preservation or according to the nearness of

molting; when the lower lip is strongly flattened by a

cover slip the notches practically disappear. Article 3

of the mandibular palp varies in length from side to

side.

One of the 2 males at hand is like that shown by

Sars (pi. 139), the erect protuberance of pleonite 6

being fully developed. The male figured herein lacks

that protuberance but has the rudimentary dorsal

tooth of pleonite 5 and the second antennal flagellum

is just commencing its elongation.

The ventral cephalic notch is highly variable. On

the 3.9 mm figured male the notch is easily discernible from lateral view; it is really composed of a posteroventral invagination of sclerotized tissue from one

of the mouth part somites (either maxilla 2 or maxilliped or a combination of the two) overlain by a thin

sheet of slightly invaginated marginal cephalic chit in.

In the 4.0 mm female the notch is not visible on the

left from lateral view (see figures) but is slightly evident from an oblique ventral view. The notch on the

right side is like that shown for the male. The outlines of lateral cephalic lobes and rostra very consid-
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

erably from specimen to specimen. The extremes have

been illustrated in choosing the male and female to be

figured. Presumably there is no connection between

cephalic variation and sexual dimorphism. To some

extent it appears as if notch morphology is disturbed

by large posteroventral concretions which occur irregularly in the 4 specimens available.

RECORD.—Western Mediterranean Sea, off Oran,

2070 m. Northeastern Atlantic along Norway coastline from Skraven southward, and south to British

Isles, about 160-600 m, Mediterranean Sea, 1938—

2070 m.

Bruzeliopsis Chevreux

Bruzeliopsis Chevreux, 1911:3-7.

DIAGNOSIS.—Forehead not protuberant, lateral

cephalic lobe not sharp; eyes absent; mandible with

weak palp, molar greatly enlarged, not triturative,

fuzzy; mouthparts basic; article 1 of antenna 1 elongate and bearing dorso (antero) distal tooth, article 2

lacking tooth; coxa 1 ordinary; coxae 3—4 pelagont;

gnathopods weakly subchelate, palms oblique, with

1-2 nonserrate defining spines; dactyl of gnathopod 2

normal; pereopods 3—5 elongate, dactyls elongate,

article 2 of pereopod 5 posteroventrally truncate,

weakly so in B. cuspidata; pleonites 1-3 not denticulate; uropod 3 not exceeding uropods 1-2, peduncle

typically long but short in B. turba; telson elongate,

entire, or very weakly cleft.

TYPE-SPECIES.—B. alberti Chevreux, 1911.

REMARKS.—See Bruzelia popolocan for remarks on

similarities of that species to Bruzeliopsis.

Bruzeliopsis alberti Chevreux

Bruzeliopsis alberti Chevreux, 1911:5-7, figures, 2, 3.

DIAGNOSIS.—Article 3 of mandibular palp as long

as article 2 and nearly as long as article 1; tooth on

article 1 of antenna 1 reaching about three-fourths

along article 2; coxa 4 as long as broad; article 2 of

pereopod 5 beveled posteroventrally; epimera 1-2

rounded posteroventrally; epimeron 3 with very large

posteroventral tooth; dorsal teeth on pleonites 1-2

very large; pereon and pleonite 3 lacking dorsal teeth.

Northeastern Atlantic, 4380 m.

Bruzeliopsis cuspidata (J. L. Barnard)

Bruzelia cuspidata J. L. Barnard, 1962a: 54, figure 43.

DIAGNOSIS.—Article 3 of mandibular palp shorter

than article 2, as long as article 1; tooth on article 1

of antenna 1 reaching about halfway along article 2;

coxa 4 longer than broad; article 2 of pereopod 5

rounded posteroventrally; epimera 2-3 with medium

posteroventral tooth; pereonites 6-7, pleonites 1-3

with small dorsal tooth.

Off South Africa, 4050 m.

Bruzeliopsis turba J. L. Barnard

FIGURE 12

Bruzeliopsis turba J. L. Barnard, 1964:29, figure 23.

DIAGNOSIS.—Article 3 of mandibular palp much

shorter than article 2 or article 1; tooth on article 1

of antenna 1 reaching about one-fourth along article

2; coxa 4 slightly longer than broad; article 2 of

pereopod 5 beveled posteroventrally; epimera 2—3

with small posteroventral tooth; pleonites 1-3 with

very small dorsal tooth (may appear absent in damaged specimens), reverted on pleonites 2-3.

MATERIAL.—Vema 15-20, ?male, 2.9 mm. Caribbean Panama 09°46.3'N, 79°37.5'W, 825-860 m,

10 November, 1958.

REMARKS.—This specimen comes from a sample

taken at the precise type-locality of this species. In

some parts it is better preserved than the holotype and

Key to the Species of Bruzeliopsis

1. Pleonites 1-2 lacking teeth or teeth extremely small; tooth on article 1 of antenna 1 reaching about one fourth along article 2; (article 2 of pereopod 5 beveled posteroventrally;

pleonite 3 lacking dorsal tooth) g^ turba

Pleonites 1-2 with distinct dorsal teeth; tooth on article 1 of antenna 1 reaching halfway

or more along article 2 2

2. Article 2 of pereopod 5 rounded posteroventrally; teeth of pleonites 1-2 small, pleonite 3

with dorsal tooth; coxa 4 longer than broad B. cuspidata

Article 2 of pereopod 5 beveled posteroventrally; teeth of pleonites 1-2 very large, pleonite

3 lacking dorsal tooth; coxa 4 as broad as long jj. alberti
NUMBE R 12 4 33

FIGURE 12.—Bruzeliopsis turba J. L. Barnard, male, 2.9 mm, Vema 15-20; /=ramus.

thus some refinement of details not adequately outlined in the holotype can be made. As it is much

smaller than the female holotype and possibly a male,

some of the differences might be attributed to these

circumstances. It distinctly has had a small reverted

tooth (now damaged) on the dorsal sides of pleonites

2 and 3 that were not seen in the holotype and

pleonal epimeron 3 is deeper. It has only one locking

spine on the palms of the gnathopods but this is probably a result of its juvenility. The outer ramus of uropod 3 is distinctly 2-articulate, a feature probably overlooked in the holotype, a frequent happening in such

poorly preserved individuals. The head is in much

better condition than it was in the holotype but that

reconstruction is relatively accurate.

Caribbean Sea, 825-975 m.

Garosyrrhoe J. L Barnard

Garosyrrhoe J. L. Barnard, 1964:29.

DIAGNOSIS.—Forehead not protuberant; lateral

cephalic lobe not sharp; mandible with palp, molar of

medium size, weakly triturative or fuzzy; mouthparts

basic; articles 1-2 of antenna 1 basic; coxa 1 ordinary; coxae 3-4 not pelagont, coxa 3 softly rectangular, posterior margin nearly parallel with anterior, and

nonexcavate, coxa 4 expanded midposteriorly, posterodorsal margin sloping, not concave but appearing

excavate, coxa 4 larger than 3; gnathopods subchelate, palms nearly transverse, defined by large

serrate spine giving hand chelate appearance; dactyl

of gnathopod 2 normal; pereopods 3—5 elongate,

dactyls elongate, article 2 of pereopod 5 rounded
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species of Garosyrrhoe

Pleonites 1-2 and pereonite 7 each with pair of dorsal teeth G. disjuncta

Pleonites 1-2 and pereonite 7 each with single dorsal tooth G. bigarra

posteroventrally; pleonites 1-3 not denticulate or very

weakly so; uropod 3 not exceeding uropods 1-2,

peduncle elongate; telson elongate, deeply cleft.

TYPE-SPECIES.—Syrrhoites bigarra J. L. Barnard

(1962b).

RELATIONSHIP.—The gnathopods relate this genus

to Syrrhoe but coxae 3 and 4 are distinct. In Syrrhoe

coxa 3 is very large and posterodorsally excavate while

coxa 4 is much shorter and narrower. Syrrhoites has

oblique palms on the gnathopods and the palmar

spines are relatively simple. Syrrhoites has the enlarged form of mandibular molar. Austrosyrrhoe has

oblique palms on the gnathopods, coxa 4 is slightly

smaller than coxa 3 and is distally adz-shaped, and

the telson is never deeply cleft.

Garosyrrhoe bigarra (J. L. Barnard)

Syrrhoites bigarra J. L. Barnard, 1962b: 73-75, figure 1.

Garosyrrhoe bigarra.—J. L. Barnard, 1966:94.

DIAGNOSIS.—Rostrum bluntly rounded apically;

pereonite 7 and pleonites 1—2 each with one dorsal

tooth.

Southern and Baja Galifornias, sublittoral.

Garosyrrhoe disjuncta J. L. Barnard

Garosyrrhoe disjuncta J. L. Barnard, 1969b: 224-225, figure

30.

DIAGNOSIS.—Rostrum sharp apically; pereonite 7

and pleonites 1—2 each with pair of blunt dorsal

teeth transversely adjacent to one another.

Upper Gulf of California, sublittoral.

Ileraustroe, new genus

DIAGNOSIS.—Forehead not protuberant, lateral

cephalic lobe not sharp; eyes absent; mandible with

palp, molar classified as enlarged but not strongly so,

not triturative or weakly so, fuzzy; mouthparts basic;

articles 1—2 of antenna 1 basic; coxa 1 ordinary;

coxae 3-4 pelagont or tending to be so, coxa 3 expanded distally and posterodorsal margin excavate,

coxa 4 small and intermediate between adz- and

comma-shaped; gnathopods typically subchelate,

palms oblique, defined by one serrate and one simple

spine; dactyl of gnathopod 2 normal; pereopods 3-5

elongate, dactyls elongate, article 2 of pereopod 5

subtruncate posteroventrally in type, rounded in second species but generally expanded; pleonites 1—3

not denticulate; uropod 3 not exceeding uropods 1—2,

peduncle elongate; telson highly elongate, cleft about

one-fifth its length.

TYPE-SPECIES.—Austrosyrrhoe ilergetes J. L. Barnard, 1964 (here selected).

REMARKS.—The type-species and Syrrhoites torpens J. L. Barnard (1962a) were at first placed by me

in Austrosyrrhoe as a fourth section during the early

part of this study but after establishing the dendrograms based on indices of total affinity, the generic

distinctions of these two species were recognized. Despite my initial interpretation of the mandibular

molar as belonging to the columnar or triturative

class, the two species were shown to have their greatest affinities to species of Bruzelia in all other characters. The molars are definitely not columnar and are

scarcely triturative so that they may be classed in the

enlarged and fuzzy category.

Ileraustroe differs from Bruzelia in the telsonic

elongation, the slight cleft in the telson, the subpelagont coxae. Whether the telsonic cleft is a secondary

acquisition after having passed through a stage like

that of Bruzelia is unknown. The telson is much more

elongate than that of Bruzelia; Ileraustroe may therefore have evolved from ancestors with cleft telson

independent from Bruzelia.

The affinities with Bruzeliopsis are very strong,

especially with Bruzeliopsis cuspidata which has a

weak apical cleft on the telson. The telson is highly

elongate in typical species of Bruzeliopsis but not in

B. turba. The type-species of Ileraustroe and Bruzeliopsis have the truncate kind of article 2 on pereopod 5 but the bevelment is oblique in Bruzeliopsis.

Other members of the two genera do not, however,

share precisely similar pereopod 5 so that this characteristic is not highly valuable generically in the

present interpretation. Bruzeliopsis has fully pelagont

coxae and article 1 of antenna 1 is elongate and
NUMBER 1 2 4 35

bears an anterodistal tooth. The gnathopods of Bruzeliopsis are nearly simple like those in Jeddo but they

do bear two simple denning spines whereas the

gnathopods of Ileraustroe are fully subchelate and

bear at least one serrate spine. The molars of Bruzeliopsis are also fully enlarged.

Ileraustroe ilergetes (J. L. Barnard)

FIGURES 13-15

Austrosyrrhoe ilergetes J. L. Barnard, 1964:27-28, figure 21.

Austrosyrrhoe ilergetes inconstant J. L. Barnard, 1967:155-

157, figure 77.

DIAGNOSIS.—Article 2 of antenna 1 simple; coxa 3

with acute anteroventral corner; article 2 of pereopod 5 expanded, slightly truncate posteroventrally;

epimeron 3 with small bifidation at posteroventral

corner; pleonites 1—5 each with dorsal tooth.

Article 6 of pereopods 1-2 tumid or not.

MATERIAL.—Vema 15-58 (4), 15-62 (1); Eltanin

75(1).

REMARKS.—Presumably /. ilergetes occurs throughout the world in deep bathyal (and ?abyssal) depths

for it has been recorded from the Mediterranean

Sea, off California and now from Chile. The typical

subspecies, from Mediterranean waters, is characterized by slightly inflated sixth articles of pereopods

1—2, an evenly rounded sinus above the tooth on

pleonal epimeron 3, a slightly oblique ventral edge

on article 2 of pereopod 5, and a long article 5 and

short article 6 on gnathopod 2. Ileraustroe ilergetes

inconstans (J. L. Barnard, 1967) from Baja California has a slender sixth article of pereopods 1—2,

a quadrate sinus on pleonal epimeron 3, a horizontal

ventral margin on article 2 of pereopod 5, and articles

5—6 of medium length on gnathopod 2. The Eltanin

form has an evenly rounded but weak sinus on

pleonal epimeron 3, a ventrally truncate edge on

article 2 of pereopod 5, and on gnathopod 2 has

article 5 of medium length but a long article 6. The

Eltanin form also differs from the Mediterranean

type by the longer article 3 of antenna 1.

Ileraustroe ilergetes differs from I. tor pens (J. L.

Barnard) in the acute point at the anteroventral corner of coxa 3, the presence of a sinus on the margin

above the tooth of the third pleonal epimeron, and

the presence of a small notch on the ventral margin

of article 2 on pereopod 5. Ileraustroe torpens still

requires a more exacting analysis than has been possible heretofore.

Mediterranean, 1938 m; Baja California to Middle

Chile, 1363-5690 m.

Ileraustroe torpens (J. L. Barnard)

Syrrhoites torpens J. L. Barnard, 1962a:61, figures 52, 53.

Austrosyrrhoe Ptorpens.—J. L. Barnard, 1964:28-29, figure

22.

DIAGNOSIS.—(Article 2 of antenna 1 presumably

simple); coxa 3 with softly quadrate anteroventral

corner; article 2 of pereopod 5 expanded, deeply

rounded posteroventrally; epimeron 3 with acute,

simple posteroventral corner; pleonites 1—5 each with

dorsal tooth but also teeth obsolescent on certain segments (possibly owing to damage).

Cape Basin, 2972-3045 m; ?Puerto Rico Trench,

5451-5419 m.

Jeddo J. L Barnard

Jeddo J. L. Barnard, 1962a: 54-55.

DIAGNOSIS.—Forehead not protuberant, lateral

cephalic lobe not sharp; eyes absent; mandible lacking palp, molar greatly enlarged, not triturative,

fuzzy; mouthparts basic; articles 1—2 of antenna 1

basic; coxa 1 ordinary; coxae 3-4 pelagont; gnathopods simple, lacking distinctive spines; dactyl of

gnathopod 2 normal; pereopods 3-5 elongate, dactyls elongate, article 2 of pereopod 5 rounded posteroventrally; pleonites 1—3 not denticulate; uropod

3 not exceeding uropods 1-2, peduncle elongate;

telson elongate and cleft.

TYPE-SPECIES.—/. simplisyrrhis J. L. Barnard

(1962a).

Jeddo simplisyrrhis J. L. Barnard

Jeddo simplisyrrhis J. L. Barnard, 1962a:57, figures 46,

47.

South Africa, 1861 m.

Latacunga, new genus

DIAGNOSIS.—Forehead not protuberant, lateral

cephalic lobe sharp; eyes absent; mandible with palp,

molar greatly enlarged, not triturative, fuzzy; mouth-
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Ileraustroe ilergetes (J. L. Barnard), female, 4.0 nun, Vema 15-58;

n=male, 3.4 mm, Eltanin 75.
NUMBK R 12 4 37

FIGURE 14.—Ileraustroe ilergetes (J. L. Barnard), female, 4.0 mm, Vema 15-58;

n=male, 3.4 mm, Eltanin 75.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

oGl

FIGURE 15.—Ileraustroe ilergetes (J. L. Barnard), female, 4.0 mm, Vema 15-58;

F.ltanin 75; « — edge.
NUMBK R 12 4 39

FIGUKE 16.—Latacunga latacunga, new genus, new species, holotype, female,

3.5 mm, Vema 15-62.
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 17.—Latacunga latacunga, new genus, new species, holotype, female,

3.5 mm, Vema 15-62.
NUMBER 1 2 4 41

Ul

FIGURE 18.—Latacunga latacunga, new genus, new species, holotype, female,

3.5 mm, Vema 15-62.

parts basic; articles 1-2 of antenna 1 basic, article 1

with small medioterminal tooth; coxa 1 tapering distally; coxae '5-A not pelagont, coxa 3 not expanded

distally except for anteroventral cusp, posterior margin nearly parallel with anterior margin and not

excavate, coxa 4 nearly as long as 3, surface area

subequal to 3, adz-shaped; gnathopods ordinarily

subchelate, defined by 1—2 serrate spines; dactyl of

gnathopod 2 normal; pereopods 3-5 elongate, dactyls elongate, article 2 of pereopod 5 rounded postoventrally; pleonites 1—3 not denticulate; uropod 3

not exceeding uropods 1-2, peduncle elongate; telson

elongate, deeply cleft.

TYPE-SPECIES.—Latacunga latacunga, new species

(here designated).

Latacunga latacunga, new species

FIGURES 16-18

DIAGNOSIS OF FEMALE.—Rostrum of medium

length, slender, lateral cephalic lobes narrowly truncate anteriorly; body scarcely toothed or carinate,

only pleonites 1 and 2 with small posterodorsal tooth

each; pleonites 4 and 5 each with small, posterolateral hump above insertions of uropods; pleonal

epimeron 1 large, posteroventrally quadrangular,

epimeron 2 with posterior margin swept into mediumsized tooth, epimeron 3 with large but simple posteroventral tooth; article 1 of antenna 1 with one mediodistal subconical process; posterior lobe of coxa 5

weak; one of 2 locking spines of gnathopod 1 simple,

other serrate, both spines of gnathopod 2 serrate;

dactyls of pereopods 1-2 about one-fourth as long

as sixth articles; article 2 of pereopods 3-5 of medium

width; fifth article of pereopods 3—5 scarcely longer

than fourth article and that of pereopod 4 two-thirds

as long as article 2; article 4 of pereopod 5 strongly

inflated proximally; outer rami of uropods 1—2 distally spinose; outer ramus of uropod 3 with article 2

about 40 percent of total length of article 1; telsonj

triangular, tapering its full length, cleft slightly less

than halfway.

HOLOTYPE.—AMNH 14,393, female, 3.5 mm.

TYPE-LOCALITY.—Vema 15-62, off Ecuador,

01°30'S, 82°19'W, 1363-1369 m, 3 December 1958.

MATERIAL.—2 specimens from the type-locality.

Ecuador, 1363-1369 m.

Latacunga comanita, new species

FIGURES 19, 20

DIAGNOSIS OF MALE.—Rostrum of medium length,

stout, lateral cephalic lobes narrowly truncate anteriorly; body without dorsal teeth except for long

tooth often typical of male synopiids on pleonite 5;

pleonite 4 with very small ventrally pointing cusp
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

above insertion of uropod 1; pleonal epimeron 1 large,

posteroventrally subquadrangular, epimeron 2 with

posterior margin swept into medium-sized tooth, epimeron 3 with medium-sized simple posteroventral

tooth; coxa 5 with strong posterior lobe; article 1 of

antenna 1 with one mediodistal subconical process;

both locking spines of gnathopod 1 simple, of gnathopod 2 serrate; dactyls of pereopods 1—2 half as long

as their sixth articles; article 2 of pereopods 3-5 of

medium width; fifth article of pereopods 3—5 conspicuously longer than fourth article and that of

pereopod 4 as long as article 2; article 4 of

pereopod 5 strongly inflated proximally; outer rami

of uropods 1—2 distally spinose; outer ramus of uropod 3 with article 2 about 50 percent of total length

of article 1; telson subrectangular, tapering only in

FIGURE 19.~Latacunga comanita, new species, holotype, male, 3.4 mm, Vema 15-13.
NUMBER 12 4 43

FIGURE 20.—Latacunga comanita, new species, holotype, male, 3.4 mm, Vema 15-13.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

distal one-third of its length, cleft slightly less than

halfway.

MOUTHPARTS.—Resembling those of L. latacunga,

new species, but the palp of maxilla 1 with a few

more setae on its lateral margin than in L. latacunga;

small details of the mandibles differing as shown in

the drawings.

HOLOTYPE.—AMNH 14,391, male, 3.4 mm.

Unique.

TYPE-LOCALITY.—Vema 15—13, off Caribbean

Colombia, ll^N , 75°50'W, 2875-2944 m, 8 November 1958.

RELATIONSHIP.—This species differs from the typespecies in the following characters: lack of teeth on

pleonites 1—2; stouter rostrum; the nonserrate condition of either spine on gnathopod 1, the longer fifth

articles of pereopods 3—5; the longer posterior lobe of

coxa 5; the longer dactyls of pereopods 1-2; and the

more rectangular, less triangular telson. Until males

and females of both species have been studied one

must assume that the presence of a tooth on pleonite

5 and the absence of humps on pleonites 4—5 above

the insertions of uropods 1-2 are characters of sexual

dimorphism in L. comanita. These species should be

considered as an analogous Pacific-Caribbean pair.

Caribbean, 2875-2944 m.

Priscosyrrhoe, new genus

DIAGNOSIS.—Forehead weakly protuberant, lateral

cephalic lobe not sharp; eyes absent; mandible with

palp, molar small and not dominating mandible, triturative surface obsolescent; article 2 of antenna 1

elongate and bearing apicodorsal tooth; coxa 1 ordinary; coxae 3—4 weakly pelagont; gnathopods typically subchelate, palms oblique and bearing one large

serrate spine; dactyl of gnathopod 2 normal; pereopods 3—5 elongate, article 2 of pereopod 5 weakly

expanded and ventrally rounded; pleonites 1-3 not

denticulate; uropod 3 not exceeding uropods 1-2,

peduncle elongate; telson highly elongate, entire.

TYPE-SPECIES.—Austrosyrrhoe priscis J. L. Barnard

(1967) (here designated).

RELATIONSHIP.—This monotypic genus differs from

Austrosyrrhoe in the elongate and uncleft telson but

it resembles the type-species of Austrosyrrhoe in the

tooth on article 2 of antenna 1 whereas it differs from

all other species now remaining in Austrosyrrhoe by

the weakly pelagont coxae; the coxae of the typespecies of Austrosyrrhoe are unknown.

This genus is erected on my interpretations of the

similarity analysis (Figure 1). The elongate, narrow

and entire telson is considered to be a generically

significant character. This same characteristic is typical of Ileraustroe but in that genus a small cleft occurs on the apex of the telson. Priscosyrrhoe also

differs from Ileraustroe in the small molar and the

tooth on article 2 of antenna 1.

Priscosyrrhoe priscis (J. L. Barnard)

Austrosyrrhoe priscis J. L. Barnard, 1967:157-160, figures

52 i-k, 78.

DESCRIPTION.—Rostrum large, medium, sharp;

coxa 3 with acute anteroventral corner, posterior

margin weakly excavate; coxa 4 relatively large, thick,

subquadrate and scarcely comma-shaped; gnathopod

2 with palm, gnathopods each with only one palmar

spine; article 2 of pereopod 5 moderately expanded,

subtruncate posteroventrally; outer rami of uropods

1—2 simple; telson entire, if not obscurely trifoliate;

epimeron 3 with sharp posteroventral tooth below

sinus; pleonites 1-5 each with dorsal tooth.

Article 6 of pereopods 1—2 slightly tumid.

Baja California, 842-1720 m.

Pseudotiron Chevreux

Pseudotiron Chevreux, 1895:166-170.—Stebbing, 1906:284.

DIAGNOSIS.—Forehead protuberant, lateral cephalic

lobe not sharp; eyes absent; mandible with palp,

molar of medium size, columnar and triturative;

mouthparts basic; articles 1-2 of antenna 1 basic;

coxa 1 ordinary; coxae 3-4 pelagont or weakly so or

not pelagont in P. coas; gnathopods simple, lacking

distinctive spines; dactyl of gnathopod 2 normal;

pereopods 3-5 typically elongate but short in P. coas,

article 2 of pereopod 5 typically rounded posteroventrally but truncate in P. longicaudatus; pleonites 1-3

typically denticulate dorsally but apparently smooth in

P. golens; uropod 3 greatly exceeding apices of

uropods 1-2, peduncle short; telson elongate, deeply

cleft. Pleonite 6 elongate.

Uropod 1 not reaching apex of uropod 2 (see

Tiron).

TYPE-SPECIES.—P. bouvieri Chevreux (1895).

The cyphos angle (modified from Bowman and

McCain, 1967) on the forehead refers to the generally

acute angle between the anteroposterior cephalic
NUMBE R 12 4 45

Key to the Species of Pseudotiron

1. Dactyls of pereopods 4-5 elongate, scarcely curved and bearing extremely weak subterminal inner seta, posteroventral corner of article 2 on pereopod 5 softly quadrate.

P. longicaudalus

Dactyls of pereopods 4-5 short, curved, claw-shaped, bearing large inner seta at middle,

posteroventral corner of article 2 on pereopod 5 rounded 2

2. Coxa 3 small, rectangular, and scarcely excavate posteriorly P. coas

Coxa 3 large, trapezoidal to L-shaped, posteriorly excavate 3

3. Coxa 3 angularly and regularly L-shaped P. bouvieri

Coxa 3 with sweeping posterodorsal curve 4

4. Posteroventral angle of coxa 3 softly quadrate, posteroventral tail of coxa 4 as long as basal

portion, anterior margin of coxa 1 straight, uropod 1 reaching end of uropod 2.

P. pervicax

Posteroventral angle of coxa 3 rounded, posteroventral tail of coxa 4 twice as long as base,

anterior margin of coxa 1 concave, uropod 1 reaching less than halfway along rami of

uropod 2 P. golens

axis and the oblique anteroventral margin of the lateral cephalic lobe.

Diagnostic values of antennae are incomplete and

mentioned by sex where known.

Pseudotiron bouvieri Chevreux

Pseudotiron Bouvieri Chevreux, 1895:166-170.—J. L. Barnard, 1964: 29-30, figure 24.

DIAGNOSIS.—Cyphos angle approximately 75-85°;

article 1 of flagellum on female antenna 1 less than

half as long as accessory flagellum, in male this article elongate and about as long as accessory flagellum, latter 3-articulate; article 1 of flagellum on

antenna 2 not elongate; anterior margin of coxa 1

concave; coxa 3 large and evenly L-shaped, as long as

coxa 2; pereopods 3—5 progressively more elongate;

posteroventral corner of article 2 on pereopod 5

rounded and slightly serrate; article 7 of pereopods

3-5 short, curved, claw-shaped, with large inner seta

at middle; lengths of uropods 1—2 poorly known but

uropod 1 apparently reaching more than halfway

along uropod 2; pleonites 1—3 posterodorsally denticulate, pleonite 4 with 3 small posterodorsal teeth,

pleonite 5 with one long tooth.

Mediterranean Sea, 170-1938 m.

Pseudotiron coas J. L. Barnard

Pseudotiron (?) coas J. L. Barnard, 1967:164-167, figure

81.

DIAGNOSIS.—Cyphos angle approximately 70°;

article 1 of flagellum on male antenna 1 more than

twice length of accessory flagellum, latter 2-articulate;

article 1 of flagellum on male antenna 2 not elongate;

anterior margin of coxa 1 slightly concave; coxa 3

abnormal for genus, nearly evenly rectangular and

scarcely excavate posteriorly, coxa 4 thus not strongly

smaller than coxa 3; pereopods 3-4 short, pereopod

5 slightly elongate, posteroventral corner on article

2 of pereopod 5 rounded, softly quadrate on pereopods 3-4, article 7 of pereopods 3—5 short, weakly

claw-shaped, with large inner seta near middle;

uropods 1-2 poorly known; pleonites 1-3 posterodorsally denticulate, pleonites 4—5 each with small

posterodorsal tooth.

Baja California, 2667-2706 m.

Pseudotiron golens J. L. Barnard

Pseudotiron golens J. L. Barnard, 1962a:57, figures 50, 51.

DIAGNOSIS.—Cyphos angle approximately 65°;

article 1 of flagellum on female-like antenna 1 as

long as accessory flagellum, latter approximately 3-

articulate; article 1 of flagellum on female-like antenna 2 elongate; anterior margin of coxa 1 concave;

coxa 3 as short as coxa 2, broadly comma-shaped,

posterodorsal margin curving, not angular; pereopods

3-5 short, similar to each other in length, posteroventral corner of article 2 on pereopods 3-5 rounded,

article 7 short, weakly claw-shaped, with large inner

seta near middle; uropod 1 reaching about halfway

along uropod 2; pleonites 1-3 apparently smooth

dorsally, pleonites 4—5 each with small posterodorsal

tooth (latter phrase corrected from original diagnosis) .

South Africa, 4893 m.
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pseudotiron longicaudatus Pirlot

FIGURES 21-23

Pseudotiron longicaudatus Pirlot, 1934:185-189, figures 73-

75.

Reference: P. 1. greteus J. L. Barnard, 1967:167-170, figures 82, 83.

DIAGNOSIS.—Cyphos angle approximately 45°;

article 1 of flagellum on female antenna 1 half

length of accessory flagellum, in male article 1 very

elongate and twice as long as accessory flagellum,

latter 3-articulate; article 1 of flagellum on female

antenna 2 not elongate; anterior margin of coxa 1

concave; coxa 3 conspicuously longer than coxa 2,

angularly L-shaped; pereopods 4—5 (3 unknown)

long, pereopod 5 longer than pereopod 4; posteroventral corner on article 2 of pereopods 4—5 softly

quadrate; article 7 of pereopods 4-5 elongate, not

claw-shaped, with weak seta distally; uropod 1 exceeding uropod 2; pleonites 1—3 posterodorsally denticulate; pleonites 4-5 each with small posterodorsal

tooth.

MATERIAL.—Vema 15-46 (3), 15-49 (1), 15-51

(2), 15-52 (1), 15-53 (3), 15-59 (5), 15-153 (2).

REMARKS.—These specimens resemble the holotype, described from a depth of 835 meters in the

Indonesian tropics, more than do specimens reported

from the northeastern Pacific Ocean by J. L. Barnard

(1967). There are significant differences in the sexes,

not heretofore shown, and a few details not described

by Pirlot, which may have subspecific importance.

Since these details have not been described for tropical specimens it is prudent not to segregate Vema

specimens nomenclaturally.

Specimens at hand range in length from 4.25 mm

to 13.0 mm. Sexual maturity appears to be reached

at a length of about 6.5 mm when males and females

become distinguishable. Individuals of both sexes,

when smaller than about 8.0 mm, have the typical

attenuated head, but large females have the head

slightly foreshortened and deeper than do juveniles

and large males. The attenuated head is the kind

described by Pirlot for the holotype.

Extensive variation occurs in the shapes of the

processes on the anteroventral midcephalic line; apparently these variations have no relationship to

sexual dimorphism. The epistomal keel (marked

"E" in the figures) varies from sharp and beaklike

to blunt to nearly obsolescent, from specimen to specimen. The process on the upper lip also varies from

strongly projecting to evanescent; usually the two

processes, epistomal and labral, are synmorphic, both

being either sharp or blunt simultaneously. Pirlot did

not illustrate these processes for the holotype. Pseudotiron I. greteus is shown by Barnard (1967) to have

strongly projecting processes.

Coxa 3 of Vema specimens resembles that of the

Indonesian holotype, whereas coxa 3 of P. /. greteus

is strongly shortened.

Pereopod 5, article 2 of males has a different shape

than that of females (see figures herein). The female

holotype described by Pirlot has article 2 resembling

that drawn herein for females, whereas the male

holotype of P. /. greteus has article 2 resembling that

of males herein. This difference in shape must therefore be a consistent sexual difference in this epigenotype.

The process on the dorsoposterior margin of pleonite 5 is elongate in males but short in females, a

sexual difference common to synopiids and pardaliscids.

The lower lip is well preserved in Vema specimens

and is illustrated herein. Apparently that drawn by

Pirlot was crushed, for the inner lobes are more discrete than shown by him and have basolateral winglike projections.

An increase in length of the dactyls of pereopods

1—2 occurs with increasing size of the specimens at

hand.

Indonesia, 835 m; Baja California, 1720-1748 m;

Pacific Costa Rica and Panama, 1746-3563 m.

Pseudotiron pervicax J. L. Barnard

Pseudotiron pervicax J. L. Barnard, 1967:170-172, figure

84.

DIAGNOSIS.—Cyphos angle approximately 75°;

article 1 of flagellum on male antenna 1 elongate but

accessory flagellum of about 4 articles also elongate

and longer than article 1; article 1 of flagellum on

male antenna 2 not elongate; anterior margin of

coxa 1 straight; coxa 3 slightly longer than coxa 2,

neither L-shaped nor comma-shaped, apically expanded and posterodorsal margin evenly curved;

pereopods 3-5 very short, 4-5 of equal length; posteroventral corner of article 2 on pereopods 3-5

rounded; article 7 of pereopods 3—5 short, clawshaped, with large inner seta near middle; uropod
NUMBER 12 4 47

FIGURE 21.—Pseudotiron longicaudatus Pirlot, male, 7.2 mm, Vema 15-59.
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 22.—Pseudotiron longicaudatus Pirlot, a>=female, 12.0 mm, Vema 15-53;

y=male, 7.2 mm, Vema 15-59.
NUMBER 12 4 49

Mr

FIGURE 23.—Pseudotiron longicaudatus Pirlot, female, 12.0 mm, Vema 15—53; e=female;

7.5 mm, Vema 15-51.
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1 reaching apex of uropod 2; pleonites 1—3 with

single posterodorsal tooth, denticulations obsolete,

pleonite 4 with small posterodorsal tooth, pleonite 5

(male) with elongate tooth.

Baja California, 1095-1205 m.

Stephobruzelia J. L. Barnard

Stephobruzelia J. L. Barnard, 1969a: 461-462.

DIAGNOSIS.—Forehead not protuberant, lateral

cephalic lobe sharp or weakly so; eyes absent; mandible with palp, molar greatly enlarged, not triturative, fuzzy; mouthparts basic, articles 1-2 of antenna

1 basic; coxa 1 ordinary; coxae 3-A not pelagont,

coxa 3 rectangular, not distally expanded except for

anteroventral cusp, posterior margin parallel with

anterior and not excavate, coxa 4 as long as and as

large as coxa 3, subrectangular, posteroventrally excavate but ventral margin truncate, with midposterior

tooth; gnathopods typically subchelate, palms oblique,

defined by about 4 simple spines; dactyl of gnathopod

2 normal; pereopods 3-5 elongate, dactyls elongate,

article 2 of pereopod 5 rounded posteroventrally;

pleonites 1—3 not denticulate; uropod 3 not exceeding uropods 1-2, peduncle elongate; telson elongate,

entire.

TYPE-SPECIES.—Bruzelia dent at a Stephensen

(1931).

Stephobruzelia dentata (Stephensen)

Bruzelia dentata Stephensen, 1931:252-255. fig. 73.

Norwegian Basin, 1096-1996 m.

Synopia Dana

Synopia Dana, 1852:315.—Stebbing, 1906:271.

DIAGNOSIS.—Forehead protuberant, lateral cephalic

lobe not sharp; eyes present, accessory eyes present;

mandible with palp, molar large, columnar, triturative (type) or swollen and pillowlike and poorly

triturative (S. variabilis) ; mouthparts, especially

maxillipeds, subfoliaceous; articles 1-2 of antenna 1

basic; coxa 1 ordinary (if small, not tapering) ; coxae

3-4 pelagont; gnathopods simple, hands slender or

tumid, often heavily setose but lacking defining spines,

tumid hands appearing subchelate; dactyl of gnathopod 2 vestigial or very small; pereopods 3-5 elongate,

dactyls elongate, article 2 of pereopod 5 weakly basic

or tending towards truncation; pleonites 1-3 not

denticulate; uropod 3 greatly exceeding apices of

uropods 1-2, peduncle short; telson short, cleft, or

entire and trifoliate.

TYPE-SPECIES.—S. ultramarina Dana (1853).

REMARKS.—The molar in S. variabilis may be interpreted as swollen but not grossly dominating the

mandible; since the molar is nontriturative and the

telson is uncleft, S. variabilis may deserve a distinct

generic name.

The species of this genus are not well known and

have not been reviewed since Bovallius (1886), except

by Spandl (1924) who described an obviously valid

new species, S. variabilis, but relied on the literature

for his systematic review. Stebbing (1906) recognized

2 valid species and 3 dubious species. Bovallius was

able to write a key to all 6 species described at that

time, only S. variabilis being added since 1886. Stebbing synonymized S. gracilis with S. ultramarina but

there does not seem to be any justification for accepting that decision until a wide series of materials from

all oceans can be studied. Smithsonian collections appear to have only a few specimens of what is presumed to be S. ultramarina, several of which have

been dissected but little knowledge can be added

since this is the best known of the several species and

it generally fits Bovallius' concept. One important

fact is that this, the type-species of the genus, has a

strongly columnar and heavily triturative mandibular

molar, whereas S. variabilis, as illustrated by J. L.

Barnard (1965), has a swollen spheroid molar which

is poorly triturative. Presumably this genus is widely

represented in nearshore epipelagic net collections in

various plankton laboratories and probably a large

series of specimens could be assembled by studying

these collections. Whether the characteristics of several species pointed out by Bovallius are valid would

be the first problem to attack.

Two keys are given below, the first modified after

Bovallius to include S. variabilis and to eliminate the

belief that S. gracilis is simply a male of S. ultramarina, as indicated in synonymy by Stebbing (1906).

I have found in another genus, Parelasmopus, that

Dana's perception of species or at least of strong

character differences among groups of specimens was

very good and thus I am willing to accept his differences in illustrations, poor as they are, until other
NUMBER 1 2 4 51

Key to Seven Species of Synopia

1. Telson entire S. variabilis

Telson cleft 2

2. Forehead not protuberant S. orientalis

Forehead protuberant 3

3. Article 2 of pereopods 3-4 rectangular or with quadrate posteroventral corners 4

Article 2 of pereopods 3-4 ovate or with rounded posteroventral corners 5

4. Article 2 of pereopod 3 narrowly rectangular, scarcely twice as broad as article 3 and

with scarcely protruding posteroventral corner S. angustifrons

Article 2 of pereopod 3 widely rectangular, nearly thrice as broad as article 3 and with

strongly protruding posteroventral corner S. gracilis

5. Flagellar articles 2, 3, 4 .. . n, of antenna 1 with long hairs S. caraibica

Flagellar articles 2, 3, 4 .. . n, of antenna 1 lacking long hairs 6

6. Telson scarcely longer than broad, apicolateral margins smooth, each apex with one stout

spine S. scheeleana

Telson much longer than broad, apicolateral margins serrate, each apex with at least 2

setae S. ultramarina

Key to Three Species of Synopia

1. Telson entire and apically trifoliate S. variabilis

Telson cleft 2

2. Telson about 1.25+ times as long as broad, each lobe with one apical spine, apicolateral

margins of lobes smooth 5. scheeleana

Telson about twice as long as broad, each lobe with at least two apical setae, one seta

short, apicolateral margins of lobes serrate S. utramarina

good evidence is found that Synopia has only 3 valid

species. The second key considers only the 3 species

currently accepted.

Synopia angustifrons Dana

Synopia angustifrons Dana, 1853: 998, pi. 68, figure 8a-d.—

Bovallius, 1886:20-21, pi. 2, figures 36-39 (after Dana)

Tropical Pacific.

Synopia caraibica Bovallius

Synopia caraibica Bovallius, 1886: 14-15, pi. 2, figure 30.

Caribbean Sea.

Synopia gracilis Dana

Synopia gracilis Dana, 1853: 995, pi. 7a-c.—Bovallius,

1886: 18-19, pi. 2, figures 31-35 [after Dana].

Stebbing (1906) indicates this is the male of S.

ultramarina but one may note how the quadrate

second articles of pereopods 3—4 separate this species

from S. ultramarina in the key to seven species

written above.

Tropical Atlantic.

Synopia orientalis Kossmann

Synopia orientalis Kossmann, 1880: 137, pi. 15, figures 11-

13 (not seen) .—Bovallius, 1886: 21.

Red Sea.

Synopia scheeleana Bovallius

Synopia Scheeleana Bovallius, 1886: 16-18, pi. 2, figures

22-29.

?Synopia scheeleana.—Stebbing, 1888: 799-804, pi. 52.

Stebbing may be correct that his Pacific specimens

are identifiable with the Atlantic S. scheeleana but he

did not see the coxae clearly and he shows apically

notched telsonic lobes whereas Bovallius shows evenly

rounded lobes.

Tropical Atlantic, especially off the Barbadoes

(Bovallius); tropical Pacific (Stebbing).

Synopia ultramarina Dana

Synopia ultramarina Dana, 1853: 955, pi. 68, figure 6a-h.—

Bovallius 1886: 6-13, pi. 1, figures 1-21.

I have examined reputed specimens of this species in

Smithsonian collections. They fit Bovallius' drawings
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and descriptions favorably except that in Bovallius' in

toto view coxae 1-2 are erroneously attached too tightly

on the long sides to the pereonites thus obfuscating

the crescentic curve these coxae make with the ventral margin of coxa 3, which in Bovallius' drawing

hangs disjunctly below the level of coxae 1-2.

Bovallius' separate drawings of coxae 1-2 are adequate. Male antenna 1 is longer than that of the

female.

MATERIAL.—Smithsonian: USNM Ace. 57608,

Crustacea Cat. 53778, Cape San Antonio, Cuba, 25

May 1914, Thomas-Barrera Exped., coll. HendersonBartsch, id. C. R. Shoemaker (10 specimens) ; USNM

Ace. 11568, Run Bay, Bahamas, no date, surface (20

specimens).

Tropical Atlantic.

Synopia variabilis Spandl

Synopia variabilis Spandl, 1923 (not seen); 1924: 48-50,

figure 17a-g.—J. L. Barnard, 1965: 494, figure 9.

Red Sea to Micronesia.

Syrrhoe Goes

Syrrhoe Goes, 1866: 527.—Stebbing, 1906: 281-282.

DIAGNOSIS.—Forehead in type-species not protuberant but strongly so in 3 other species, lateral

cephalic lobe sharp in type but rounded in species

with protuberant forehead and others; eyes typically

present but occasionally absent, accessory eyes absent;

mandible with palp, molar small, weakly triturative;

mouthparts basic; articles 1-2 of antenna 1 usually

basic but in type article 1 bearing large posteroterminal tooth and article 2 in S. papyracea with weak

tooth, peduncle slightly elongate (female) ; coxa 1

ordinary; coxae 3-4 pelagont; gnathopods with transverse or subtransverse palms bearing enlarged serrate

defining spine giving chelate appearance to hand,

spine possibly unserrate in S. affinis; dactyl of gnathopod 2 normal; pereopods 3-5 elongate, dactyls

elongate, second articles heavily serrate or not, article

2 of pereopod 5 typically rounded posteroventrally

but in few other species becoming truncate; pleonites

1-3 typically denticulate dorsally but often smooth or

Key to the Species of Syrrhoe

1. Cephalon extended as a forehead, forming protrusive appearance as in Figure 24 2

Cephalon with anterodorsal margin curving hemispherically, dorsal and anterior margins

of head nearly perpendicular 4

2. Coxa 1 strongly broadened, distal margin about twice as broad as proximal margin S. oluta

Coxa 1 not broadened, anterior and posterior margins nearly parallel 3

3. Telson of medium length and cleft halfway S. papyracea

Telson long and cleft about three-fourths of its length 5. longifrons

4. Pereonite 7 and pleonites 1-3 each with middorsal nodule in addition to posterodorsal

carinal tooth, pereonites 6—7 each with lateral nodulocarina S. nodulosa

Supernumerary nodules absent 5

5. Pleonites 1—2 with strong serrations on posterodorsal and posterolateral margins, no distinct

middle teeth 6

Pleonites 1—2 with one distinct posterodorsal tooth and few or no rudimentary serrations

or widely separated teeth 7

6. Posterior margin of pleonite 3 including both epimera fully serrate iS. crenulata

Posterior margin of pleonite 3 with gap in serrations represented by a smooth margin on

each dorsoposterior epimeron S. affinis and S. psychrophila (see diagnoses)

7. Coxa 1 distally expanded, coxa 2 distally tapering, posterior lobe of coxa 3 more than half

as broad as full depth of coxa S. serrima

Coxae 1 and 2 evenly extended, posterior lobe of coxa 3 less than half as broad as full

depth of coxa or lobe indistinct 8

8. Proximal serration on palmar spine of gnathopods enlarged, blunt posterior lobe of coxa

3 not readily distinct from remainder, posteroventral angle not truncate ... S. tuberculata

Proximal serration on palmar spine of gnathopods small and pointed, posterior lobe of

coxa 3 readily distinguished, posteroventral angle with truncate vertical margin

S. semiserrata
NUMBER 1 24 53

bearing single dorsal tooth, uropod 3 not exceeding

apices of uropods 1-2 (possible exception in S. longifrons group), peduncle short (except S. nodulosa);

telson elongate, deeply cleft.

TYPE-SPECIES.—S. crenulata Goes (1866).

REMARKS.—Syrrhoe appears to be the most diverse

genus of Synopiidae in terms of character similarities

picked for the affinity study (Figure 1). From that

study one would assume at least 3 genera are represented by several species now assigned to Syrrhoe but

the division, if it ever was necessary, is not now prudent because no definite disjunctions seem apparent.

Undoubtedly more species remain undescribed.

Syrrhoe is divisible into 2 groups based on protuberant forehead and basic forehead. Two subgroups

in the category of protuberant forehead are represented by (1) Syrrhoe oluta and S. longifrons and

(2) S. papyracea. The latter differs from the typespecies group (S. crenulata) in the head, short telson,

presence of 2 palmar spines, unmodified antenna 1,

and slightly toothed article 2 of antenna 1. Group

1 differs from the type group also in the head, the

elongate uropod 3, basic antenna 1, dorsally unserrate

pleonites and the slightly truncate ventral margin of

pereopod 5 article 2. Various of the noncephalic

characters however occur one at a time or two at a

time in the typical group of Syrrhoe, such as the weak

dorsal serrations and development of discrete teeth

in S. nodulosa and S. tuberculata. Syrrhoe affinis

differs from the type group in the absence of eyes

and the presumed unserrate palmar spines.

Syrrhoe affinis Chevreux

Syrrhoe affinis Chevreux, 1908: 7-9, figure 4.

DIAGNOSIS.—Head not protuberant, lateral cephalic

lobe sharp (assumed) ; eyes absent; article 1 of

antenna 1 not elongate, with distomedial tooth, article

2 lacking tooth; coxa 1 slightly expanded distally,

coxa 2 tapering, coxa 3 with posterior lobe about

half total coxal height; gnathopodal palms with one

spine, apparently simple (from drawing), palm of

gnathopod 2 especially oblique; article 2 of pereopod

5 weakly truncate posteriorly, generally ovate, hind

teeth of medium size to large; pleonites 1-3 dorsally

serrate (1-2 assumed) ; epimeron 3 posteriorly serrate

on lower two thirds, with smooth gap between lower

epimeral serrations and upper body serrations;

uropod 3 apparently not exceeding uropods 1-2,

peduncle apparently short; telson elongate, cleft

about three-fourths its length.

SPECIAL CHARACTER (compare S. psychrophila).—

Coxa 2 about one-third as broad as long, with long

tapering extension.

East Atlantic, off Morocco, 851 m.

Syrrhoe crenulata Goes

Syrrhoe crenulata Goes, 1866: 527, pi. 40, figure 25.—Sars,

1895: 390-391, pi. 136.—Stebbing, 1906: 282-283.—

Shoemaker, 1930: 73.—Stephensen, 1931: 257-258.

DIAGNOSIS.—Head not protuberant, lateral cephalic

lobe sharp; eyes present; article 1 of antenna 1

elongate and with posterodistal tooth, article 2 lacking tooth; coxae 1—2 scarcely tapering distally, posterior lobe of coxa 3 less than half total coxal height;

gnathopodal palms with one serrate spine; article 2

of pereopod 5 truncate posteroventrally, posterior

teeth large; pereonite 7 and pleonites 1-3 dorsally

serrate; epimera 1-2 each with small to medium

posteroventral tooth, epimeron 3 fully serrate posteriorly (smooth gap very small); uropod 3 not exceeding uropods 1-2, peduncle short; telson elongate,

cleft two-thirds or three-fourths.

MATERIAL.—Oregon 51 (1), off Oregon, Columbia

River, 150 m.

North boreal, south to about Skagerrak, New England, Japan Sea, Oregon, generally 40—200 m.

Syrrhoe longifrons Shoemaker

Syrrhoe longifrons Shoemaker, 1964: 404—405, figure 7.

DIAGNOSIS.—Forehead protuberant, lateral cephalic

lobe blunt; eyes present; article 1 of antenna 1

not elongate but with small anterodistal tooth,

article 2 also with small tooth; coxae 1—2 scarcely

expanded distally, posterior lobe of coxa 3 half total

coxal height; gnathopodal palms with one serrate

spine; article 2 of pereopod 5 apparently expanded

and ventrally truncate, posterior teeth medium to

large; pereonite 7 and pleonites 1-3 apparently

dorsally serrate; epimeron 3 apparently posteriorly

serrate; uropod 3 apparently not exceeding uropods

1—2; telson apparently elongate and cleft two-thirds

to three-fourths.

British Columbia, shallow water.
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Syrrhoe nodulosa K. H. Barnard

Syrrhoe nodulosa K. H. Barnard, 1932: 150-151, figure 88.

DIAGNOSIS.—Head not protuberant, lateral cephalic

lobe sharp; eyes present; article 1 of antenna 1

keeled dorsally and with mediodistal tooth; article

2 lacking tooth; coxa 1 slightly expanded distally,

coxa 2 evenly extended, posterior lobe of coxa 3 half

total coxal height; gnathopodal palms with one

serrate spine; article 2 of pereopod 5 expanded, angular, then weakly rounded ventrally, posterior teeth

small; pereonite 7 and pleonites 1-3 each with 2

dorsal teeth in tandem, anterior member noduliform,

posterior member bifid transversely, pleonites 1—4 and

epimera 1-3 fully serrate posteriorly; uropod 3

scarcely exceeding uropods 1—2, peduncle elongate;

telson elongate, cleft almost to base.

The elongate peduncle of uropod 3 is unusual in

Syrrhoe.

Antarctic, tows between 160 and 500 in.

Syrrhoe oluta, new species

FIGURES 24 28

DIAGNOSIS.—Forehead protuberant, lateral cephalic

lobe not sharp; eyes absent; article 1 of antenna 1

not elongate and lacking tooth, article 2 elongate

and with mediodistal tooth; coxa 1 extremely broadened apically and acutely produced forward, coxa 2

scarcely tapering, posterior lobe of coxa 3 half or more

total coxal height; gnathopodal palms with one serrate spine; article 2 of pereopod 5 expanded and

ventrally truncate, posterior teeth small to medium;

pereonite 7 and pleonites 1-5 each with one small

to medium dorsoposterior tooth; epimera 1-2 with

small posteroventral tooth, epimeron 3 with small

double tooth; uropod 3 not exceeding uropods 1—2,

peduncle short; telson elongate, cleft more than 85

percent its length.

DESCRIPTION.—Cephalon with strong forehead,

anterior and dorsal margins forming acute angle,

rostrum sharp and pointing ventrally, rostral base

forming proximal keel separated from secondary midsagittal keel projecting anteriorly by falciform notch;

articles 1 and 2 of antenna 1 subequal in length,

article 3 about half as long, article 1 with rudimentary

(type) or strong (Oregon) distomedial cusp, article

2 with strong reverted cusp; article 4 of antenna 2

much shorter than article 5, gland cone weak; coxa

1 strongly produced forward by means of anterior

conical process but unlike other syrrhoes, coxa 1

broadly expanded, thus coxa 2 not bent forward and

coxa 3 not extended forward by anteroventral prolongation to fit unusual posterior margin of coxa 1

in sequence; coxa 2 tapering slightly distally and

2-serrate distally; coxa 4 comma-shaped; gnathopod

1 of medium stoutness for genus, gnathopod 2 extremely slender and elongate; article 6 of pereopods

1-2 stout and bearing very dense brush of distal

setae; posterior margins of second articles on pereopods 3-5 coarsely and sparsely serrate, posteroventral

corners more (Oregon) or less (Colombia) protruding; distolateral peduncular margins of uropods 1-2

with long and short cusp respectively, outer rami

distally simple; telson elongate and cleft almost to

base; body essentially without serrations, only dorsoposterior angles of pereonites 7 and pleonites 1-5

with one tooth each, that on pleonite 3 larger than

on others; no nodules present; pleonal epimera 1-2

with small posteroventral point, epimeron 3 with

sinuous posterior margin and apparently bifid at

posteroventral corner.

HOLOTYPE.—AMNH 14,402 male, 7.0 mm, pereopod 5 missing.

TYPE-LOCALITY.—Vema 15-38, off Pacific Colombia, 05°00'N, 79°04'W, 3023-3251 in, 15 November 1958.

MATERIAL.—The holotype and Oregon 81 (1);

Oregon 82 (1).

RELATIONSHIP.—This species has closest affinities

with S. papyracea Stebbing (1888) in view of its

head, but the following strong differences occur in

the new species: (1) the telson is elongate and more

deeply cleft; (2) coxa 1 is broadly expanded ventrally and therefore coxa 2 is not bent forward and

coxa 3 follows suit; coxa 3 has a smooth posterior

margin and coxa 4 is more comma-shaped; (3) no

accessory serrations occur on the pleonites: (4) the

outer ramus of uropod 1 lacks a distal spine; (5)

article 4 of antenna 2 is relatively much shorter and

the gland cone is much weaker; (6) pereopods 1-2

are more strongly developed, article 6 is stouter and

the distal setal brush is more highly elaborated; (7)

the posterior margin of article 2 on pereopod 3 (and

usually 4-5) protrudes more strongly.

Most of the differences noted above also apply to

S. longifrons Shoemaker (1964) except that the
NUMBER 12 4 55

latter has a deeply cleft telson like S. oluta.

Oregon to Colombia, 2798-3251 m.

Syrrhoe papyracea Stebbing

Syrrhoe papyracea Stebbing, 1888: 789-793, pi. 50.

DIAGNOSIS.—Forehead protuberant, lateral cephalic

lobe not sharp; eyes absent; article 1 of antenna 1

not elongate and with small mediodistal tooth, article

2 lacking tooth; coxa 1 narrow and evenly expanded

base to apex, coxa 2 scarcely tapering, posterior lobe

of coxa 3 half or slightly more total coxal height;

gnathopodal palms with one serrate and one bifid

spine; article 2 of pereopod 5 weakly expanded and

rounded ventrally, posterior teeth medium in size;

pereonite 7 and pleonites 1-4 dorsally serrate; epimeron 1 with small posteroventral tooth, epimeron 2

with large tooth, epimeron 3 with medium tooth, no

posterior serrations; uropod 3, if exceeding uropods

1-2, not strongly so (broken), peduncle short, telson

of medium length, cleft halfway.

Culebra Island, West Indies, 390 fms.

Q

FIGURE 24.—Syrrhoe oluta, new species, holotype, male, 7.0 mm, Vema 15-38.
56

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 25.—Syrrhoe oluta, new species, holotype, male, 7.0 mm, Vema 15-38.

Syrrhoe psychrophila Monod

Syrrhoe crenulata var. psychrophila Monod, 1926: 54, figure

52 AH.

Syrrhoe psychrophila. Schellenberg, 1931, 159, figure 83.—

K. H. Barnard, 1932: 149-150, figure 87.

DIAGNOSIS.—Head apparently not protuberant,

lateral cephalic lobe apparently sharp; eyes present

(Schellenberg); article 1 of antenna 1 not elongate,

with mediodistal tooth, article 2 probably without

tooth; coxae 1-2 presumably extended evenly but

K. H. Barnard and Schellenberg state coxa 2 tapering at least anteroventrally and coxa 1 expanded

slightly apically vide Schellenberg; posterior lobe of

coxa 3 presumably about one-third or one-half total

coxal height; gnathopodal palms with one serrate

spine; article 2 of pereopod 5 slightly expanded and

ventrally truncate (Schellenberg), posterior teeth
NUMBER 12 4 57

FIGURE 26.—Syrrhoe oluta, new species, holotype, male, 7.0 mm, Vema 15—38.
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 27.—Syrrhoe oluta, new species, female, 6.3 mm, Oregon 81;

y=?male, 6.4 mm, Oregon 82.
NUMBER 1 24 59

P5

FIGURE 28.—Syrrhoe oluta, new species, female, 6.3 mm, Oregon 81;

y** ?male, 6.4 mm, Oregon 82.
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

small (K. H. Barnard), pleonites 1-3 presumably

serrate dorsally (K. H. Barnard states pereonite 7

smooth) ; epimeron 3 posteriorly serrate on lower twothirds, long smooth gap between there and dorsal

serrations; uropod 3 presumably not exceeding

uropods 1-2, peduncle presumably short; telson

elongate, cleft three-fourths its length.

SPECIAL CHARACTER.—Coxa 2 about two-thirds as

broad as long (see S. affinis).

Antarctica, about 136 to 400 in.

Syrrhoe semiserrata Stebbing

Syrrhoe semiserrata Stebbing, 1888: 793-797, pi. 51.—

Schellenberg, 1938: 34.

DIAGNOSIS.—Head not protuberant, lateral cephalic

lobe rounded; eyes present; articles 1-2 of antenna

1 not elongate, lacking teeth (male); coxae 1 and 2

not tapering, posterior lobe of coxa 3 about one-third

total coxal height; gnathopodal palms with one serrate spine; article 2 of pereopod 5 of medium expansion, rounded ventrally, posterior teeth small;

pereonite 7 and pleonites 1-2 with one small dorsal

tooth each, pleonite 3 apparently with small dorsal

denticle (s) ; epimeron 1 lacking posteroventral tooth,

epimeron 2 with medium tooth, epimeron 3 with

lower half of posterior margin serrate; uropod 3

not exceeding uropod 2 but uropod 2 exceeding

uropod 1, peduncle of uropod 3 short or medium

in length; telson long and deeply cleft, perhaps threefourths its length.

SPECIAL CHARACTER.—Proximal serration on spine

of gnathopods enlarged and blunt.

Melbourne, Victoria, 33 fms; Bass Strait.

Syrrhoe serrima, new species

FIGURES 29-31

DIAGNOSIS.—Head not protuberant, lateral cephalic

lobe sharp; eyes apparently present; article 1 of

antenna 1 not elongate, with distomedial tooth,

article 2 as long as article 1 but lacking tooth; coxa

1 slightly expanded distally, coxa 2 tapering distally,

posterior lobe of coxa 3 about half or more total coxal

height; gnathopodal palms with one serrate spine;

article 2 of pereopod 5 expanded and subtruncate

ventrally, posterior teeth medium to large; pleonites

1-2 each with small dorsoposterior tooth; epimera

1-2 each with small posteroventral tooth; epimeron

3 with lower half of posterior margin serrate; uropods

poorly known; telson elongate, cleft about threefourths its length.

DESCRIPTION.—Cephalon without extended forehead, anterodorsal margin evenly hemispherical,

rostrum sharp and pointing ventrally, rostral base

with midsagittal keel separated from secondary keel

on main body of head by deep notch; article 1 of

antenna 1 with reverted distal tooth, article 2 as long

as 1, article 3 about two-thirds as long as article 2;

coxa 1 extended forward and distally expanded but

relatively narrow (compared with S. oluta), anteroventral corner rounded, posteromedial margin with

strong declivity, coxa 2 tapering distally, very narrow,

also with posterior declivity; anteroventral corner of

coxa 3 slightly extended but rounded, posterior lobe

broad and quadrate; coxa 4 comma-shaped but with

posteroventral oblique truncation, coxa 7 with posterior incision; gnathopods 1-2 of medium stoutness

for genus, large locking spine of each palm almost

evenly serrate; article 6 of pereopods 1-2 slender

and bearing distal brush of setae of medium density,

dactyls long and anteriorly serratopinnate; article 2

of pereopods 3-5 with sparse and coarse posterior serrations, posteroventral corner scarcely protruding,

pereopods 3-4 with one bulbous castellation at that

corner; peduncles of uropods 1—2 with long and

medium distolateral cusps respectively, outer rami

slender, tapering, distally simple (otherwise uropods

1-2 in poor condition); telson elongate and cleft

about three-fourths of its length, lobes tapering

distally; body essentially without serrations, only

pleonites 1-2 each with weak dorsoposterior tooth

and several accessory serrations, epimeron 1 with

strongly quadratoconvex posterior margin and small

posteroventral tooth, epimeron 2 with weaker posterior bulge, sinuous posterior margin and small posteroventral tooth, epimeron 3 with subquadrate

posteroventral plate deeply serrate on ventral half of

posterior margin.

HOLOTYPE.—AMNH 14,397, ?male, 5.8 mm.

TYPE-LOCALITY.—Vema 15-131, east of Bahia

Blanca, Argentina, 40° 14.6' S, 55°24.7' W, 1475 m,

3 April 1959.

MATERIAL.—Two specimens from the type-locality,

sex unclear.

RELATIONSHIP.—Even though this species is repre-
NUMBE R 12 4 61

FIGURE 29.—Syrrhoe serrima, new species, holotype, ?male, 5.8 mm, Vema 15-131.
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

X2

oAl ^ K

FIGURE 30.—Syrrhoe serrima, new species, holotype, ?male, 5.8 mm, Vema 15—131.

sented by 2 partially defective individuals (uropods

1-2 and antenna 1 damaged, antenna 2 missing), it

ir. sufficiently unique from other syrrhoes to warrant

description. Because materials of this genus are rarely

abundant in collections, any material is highly valuable and usually worth describing. At first this species

was believed to be an individual of Syrrhoe semiserrata Stebbing (1888, Bass Strait and Bismarck

Archipelago, 60-(160) m), but numerous small differences point to its specific identity. If the present

species has eyes they have been dispersed into 2

masses, possibly by preservational effects. A few details of Stebbing's species are unknown just as are

several characters of S. serrima. The epistome of

S. serrima forms a clavotruncatiform lobe erectly

overriding the anterior cephalic surface.

Syrrhoe serrima differs from S. semiserrata in the

broader coxa 1, the distally narrower coxa 2, the

weaker anterior extension and broader posterior lobe

of coxa 3 and apparently by the less posteriorly

directed coxa 4 extension; coxa 7 also has a posterior

notch not shown for 5". semiserrata. The large palmar

spine of the gnathopods is almost evenly serrate in

S. serrima whereas in S. semiserrata it has one large
NUMBE R 12 4 63

U2

FIGURE 31.—Syrrhoe serrima, new species, holotype, ?male, 5.8 mm, Vema 15—131.

proximal cusp plus serrations. Pereonite 7 of S. serrima is evenly truncate posterodorsally whereas on

S. semiserrata it appears to be slightly cuspidate.

Uropods 1-2 of the new species have large distolateral

cusps on the peduncle in contrast to weak or absent

cusps on S. semiserrata. The telson of S. serrima has

strongly tapering lobes, not truncate as in S. semiserrata. The latter species is based on a fully adult

male whereas S. serrima is based on specimens possibly of subadult male character and thus generally

like females. Some of the character differences might

therefore be of sexual dimorphism, a subject poorly

understood in Synopiidae.

Argentina, 1475 m.
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Syrrhoe tubcrculata Dahl

Syrrhoe tuberculata Dahl, 1954: 290-292, figures 36-41.

DIAGNOSIS.—Head not protuberant, lateral cephalic

lobe unknown; eyes present; article 1 of antenna 1

not elongate, with mediodistal tooth, article 2 apparently basic; coxa 1 evenly extended, coxa 2 apparently evenly extended, coxa 3 widely expanded

apically but lacking distinctive posterior lobe because

posterior margin extended obtusely and not truncate;

gnathopodal palms with one serrate spine; article 2

of pereopod 5 expanded, ventrally truncate, anteroventral corner especially projecting, posterior teeth

small; pereonites 6-7 and pleonites 1-4 each with

bifid dorsal tooth, posterior borders of pereonites 6-7

and pleonites 1-3 with other large teeth; epimera

1-2 with weak or sharp posteroventral tooth; epimeron 3 with lower two-thirds of posterior margin

bearing wavy serrations; uropod 3 not exceeding

uropods 1-2, peduncle unknown; telson elongate,

cleft almost to base.

Ross Sea, 550 m.

Syrrhoites Sars

Syrrhoites Sars, 1895: 391-392.—Stebbing, 1906: 279.

DIAGNOSIS.—Forehead not protuberant, lateral

cephalic lobe not sharp; eyes absent; mandible with

palp, molar greatly enlarged, not triturative, fuzzy;

mouthparts basic; articles 1-2 of antenna 1 basic,

article 1 with small medioterminal tooth; coxa 1

ordinary; coxae 3-4 not pelagont, coxa 3 rectangular,

not distally expanded, posterior margin parallel with

anterior margin and not excavate, coxa 4 nearly as

long as 3, with subequal surface area, adz-shaped;

gnathopods typically subchelate, palms oblique, defined by 1-2 unserrate spines but majority of species

with palms obsolescent though distinguished by

spine(s); dactyl of gnathopod 2 normal; pereopods

3-5 elongate, dactyls elongate, article 2 of pereopod

5 typically basic but posteroventrally truncate in

cohasseta-redox group; pleonites 1-3 not denticulate;

uropod 3 not exceeding other uropods (or rarely in

slight degree), peduncle elongate; telson elongate and

deeply cleft. Uropod 1 short in S. pusillus.

TYPE-SPECIES.—Bruzelia serrata Sars (1879).

In the following key, Syrrhoites species A is placed

in several positions with incorrect antecedents in

order to show its several points of relationship.

Extraordinary characters in certain diagnoses are

placed in separate paragraphs.

Key to the Species of Syrrhoites

1. Pleonite 3 dorsally bidentate longitudinally (condition sometimes very weak) 2

Pleonite 3 not bidentate dorsally 3

2. Palms of gnathopods each with 2 defining spines, epimeron 3 deeply serrate posteriorly

Palms of gnathopods each with one spine, epimeron 3 smooth posteriorly except for

S. serrata

posteroventral tooth S. pacifica

3. Pleonite 5 untoothed dorsally 4

Pleonite 5 with dorsal tooth 5

4. Dorsal body carina and teeth commencing strongly on pereonite 5, epimeron 3 with

medium-sized, acute posteroventral tooth S. sorpresa

Dorsal body carina and teeth commencing strongly on pereonite 6, epimeron 3 quadrate

posteToventrally, with minute tooth S. tenella

5. Epimeron 2 rounded posteroventrally, unarmed 6

Epimeron 2 quadrate or bearing tooth posteroventrally 8

6. Epimeron 3 prolonged posteroventrally but lacking distinct tooth S. walkeri

Epimeron 3 with distinct but small posteroventral tooth 7

7. Dorsal body carina and teeth commencing strongly on pereonite 2 S. lorida

Dorsal body carina and teeth commencing strongly on pereonite 6 S. cohasseta

8. Peduncle of uropods 1-2 lacking distolateral tooth or extension 9

9.

Peduncle of uropods 1-2 bearing distolateral tooth or extension 10

Telson cleft less than halfway S. redox

Telson cleft more than halfway Syrrhoites species A and S. terceris

10. Epimeron 3 lacking posteroventral tooth J. trux

Epimeron 3 bearing posteroventral tooth 11
NUMBER 1 2 4 65

Key to the Species of Syrrhoites—Cont.

11. Tooth of epimeron 3 long and serrate ventrally S. anaticauda

Tooth of epimeron 3 short and smooth 12

12. Rostrum very long and deflexed Syrrhoites species A and S. pusilla

Rostrum of medium length and subhorizontally extended 13

13. Palms of gnathopods each with 2 denning spines 14

Palms of gnathopods each with one defining spine 15

14. Serrations on epimeron 3 extremely large, article 2 of pereopod 5 evenly rounded posteroventrally and heavily serrate, distolateral process on peduncles of uropods 1-2

strong S. columbine

Serrations on epimeron 3 small or absent, article 2 of pereopod 5 truncate posteroventrally, posterior serrations small, distolateral process on peduncles of uropods 1-2

weak S. pantasma

15. Epimeron 2 with straight posterior margin and quadrate posteroventral corner S. cu

Epimeron 2 with sinuous posterior margin and tooth at posteroventral corner 16

16. Epimeron 3 with straight ventral margin and long posterior margin bearing 2 serrations,

article 5 of pereopods 1-2 with few short posterior setae, article 5 of gnathopod 2

scarcely elongate S. silex

Epimeron 3 with beveled ventral margin and very short posterior margin lacking serrations,

article 5 of pereopods 1-2 with numerous long posterior setae and spines, article 5

of gnathopod 2 elongate 17

17. Rostrum subhorizontally extended, telson cleft halfway, dorsal teeth on body small, article

2 of pereopod 5 widely expanded ventrally, uropods 1—2 with distolateral tooth on

peduncle & dulcis

Rostrum strongly deflexed, telson cleft more than halfway, dorsal teeth on body very large,

article 2 of pereopod 5 narrowing ventrally, uropods 1-2 lacking distolateral tooth on

peduncle (see couplets 9 and 12) Syrrhoites species A

Syrrhoites anaticauda K. H. Barnard

Syrrhoites anaticauda K. H. Barnard, 1930: 367-369, figure

37; 1932: 151-152, figure 89.

DIAGNOSIS.—Rostrum not deflexed; spines on

gnathopods unknown, article 5 of gnathopod 2 unknown ("gnathopods like S. serrata") ; article 5 of

pereopods 1—2 unknown; article 2 of pereopod 5

expanded, rounded broadly and shallowly posteroventrally, with small bifid or blunt castellations on

posteroventral margin; uropods 1-2 with distolateral

processes on peduncles; dorsal teeth extending from

pereonite 3 to pleonite 2, pleonite 3 with reverted

tooth, 4 with large hump, 5 with sharp tooth, pereonites 4-7 and pleonites 1-3 with small lateral nodule;

epimeron 1 lacking tooth, epimeron 2 with medium

tooth, epimeron 3 with large tooth serrate ventrally,

posterior margin convex; telson cleft slightly less

than halfway.

Coxae 5-7 with small facial points. See K. H.

Barnard (1932) for strong variant.

Antarctica, McMurdo; South Shetlands; 200-

342 m.

Syrrhoites cohasseta J. L. Barnard

Syrrhoites cohasseta J. L. Barnard, 1967: 173-176, figure 85.

DIAGNOSIS.—Rostrum deflexed moderately to

strongly; gnathopods with one denning spine, article

5 of gnathopod 2 very elongate; article 5 of pereopods

1-2 with long setae; article 2 of pereopod 5 expanded, subtruncate posteroventrally; uropods 1-2

with distolateral process of peduncle but weak on

uropod 1; dorsal teeth extending from pereonite 6

to pleonite 3, latter tooth reverted, dorsoposterior

margin of pleonite 4 sharply quadrate from lateral

view, pleonite 5 with tooth; epimera 1-2 without

tooth, 3 with small tooth, posterior margin nearly

truncate; telson cleft about halfway.

Baja California, 1205-1720 m.

Syrrhoites columbiae, new species

FIGURES 32-34

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with 2 defining spines; article 5 of gnathopod 2 of
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

SI

FIGURE 32.—Syrrhoites columbiae, new species, holotpye, male, 3.7 mm, Oregon 78;

/ = epistome.
NUMBER 1 2 4 67

P3

FIGURE 33.—Syrrhoites columbiae, new species, holotype, male, 3.7 mm, Oregon 78.
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

\vU3

FIGURE 34.—Syrrhoites columbine, new species, holotype, male, 3.7 mm, Oregon 78; / — ramus.

medium elongation; article 5 of pereopods 1-2 poorly

setose (setae sparse and short); article 2 of pereopod

5 expanded, ovate, rounded posteroventrally, with

medium-sized serrations posteroventrally; uropods 1-2

with distolateral process on peduncles; dorsal teeth

extending from pereonite 4 to pleonite 2, pleonites

3-5 with reverted tooth; epimera 1-2 with small

tooth, 3 with large tooth and posterior bulge with

large serrations; telson cleft slightly more than halfway.

DESCRIPTION OF ?YOUNG MALE.—Rostrum of

medium length, turned downwards slightly, lateral

lobes broadly rounded, convex and scarcely truncate

anteriorly (depending on view); body strongly

carinate dorsally, carinae separated into teeth commencing with pereonite 4 and increasing in size to

pleonite 2, tooth of pleonite 3 slightly smaller and

reverted, teeth of pleonites 4-5 small and reverted,

pleonite 6 with very small reverted dorsal tooth;

tooth of pleonite 5 weakly bifid apically in plane from

side to side; tooth of pleonite 4 above insertion of

uropod 1 obsolescent; pleonal epimeron 1 with

broadly rounded posterior margin and small posteroventral tooth, epimeron 2 with sinuous posterior

margin and small posteroventral tooth, epimeron 3

regularly quadratiform, with convex posterior margin

deeply serrate, posteroventral sharp tooth large;

article 1 of antenna 1 with one strong reverted distomedial cusp; gnathopods slender, almost indistinguishable from each other, but gnathopod 2 slightly

the longer, each palm with 2 spines, fifth articles of

gnathopods strongly spinose their full lengths; second

articles of pereopods 3-5 of medium breadth, densely

and deeply serrate, posteroventral corners evenly

rounded; outer rami of uropods 1-2 apically spinose;

article 2 of outer ramus of uropod 3 of medium

length; telson cleft about 60 percent of its length;

outer plate of maxilliped with one strong cusp

produced between 2 spine-teeth; coxa 3 with serrations on distoposterior margin, coxa 4 with 2 serrations forming anteroventral corner, coxae 6 and 7

each with small posterior cusps.

HOLOTYPE.—USNM 127134, male, 3.7 mm.

Unique.

TYPE-LOCALITY.—Oregon 78, off mouth of Columbia River, Oregon, 44°40.1'N, 125°06.7'W, 800 m,

18 June 1964.

RELATIONSHIP.—This species has close affinities

with Syrrhoites serrata (Sars) (see Sars 1895, pi.

137). The 2 species are grossly similar in dorsal

ornamentation except that S. serrata has double

teeth on some pleonites, the bidentation occurring
NUMBER 12 4 69

axially. Syrrhoites columbiae has a weakly bidentate

tooth of pleonite 5 that occurs in the lateral plane

and which presumably does not occur on S. serrata.

The serrations of coxae 3—4 and the cusps of coxae

6-7 characterize S. columbiae. The outer plate of the

maxilliped has a fixed cusp not occurring in

S. serrata.

Oregon, 800 m.

Syrrhoites cu, new species

FIGURES 35-37

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2

elongate; article 5 of pereopods 1-2 with about 3

long posterior setae; article 2 of pereopod 5 expanded,

ovate, rounded posteroventrally, serrations small;

uropods 1-2 with distolateral process on peduncles;

dorsal teeth extending from pereonites 5-6 to pleonite

4, tooth on pleonite 5 large, female with weak dorsal

serrations on pleonite 3; epimera 1-2 lacking teeth

or softly quadrate posteroventrally, epimeron 3 with

weak tooth in female, quadrate in male, posterior

margin straight, very short, posteriorly serrate in female, epimeron beveled below main tooth; telson

cleft slightly more than halfway.

DESCRIPTION OF FEMALE.—Rostrum of medium

length, turned downwards slightly, lateral cephalic

lobes broadly adz-shaped and convex anteriorly, head

short, with ventral notch strongly anteriad; body

weakly carinate dorsally, posterodorsal teeth commencing on pereonite 5 and increasing in size to pleonite 3, latter tooth formed as squared-off, serrate

carina, tooth of pleonite 4 weak, that of pleonite 5

long; pleonite 4 with weak posterolateral tooth above

insertion of uropod 1; pleonal epimeron 1 broadly

subquadrate posteroventrally, epimeron 2 with

straight posterior margin and quadrate posteroventral corner, epimeron 3 of medium depth, posterior

margin straight and minutely serrate, posteroventral

corner with small tooth; article 1 of antenna 1 with

one strong distal cusp; gnathopods with one major

palmar spine, palms essentially indistinct, article 5 of

gnathopod 2 strongly setose and not abnormally

elongate; article 2 of pereopods 3-5 of medium

breadth, posteroventral margins sloping and not distinctly quadrate or produced; outer ramus of uropod

1 with distal spines, of uropod 2 with subdistal constriction and small spine; article 2 of outer ramus

of uropod 3 not distinguished clearly; telson cleft

about 60 percent of its length.

Maxillae 1-2 are like those figured herein for

Syrrhoites pantasma, new species. The mandibular

palp is also like that of S. pantasma except the midmarginal seta (of 5) on article 2 is large and elongate.

MALE.—Differing from female in stronger ventral

extension of pleonal epimeron 3 with relatively

shorter posterior margin, in dorsad thickening of

pleonites 4-5, thinner tooth of pleonite 5 and

stronger and more erect dorsal hump of pleonite 6;

article 2 of pereopods 3-5 relatively narrower than

in female; article 3 of antenna 1 shorter, article 5 of

antenna 2 longer, and bases of flagella on antenna 1

conjoint; short article 2 on outer ramus of uropod

3 more clearly distinct in male than in female;

apices of telson shown to be bifid because of better

preservation in male; rostrum slightly sharper than

in female; coxa 6 with rounded, not subquadrate

posterior lobe.

HOLOTYPE.—AMNH 14,395, female, 3.2 mm.

TYPE-LOCALITY.—Vema 15-38, off Pacific Colombia, 05°00/N, 79°04'W, 3023-3251 m, 15 November

1958.

MATERIAL.—Male and female from the typelocality.

RELATIONSHIP.—This species resembles Syrrhoites

walkeri Bonnier (1896) in most of its major qualitative characters but differs by several quantitative

characters and a few of qualitative significance. The

dorsal teeth are generally much larger in S. walkeri

than in S. cu, except for the tooth of pleonite 5.

Pleonal epimeron 3 of S. walkeri lacks posterior

serrations (a possible sexual characteristic) but has

a larger epimeral tooth than does S. cu; the posterior

lobe of coxa 4 is sharper in S. walkeri and the major

spines on the outer plate of the maxilliped smaller

in S. walkeri than in S. cu.

Syrrhoites tenellus K. H. Barnard (1925) lacks a

tooth and serrations on pleonal epimeron 3 and apparently has distinct palms on gnathopods 1-2.

There is a strong resemblance between S. cu and

S. trux J. L. Barnard (1967) but S. trux has a

distinct but minute tooth on pleonal epimeron 1, a

sinuous posterior margin of epimeron 2, a vertically

shortened epimeron 3 with the shape of a hatchet

and coxa 4 has sharp corners.

Syrrhoites cohasseta J. L. Barnard (1967) differs
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 35.—Syrrhoites cu, new species, holotype, female, 3.2 mm, Vema 15-38;

y=male, 3.5 mm; /—lower lip.
NUMBER 1 2 4 71

Dl

FIGURE 36.—Syrrhoites cu, new species, holotype, female, 3.2 mm, Vema 15-38;

y=male, 3.5 mm; /?=ramus.
72

SMITHSONIAN CONTRIBUTION TO ZOOLOGY

FIGURE 37.—Syrrhoites cu, new species, holotype, female, 3.2 mm, Vema 15-38; y=male, 3.5 mm.

from S. cu in the absence of distal spines on the

outer ramus of uropod, 1, the very elongate article

5 of gnathopod 2 lacking posterior setae, the quadrate posteroventral corner of article 2 on pereopod 5

in the female.

Syrrhoites silex J. L. Barnard (1967) is very similar to the species at hand but differs primarily in the

presence of a large posteroventral sinuosity and tooth

on pleonal epimeron 2; S. silex also has distinctly

articulate distal spines on the outer ramus of uropod

2 and a narrow article 5 of pereopod 5 with subconically produced posteroventral corner.

Pacific Colombia, 3023-3251 m.

Syrrhoites dulcis J. L. Barnard

Syrrhoites dulcis J. L. Barnard, 1967: 176-178, figure 86.

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2

elongate; article 5 of pereopods 1-2 with long spinesetae ; article 2 of pereopod 5 expanded and ventrally

truncate; uropods 1-2 with distolateral process on

peduncles; dorsal teeth extending from pereonite 3

to pleonite 5, tooth of pleonite 3 slightly reverted;

epimeron 1 lacking tooth, epimera 2-3 with small

tooth, posterior margin of epimeron 3 short, nearly

straight, epimeron below tooth beveled; telson cleft

halfway.

Baja California, 1095-1205 m.

Syrrhoites lorida (J. L. Barnard)

Kindia lorida J. L. Barnard, 1962a: 57, figure 49.

DIAGNOSIS.—Rostrum deflexed; gnathopods with

one defining spine; following parts unknown: gnathopod 2, pereopods 1-2, pereopod 5, uropods 1-2;

dorsal teeth extending from pereonite 2 to pleonite 5

except tooth on pleonite 4 damaged and possibly

very small; epimera 1-2 lacking tooth, 3 with medium

sharp tooth and convex posterior margin; telson cleft

about five-eighths its length.

South Africa, 3045 m.
NUMBE R 12 4 73

Syrrhoites pacifica Nagata

Syrrhoites paciftcus Nagata, 1965: 171-174, figure 16.

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2

elongate; article 5 of pereopods 1-2 with long setae;

article 2 of pereopod 5 expanded and posteroventrally

rounded; uropods 1-2 with distolateral process on

peduncles, that on uropod 2 weak; dorsal teeth in

female extending from pereonite 2 to pleonite 3,

obsolescent on pleonites 4-5, in male from pereonite

7 to pleonite 5 but weak on pleonite 4, pleonite 3

in both sexes with tooth forming crest and on posterior segments generally dorsal configuration saddleshaped, thus with hump in middle of segment dorsally; epimera 1-2 unknown, 3 with sharply extended

acute posteroventral corner, and concave short posterior margin; telson cleft halfway.

Japan, 20-100 m.

Syrrhoites pantasma, new species

FIGURES 38, 39

DIAGNOSIS.—Rostrum especially small and moderately deflexed; gnathopods with 2 defining spines,

article 5 of gnathopod 2 elongate; article 5 of pereopods 1-2 with posterior setae short to medium in

length; article 2 of pereopod 5 widely expanded

and ventrally truncate; uropods 1-2 with weak distolateral process on peduncles; dorsal teeth extending from pereonite 5 to pleonite 5; tooth on pleonite

3 slightly reverted, tooth on pleonites 4-5 of medium

size; epimeron 1 lacking tooth, epimeron 2 with

medium tooth, 3 with small tooth, posterior margin

weak to strongly convex, serrate in male; telson cleft

about five-eighths its length.

DESCRIPTION OF FEMALE.—Rostrum of medium

length, lateral cephalic lobes narrowly truncate anteriorly; body strongly carinate dorsally, pereonites

5-7 and pleonites 1-6 all with strong posterodorsal

teeth, teeth anterior to urosome largest, tooth of

pleonite 3 slightly reverted; pleonite 4 with strong

subacute lateral tooth above insertion of uropod 1;

pleonal epimeron 1 broadly rounded posteroventrally,

epimeron 2 with posterior margin swept into mediumsized tooth, epimeron 3 deep, with strongly convex

posterior margin and minute posteroventral tooth;

article 1 of antenna 1 with one strong distolateral

cusp, a small distomedial cusp; article 2 of pereopods

3-5 very broad, articles 4-5 slender, highly elongate

in comparison to Syrrhoites sorpresa (J. L. Barnard);

outer rami of uropods 1-2 distally spinose; outer ramus

of uropod 3 with article 2 almost 30 percent as long

as article 1, telson cleft slightly more than halfway.

?MALE, 3.0 mm: Bearing spined penial processes

on seventh pereonal sternite; differing from female

in larger tooth of pleonal epimeron 3 with small serrations on posterior margin above tooth; all other

parts figured and described for female similar, including antenna 1, but dorsal ornamentation of

pleonites 5 and 6 damaged and not analyzed; head

illustrated without need for restoration.

HOLOTYPE— AMNH 14,399, female, 4.0 mm.

TYPE-LOCALITY.—Vema 15-62, off Ecuador, 01°30'

S, 82°19/W, 1363-1369 m, 3 December 1958.

MATERIAL.—Vema 15-59 (1 male), 15-62 (holotype).

RELATIONSHIP.—This species has approximately

the same dorsal configuration as Syrrhoites sorpresa

(J. L. Barnard, 1962a, as Kindia sorpresa) from Cape

Basin, South Africa. The unique specimen of that

species was badly damaged and several characters are

unknown except for a specimen reported herein

believed to be identical with S. sorpresa: the condition

of the uropods, the shape of pleonal epimeron 2, the

dorsal margins of urosomites 2 and 3. Like the specimen at hand and so many deep-sea synopiid specimens the rostrum is forced upward and damaged.

The rostrum is restored in the drawings of S. pantasma, new species. Nevertheless, the specimen at

hand is in good condition and differs from S. sorpresa in the following characters: (1) the deeper

pleonal epimeron 3, with long, convex posterior

margin and small, nearly obsolete posteroventral

tooth (in S. sorpresa the posterior margin is short,

concave, and the tooth is large and dominates the

epimeron) ; (2) the presence of 2 palmar defining

spines on gnathopods 1 and 2 (1 spine in S. sorpresa) ;

(3) the much longer articles 5 and 6 of pereopods

3 and 4.

If the specimen of S. sorpresa reported herein is

correctly identified then S. pantasma differs from S.

sorpresa in the presence of a tooth on pleonal

epimeron 2, a vestigial cusp on the peduncle of

uropod 1, a cusp on coxa 3 and a strongly setose

gnathopod 2 among other more minor characters.
74 SMITHSONIAN CONTRIBUTION TO ZOOLOGY

FIGURE 38.—Syrrhoites pantasma, new species, holotype, female, 4.0 mm, Vema 15-62.
NUMBER 12 4 75

oP3

FIGURE 39.—Syrrhoites pantasma, new species, holotype, female, 4.0 mm, Vema 15—62 3"

y=male, 3.0 mm, Vema 15-59.
76 SMITHSONIAN CONTRIBUTION TO ZOOLOGY

Syrrhoites pantasma has a character which is

unique, so far as is known, among members of

Syrrhoites. The anteroventral corner of coxa 3 has

a small cusp, typical of many members of Austrosyrrhoe, Bruzelia and Latacunga.

Syrrhoites pantasma resembles S. silex J. L. Barnard

(1967) very strongly but differs in the presence of a

cusp on coxa 3, a very enlarged article 2 of pereopod

5, a vestigial cusp on peduncle of uropod 1 and a

relatively elongate inner ramus of uropod 1 among

other more minor characters.

Off Pacific Panama and Ecuador, 1369-2853 m.

Syrrhoites pusilla Enequist

Syrrhoites pusillus Enequist, 1950: 338-340, figures 57-60.

DIAGNOSIS.—Rostrum deflexed; gnathopods with

one defining spine, article 5 of gnathopod 2 elongate;

article 5 of pereopods 1-2 with long spines; article

2 of pereopod 5 expanded and posteroventrally

rounded; uropods 1-2 with weak distolateral process

on peduncles; dorsal teeth extending from pereonite

5 to pleonite 5 but tooth on pereonite 4 obsolescent,

large on pereonite 5, reverted on pereonite 3;

epimeron 1 lacking tooth, epimeron 2 with small

tooth, 3 with small, sharp upturned tooth, posterior

margin short, slightly concave; telson cleft two-thirds

its length.

Skagerrak.

Syrrhoites redox J. L. Barnard

Syrrhoites redox J. L. Barnard, 1967: 178-183, figures 87,

88.

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2

elongate; article 5 of pereopods 1-2 lacking long

spines; article 2 of pereopod 5 expanded and ventrally

truncate; uropods 1-2 with no peduncular processes;

small dorsal teeth extending from pereonite 5 to

pleonite 2, tooth of pleonites 3 and 5 large, tooth

of pleonite 4 obsolescent; epimeron 1 lacking tooth,

epimeron 2 with small tooth, epimeron 3 with sharp

posteroventral corner, posterior margin short and

convex; telson cleft about one-third its length.

Baja California, 1720-1748 m.

Syrrhoites s err at a (Sars)

Bruzelia serrata Sars, 1879: 447.

Syrrhoites serratus.—S&n, 1895: 392-394, pi. 137.—Gurjanova, 1951: 593, figure 398.

DIAGNOSIS.—Rostrum slightly deflexed; gnathopods with 2 defining spines, article 5 of gnathopod 2

not elongate; article 5 of pereopods 1-2 with short

setae; article 2 of pereopod 5 expanded and posteroventrally rounded; uropod 1 with strong peduncular

process, uropod 2 with weak process; dorsal teeth

extending from pereonite 5 to pleonite 5, teeth

paired in tandem on each segment producing dorsal

saddle crest especially on pleonites 1-4; epimeron 1

lacking tooth, epimeron 2 with small tooth, epimeron

3 with medium sharp tooth, posterior margin convex

and heavily serrate; telson cleft about two-thirds its

length.

Arctic boreal of North Atlantic, bathyal.

Syrrhoites silex J. L. Barnard

Syrrhoites silex J. L. Barnard, 1967: 183-185, figure 89.

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2

slightly elongate; article 5 of pereopods 1-2 with

3 setae of medium length; article 2 of pereopod

5 slightly expanded and ventrally truncate; uropods

1-2 with distolateral process on peduncles; dorsal

teeth extending from pereonite 5 to pleonite 5, tooth

on pleonite 3 slightly reverted; epimeron 1 lacking

tooth, epimera 2 and 3 with small tooth, epimeron

3 with 2 serrations above main tooth; telson cleft

two-thirds its length.

Baja California, 1095-1205 m.

Syrrhoites sorpresa (J. L. Barnard)

FIGURES 40, 41

Kindia sorpresa J. L. Barnard, 1962a: 57, figure 48.

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2 elongate; article 5 of pereopods 1-2 with long posterior

setae; article 2 of pereopod 5 expanded and truncate

ventrally; uropods 1-2 probably with peduncular

processes; dorsal teeth extending from pereonite 5 to

pleonite 3, tooth small on pleonite 4, probably also
NUMBE R 12 4 77

U2

FIGURE 40.—Syrrhoites sorpresa (J. L. Barnard), male, 3.8 mm, Eltanin 257.
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 41.—Syrrhoites sorpresa (J. L. Barnard), male, 3.8 mm, Eltanin 257.

present on pleonite 5; epimera 1-2 lacking tooth,

epimeron 3 with small to medium tooth, posterior

margin straight; telson cleft about two-thirds or

slightly less.

MATERIAL.—Eltanin 257 (?male, 3.8 mm.).

REMARKS.—The individual in this collection is in

much better condition than was the holotype (the

unique representative of the species at the time of its

erection). The present identification is based on congruence in the dorsal teeth of pereonites and pleonites

1-4, pleonal epimera, coxae, head, gnathopods and

pereopods. The holotype had damaged pleonites 5-6,

pleonal epimeron 2, head, and missing or damaged

uropods 1-3. The following notes and accompanying

illustrations supplement the original description and

figures:

Pleonite 5 with small dorsal tooth, pleonite 6

slightly elevated posterodorsally but untoothed (tooth

of pleonite 4 dorsally damaged) ; pleonal epimeron

2 with evenly and slightly convex posterior margin

and smoothly rounded posteroventral corner (tooth

of epimeron 3 damaged); posterolateral, supra-
NUMBER 12 4 79

uropodal tooth of pleonite 4 very long and sharp;

gnathopods essentially simple (thus resulting in

description of now synonymous genus Kindia), each

definitely bearing only one major posterior spine;

coxa 3 definitely with rounded anteroventral corner;

pereopods 1-2 with posterior setae but no comb on

article 6; serrations on posterior margin of article

2 on pereopod 5 extending full length; peduncle of

uropod 1 with long distolateral process, outer ramus

slightly more than half as long as inner, apices of

both rami and outer ramus of uropod 1 with large

distal spine and subsidiary spines; outer ramus of

uropod 2 slightly more than half as long as inner

ramus, latter narrow, lanceolate, distally simple,

peduncle with small distolateral process; outer ramus

of uropod 3 biarticulate, article 2 more than twothirds as long as article 1; left and right mandibles

similar but right molar bearing marginal spine,

absent on left molar. Other details of mouthparts

and appendages shown in figures.

There may be subspecific differences in the two

individuals of this species now known, but if so they

must be confined to pleonites 5-6 and the uropods

or on article 1 of antenna 1 (which perhaps was

inadequately observed in the holotype).

Maxillae are not illustrated herein, except for the

apex of the palp on maxilla 1; maxillae otherwise

resemble those illustrated for Syrrhoites pantasma,

new species.

RECORD.—Drake Passage, 3867^1086 m.

Drake Passage and Cape Basin, South Africa,

1861-3867 m.

Syrrhoites tenella K. H. Barnard

Syrrhoites tenellus K. H. Barnard, 1925: 353-354.

DIAGNOSIS.—Rostrum slightly deflexed; gnathopods

"like serratus" thus gnathopods probably with one

defining spine, article 5 of gnathopod 2 not elongate;

pereopods 1-2 "like walkeri" thus probably lacking

long spines on article 5; pereopod 5 "like walkeri"

thus article 2 probably expanded and posteroventrally

rounded; uropods 1-2 "like serratus" thus probably

with processes on peduncles but weak on uropod 2;

dorsal carina extending from pereonites 6 or 7 to

pleonite 4, not extended as teeth, posterodorsal

corners of segments extended quadrately; epimeron

1 lacking tooth, epimera 2-3 quadrate and with slight

point at posteroventral corner; telson cleft halfway.

South Africa, 700 fms.

Syrrhoites terceris J. L. Barnard

Syrrhoites terceris J. L. Barnard, 1964: 31-32, figure 25.

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2

elongate; article 5 of pereopods 1-2 with setae of

medium length; article 2 of pereopod 5 expanded,

ventrally subtruncate; uropods 1-2 lacking peduncular processes; dorsal teeth extending from pereonite

5 to pleonite 2, teeth obsolescent on pleonites 3-4

and small on pleonite 5; epimeron 1 lacking tooth,

epimeron 2 with medium tooth, epimeron 3 with

sharply quadrate posteroventral corner, posterior

margin finely serrate; telson cleft two-thirds its

length.

East tropical Pacific, 1609-1746 m.

Syrrhoites trux J. L. Barnard

Syrrhoites trux J. L. Barnard, 1967: 185-188, figures 90, 91.

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2

scarcely elongate; article 5 of pereopods 1-2 with

numerous short setae; article 2 of pereopod 5 expanded and broadly rounded posteroventrally; uropods 1-2 with distolateral process on peduncles;

pereonite 7 with weak tooth, pleonites 1-2 with

quadrate posterodorsal elevations, pleonite 3 with

sharp tooth, pleonites 4-5 with small tooth erect on

pleonite 4; epimeron 1 with small tooth, epimera

2-3 with quadrate posteroventral corner, epimeron 3

with short and straight posterior margin; telson cleft

halfway.

Baja California, 842-1095 m.

Syrrhoites walkeri Bonnier

Syrrhoites Walkeri Bonnier, 1896: 647-650, pi. 38, figure 4.

DIAGNOSIS.—Rostrum not deflexed; gnathopods

with one defining spine, article 5 of gnathopod 2

elongate; article 5 of pereopods 1-2 lacking long setae;

article 2 of pereopod 5 expanded and posteroventrally

rounded; uropod 1 with peducular process, uropod

2 apparently without; dorsal teeth extending from

pereonite 6 to pleonite 5; epimera 1-2 lacking tooth,

epimeron 3 extended as sharp posteroventral protu-
80 SMITHSONIAN CONTRIBUTION TO ZOOLOGY

sion, posterior margin slightly concave; telson cleft

halfway.

Northeast Atlantic, 950 m.

Syrrhoites species A

FIGURES 42, 43

DIAGNOSIS.—Rostrum moderately deflexed; gnathopods with one defining spine, article 5 of gnathopod

2 elongate; article 5 of pereopods 1-2 with long

spines; article 2 of pereopod 5 narrowly expanded

and subtruncate posteroventrally; uropod 2 lacking

peduncular process, uropod 1 unknown; dorsal teeth

extending from pereonite 5 to pleonite 3, tooth absent

on pleonite 4, small (female) to medium (male)

tooth on pleonite 5; epimeron 1 lacking tooth, subquadrate posteroventrally, epimera 2-3 with medium

tooth, posterior margin of epimeron 3 straight; telson

cleft two-thirds its length.

MALE WITH SPECIAL CHARACTER.—Sternal teeth

present on pereonites 4-6.

MATERIAL.—Eltanin 75 (2).

REMARKS.—These specimens, a male and a female,

though obviously distinct from any species heretofore described, are not named because they are

moderately damaged; they lack uropod 1, the distal

spines on the outer ramus of uropod 2 and pereopods

3-5 are badly damaged in the male so that the

posterior margins of the second articles are unclear;

the setae and spines of many of the appendages,

especially antennae and pereopods 3-5, are worn off

as if they had been sandblasted. The most remarkable aspect of the male is the presence of 3 large

sternal hooks on the pereon. Unfortunately this discovery came late in the current study and although

previously analyzed specimens of other species were

reexamined in search for these hooks, several specimens which might have had them were so damaged

by dissection and analysis that they were not clearly

useful. The sternites of the pereon are extremely weak

and fragile in many deep-sea synopiids. These hooks

may be an excellent specific character of some males

of the group and should be noted when possible.

DESCRIPTION.—Rostrum very long, lateral cephalic

lobes weakly rounded anteriorly; body strongly

carinate, dorsal teeth occurring on pereonites 5-7,

pleonites 1-3 and pleonite 4, teeth of pereonites 5-7

and pleonites 1-3 largest, tooth of pleonite 3 slightly

reverted; pleonite 4 with strong subacute tooth

(broken in male) above insertion of uropod 1;

pleonal epimeron 1 broadly rounded posteroventrally,

epimeron 2 with posterior margin swept into mediumsized tooth (presumed in male, see relationship),

epimeron 3 of medium depth, with straight posterior

margin and weak blunt (male) or sharp (female)

posteroventral tooth ("sandblasted") ; article 2 of

pereopods 4-5 moderately broad, of pereopod 3

apparently moderately slender, articles 4 and 5 not

especially elongate (where known, see figures);

outer ramus of uropod 2 with sockets indicating

presence of distal spine(s) ; outer ramus of uropod 3

with very elongate article 2, about equal in length to

article 1; telson cleft about two-thirds of its length;

male and female differing only in presence of sternal

hooks in male and brood lamellae in female.

Mouthparts like those of S. sorpresa figured herein

except as follows; both mandibles with spine on

molar, right lacinia mobilis similar and simple, left

lacinia mobilis with weakly developed bifid or trifid

fused process adjacent to its base; lower lip not

analyzed owing to damage; first maxillary palp with

similar positions of distal setae; maxilliped similar

but terminal spine of outer plate much more slender

and inner plate with 2 medioventral coupling spines

instead of one.

RELATIONSHIP.—This species has a remarkable

resemblance to S. sorpresa (J. L. Barnard, 1962a, and

see herein) but pleonal epimeron 2 (though damaged

in male) has a distinct, medium-sized posteroventral

tooth; the apex of the tooth is broken but the internal

lines of the succeeding instar demonstrate its form

in the male. It is well developed in the female. The

lateral peduncular cusp of Syrrhoites species A is not

as strong on uropod 2 as it is in S. sorpresa, coxa 4

is more elongate and coxae 5 and 6 have strongly

truncate posteroventral lobes.

Apparently Syrrhoites species A differs from the

following species in those characters stated: from

S. anaticauda K. H. Barnard in the presence of only

one strong spine on the gnathopodal palms, in the

absence of serrations on pleonal epimeron 3 (possibly

a sexually dimorphic character) and the absence of

dorsal teeth on pereonites 3 and 4; from S. redox

J. L. Barnard in the larger segmental teeth, the much

deeper cleft of the telson, the poorly setose gnathopod

2; from its sympatriot S. pantasma, new species, in

the presence of only one spine on the gnathopodal
NUMBER 12 4 81

FIGURE 42.—Syrrhoites species A, female, 4.5 mm, Eltanin 75; y=male, 4.0 mm.
82

SMITHSONIAN CONTRIBUTION TO ZOOLOGY

yG2

FIGURE 43.—Syrrhoites species A, female, 4.5 mm, Eltanin 75; y=male, 4.0 mm.

palms, the absence of a cusp on coxa 3, the absence

of a tooth on pleonite 4 (perhaps a sexual difference),

the weak setation of gnathopod 2, the elongate article

2 on the outer ramus of uropod 3, the more slender

lacinia mobili of the mandibles, and the much longer

rostrum.

Species A is similar to S. terceris J. L. Barnard

in the deeply cleft telson and the absence of processes

on the peduncles of uropods 1-2 but it differs from

S. terceris in the much larger dorsal teeth on body

segments, the posteriorly shortened epimeron 3 with

only one tooth posteroventrally, and the ventrally

narrowed article 2 of pereopod 5.

Species A is also similar to S. pusilla Enequist in

the deflexed rostrum, the configuration of dorsal

teeth, telson and long setae on pereopods 1-2 but

differs from S. pusilla by the larger tooth on pleonite

3 and the absence of peduncular processes on uropods

1-2.

Species A resembles S. dulcis J. L. Barnard in the

long spines on article 5 of pereopods 1-2 and in the

elongate article 5 of gnathopod 2 but differs from
NUMBER 12 4 83

S. dulcis in the long downturned rostrum, the

narrower and more deeply cleft telson, the much

larger dorsal teeth, the ventrally narrowing article 2

of pereopod 5, the absence of peduncular teeth on

uropods 1—2 and the absence of a dorsal tooth on

pleonite 4.

Chile, 1834 m.

Tiron Liljeborg

Tiron Liljeborg, 1865: 19.—Stebbing, 1906: 275-276.

Tessarops Norman, 1868: 412 [homonym, Arachnida].

DIAGNOSIS.—Forehead protuberant or not (typespecies), lateral cephalic lobe sharp; eyes present

and accessory eyes usually present; mandible with

palp, molar of medium size, strongly projecting,

columnar and triturative; mouthparts basic; articles

1-2 of antenna 1 basic; coxa 1 ordinary, coxae 3-4

not pelagont or weakly so, coxa 3 softly rectangular,

posterior margin almost parallel with anterior margin

and not strongly excavate; coxa 4 variable, typically

adz-shaped and almost as long as coxa 3, surface

area of latter nearly equal to coxa 4, rarely coxa 4

distinctly shorter and smaller than 3 and weakly

comma-shaped; gnathopods simple, hands elongate,

linear, lacking distinct spines; dactyl of gnathopod 2

normal; pereopods 3-5 very short, dactyls typically

very short, clawlike and bearing large inner wire-seta,

but occasionally dactyls slightly elongate and poorly

armed; article 2 of pereopod 5 posteroventrally

rounded; pleonites 1-3 typically denticulate dorsally

but apparently smooth in 3 species; uropod 3 greatly

exceeding apices of uropods 1-2, (except in T.

brevidactylus), peduncle short; telson elongate, deeply

cleft.

Uropod 1 reaching apex of uropod 2 (see

Pseudotiron).

TYPE-SPECIES.—Lysianassa spinifera Stimpson

(1853) ( = 7\ acanthurus Liljeborg, 1865, see Sars,

1895).

REMARKS.—Apparently all species of Tiron have

one or more small teeth on the dorsoposterior margin

of pleonites 1-5, the teeth usually becoming enlarged

on pleonites 4-5; on pleonites 1—3 the middle tooth

is often very small and indistinguishable from a

group of serrations running transversely across the

segmental margin; these extra serrations are mentioned in the diagnoses of species, but little account

is yet taken of the size of dorsal teeth generally until

more is known of growth and sexual variables. Males

usually have larger teeth than do females.

Accessory eyes have not been seen in all species.

In the following key T. antarcticus is omitted because several of its attributes are unknown and it

otherwise is similar to T. biocellata; see the latter for

minor distinctions known so far.

Dactyls of pereopods 1-5 are classed in 2 alternatives, either normally claw-shaped or stubby. The

condition of stubby dactyls may be seen herein on

T. tropakis; the dactyls are short, tumid and may

bear one or more accessory processes or spines. Even

if the spine extends as a point the dactyl is still regarded as stubby.

Tiron bombayensis Tembe-Deshpende (1964) is a

nomen nudum.

Key to the Species of Tiron

(T. antarcticus omitted, see text)

1. Each lobe of telson with medial row of one or more large spines (all of pereopods 1-5

with stubby dactyls) 2

Each lobe of telson lacking large medial spines in rows 5

2. Outer plate of maxilliped aberrant, apically incised and guarded by hooked wing on each

side, large spines absent T. thompsoni

Outer plate of maxilliped normal, ovate, bearing large spines on medial margin 3

3. Article 2 of pereopods 4-5 with posterior submarginal row of long plumose setae T. intermedia

Article 2 of pereopods 4—5 lacking special row of setae 4

4. Accessory eye composed of 4 appressed ommatidia, mandibular palp present T. australis

Accessory eye composed of 2 segregated ommatidia, mandibular palp absent T. tropakis

5. Dactyls of pereopods 1-5 stubby 6

Dactyls of pereopods 1-5 ordinary 7
84

SMITHSONIAN CONTRIBUTION TO ZOOLOGY

Key to the Species of Tiron—Cont.

6. Outer ramus of uropod 2 shortened, rami of uropod 3 obliquely truncate apically

T. brevidactylus

Outer ramus of uropod 2 as long as inner, rami of uropod 3 pointed apically T. altifrons

7. Posterior margin on article 2 of pereopod 5 smooth, accessory eye with 3 appressed

ommatidia T. spiniferum

Posterior margin on article 2 of pereopod 5 crenulate and setulose, accessory eye with 2

segregated ommatidia T. biocellata

Tiron altifrons Reid

Tiron altifrons Reid, 1951: 235-236, figure 31.

DIAGNOSIS.—Accessory eye apparently composed of

about 5 appressed ommatidia; mouthparts unknown;

dactyls of pereopods 1-5 stubby but sharp; article 2

of pereopods 4-5 lacking special setal row, article 2

of pereopod 5 with 3 setulose crenulations posteriorly;

pleonites 1-3 unknown; outer rami of uropods 1-2

not shortened; rami of uropod 3 apically pointed;

each lobe of telson with 3 groups of dorsal bristles.

West Africa.

Tiron antarcticus K. H. Barnard

Tiron antarcticus K. H. Barnard, 1932: 148-149, figure 86.

DIAGNOSIS.—Accessory eyes not seen; mouthparts

not fully described; dactyls of pereopods 1-5 ordinary; article 2 of pereopods 4-5 without special row

of setae, article 2 of pereopod 5 posteriorly crenulate

and setulose; pleonites 1-3 not dorsally crenulate;

rami of uropods not described; each lobe of telson with

4-5 dorsal setules.

South Shetlands and South Georgia, 135-200 m.

Tiron australis Stebbing

Tiron australis Stebbing, 1908: 79-81, pi. 38.

DIAGNOSIS.—Accessory eye with 4 appressed ommatidia; inner plate of maxilla 1 fully setose but with

smooth acclivital portion in middle; inner plate of

maxilla 2 with medial submarginal row of setae; outer

plate of maxilliped normal; dactyls of pereopods 1-5

stubby but apically sharp; article 2 of pereopods 4-5

without special row of setae, article 2 of pereopod 5

weakly and irregularly crenulate posteriorly but not

setulose; pleonites 1-3 dorsally crenulate; outer ramus

of uropod 1 not shortened, of uropod 2 shortened

slightly; rami of uropod 3 apically pointed; each lobe

of telson with medial row of large spines.

South Africa, 47 fms.

Tiron biocellata J. L. Barnard

FIGURE 44

Tiron biocellata J. L. Barnard, 1962b: 75, figure 2.

DIAGNOSIS.—Accessory eye composed of 2 segregated ommatidia; inner plate of maxilla 1 fully setose;

inner plate of maxilla 2 with medial row of submarginal setae; outer plate of maxilliped normal;

dactyls of pereopod 5 ordinary; article 2 of pereopods

4-5 without special row of setae, article 2 of pereopod

5 crenulate and setulose posteriorly; pleonites 1-3

weakly crenulate dorsally; outer rami of uropods 1-2

significantly shortened; rami of uropod 3 apically

pointed; telson with weak dorsal setae.

Distinguished from T. antarcticus K. H. Barnard

also in the narrow and more elongate article 4 of

pereopod 5.

REMARKS.—Mouthparts of this species and T.

tropakis, new species, have many similarities, especially

the maxillipeds and maxilla 2 which are similar in

all essentials, but the outer plate of maxilla 1 in T.

biocellata has 10 spines, that of T. tropakis 7 spines,

the inner plate of T. biocellata has only a small gap

between the terminal and medial setae whereas a

large gap occurs in T. tropakis. The latter has no

mandibular palp. The telson of T. biocellata has

almost no dorsal spines, occasionally one subterminal

spine occurring on each lobe.

MATERIAL.—Velero 4718 (1), 4719 (1), 4721 (4),

4759 (1), 4787 (8), 4822 (1), 4836 (2), 4840 (1),

4842 (1), 4843 (1), 4861 (2), 4869 (3), 4870 (2),

4871 (1), 4885 (5), 4886 (2), 5042 (1), 5109 (1),

5134 (6), 5167 (1), 5172 (4), 5256 (2), 5346 (1),

5373 (1), 5507 (1), 5557 (1), 5564 (4), 5631 (2),
NUMBER 12 4 85

XI \ X2

FIGURE 44.—Tiron biocellata J. L. Barnard, female, 4.6 mm, Velero 5631; e=lacinia mobilis.

565 4 (1) , 570 9 (2) , 574 0 (1) , 574 1 (1) , 576 5 (1) ,

583 8 (1) , 596 5 (2) , 596 8 (1) , 5972 (1) , 597 3 (1) ,

610 0 (1) , 610 2 (2) , 610 3 (1) , 639 3 (1) , 669 4 (1 )

(from th e Alla n Hancoc k Foundation) .

Californi a an d Baj a California , 9-18 0 m .

Tiron brevidactylus (Pillai)

Pseudotiron brevidactylus Pillai, 1957: 43-46, figure 7.

DIAGNOSIS.—Accessory eyes not described; inner

plate of maxilla 1 small and naked; inner plate of

maxilla 2 lacking medial setae; outer plate of maxilliped normal; dactyls of pereopods 1-5 stubby; article

2 of pereopods 4-5 lacking special setal row, article

2 of pereopod 5 crenulate and setulose posteriorly;

serrations on pleonites 1-3 not mentioned; outer

ramus of uropod 1 as long as inner, but shortened on

uropod 2; rami of uropod 3 with oblique truncate

apices; telson with several scattered dorsal setae.

Mandibular palp absent (special character, see

T. tropakis, new species).

Travancore.

Tiron intermedia Reid

Tiron intermedia Reid, 1951: 233-235, figure 30.

DIAGNOSIS.—Accessory eyes present but composition unknown; mouthparts unknown; dactyls of
86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

percopods 1-5 stubby; article 2 of pereopods 4-5

with posterior submarginal row of long plumose setae,

article 2 of pereopod 5 with posterodorsal ala bearing

3 crenulations, no setules; pleonites 1-3 dorsally

crenulate; rami of uropods 1-2 unknown; rami of

uropod 3 apically pointed; each lobe of telson with

row of Pmedial spines.

West Africa.

Tiron spiniferum (Stimpson)

Lysianassa spinifera Stimpson, 1853: 49.

Tiron acanthurus Liljeborg, 1865: 19.—San, 1895: 399-

401, pi. 140.—Shoemaker, 1930: 291-292.—Stephensen,

1938: 231; 1944: 76.—Gurjanova, 1951: 591, figure 397.

Tiron spiniferum Shoemaker, 1955: 38-39.

DIAGNOSIS.—Accessory eye composed of 3 appressed

ommatidia; inner plate of maxilla 1 fully setose;

inner plate of maxilla 2 with medial submarginal

row of setae; outer plate of maxilliped normal;

dactyls of pereopods 1-5 ordinary; article 2 of pereopods 4-5 lacking special row of setae, article 2 of

pereopod 5 smooth posteriorly; pleonites 1-3 dorsally

crenulate; outer rami of uropods 1-2 shortened;

rami of uropod 3 apically pointed; telson lacking

large spines or setae dorsally.

Circumboreal, south to Bay of Fundy, Skagerrak,

Sea of Japan, generally 30-200 m, rarely to 682 m.

Tiron thompsoni Walker

Tiron thompsoni Walker, 1904: 263-264, pi. 4, figure 21.

DIAGNOSIS.—Eyes not known; maxillae unknown;

cuter plate of maxilliped with apical excavation

guarded by falcate wings, large spines absent; dactyls

of pereopods 1-5 stubby but apically sharp; article 2

of pereopods 4-5 apparently without special setal

row, posterior margin on article 2 of pereopod 5

unknown; dorsal serrations on pleonites 1-3 not mentioned ; outer ramus of uropod 1 as long as inner, but

shortened on uropod 2; rami of uropod 3 with truncate apices; telson with at least one large spine on

dorsomedial margin of each lobe, perhaps full row

overlooked.

Ceylon.

Tiron tropakis, new species

FIGURES 45, 46

DIAGNOSIS.—Accessory eye composed of 2 segregated ommatidia; inner plate of maxilla 1 fully setose

but margin broken by smooth acclivity; inner plate

of maxilla 1 fully setose but margin broken by smooth

acclivity; inner plate of maxilla 2 with row of submarginal setae; outer plate of maxilliped normal;

dactyls of pereopods 1-5 stubby; article 2 of pereopods 4-5 without special setal row, article 2 of

pereopod 5 weakly crenulate and setulose posteriorly;

pleonites 1-3 dorsally crenulate; outer ramus of

uropod 1 as long as inner, but shortened on uropod

2; rami of uropod 3 apically pointed; each lobe of

telson with medial row of large spines.

Mandibular palp absent (see T. brevidactylus).

DESCRIPTION OF FEMALE.—Dorsoanterior margin

of head curving evenly to rostrum without forming

extended forehead, rostrum pointing obliquely anteroventrally, accessory eye formed of 2 slightly

separated ommatidia; mandible without palp; inner

plates of maxilla 1 large, terminally setose and

medially with long gap between setal groups, outer

plates with 7 spines, inner lobe of maxilla 2 with

dense mediosubmarginal setal row; inner and outer

plates of maxillipeds typical of genus (type-species);

gnathopodal dactyls of medium stoutness and bearing

medium-sized inner tooth; coxae unusually short for

genus, coxa 2 with slight anteroproximal wing, coxa 3

posteriorly excavate, distally expanded and lobate

posteriorly in resemblance of coxa 4; coxa 4 especially

short and broad, comma-shaped, posteriorly excavate;

posterior lobes of coxae 5-6 very large; pereopods 1-5

with short and broad sixth articles, distally appearing

as prehensile because of relative breadth and presence

of large distal spine and peculiar dactyls; dactyls of

pereopods 1-2 formed of a basal subspherical disc

imbedded in end of article 6, disc with hook tooth,

long seta matching curve of hook and accessory small

spine at base of hook; article 6 with large distal

spine; dactyls of pereopods 3-5 similar but hook-spine

only hooked distally, seta similar, basal portion of

dactyl not discoid, article 2 of pereopods 3-4 evenly

ovate, of medium breadth, article 2 of pereopod 5

proximally broad, evenly tapering distally, posterior

margin evenly convex; pleonal epimera 1-3 with

small posteroventral points, each with one or a few

posterior notches, evenly convex margins; pleonites

1-3 dorsally and laterally with small notches becoming increasingly contiguous to form dorsal serrations

but no distinct dorsal teeth present; pleonites 4-5
NUMBER 12 4 87

FIGURE 45.—Tiron tropakis, new species, holotype, female, 4.9 mm, Velero 6688; yrzmale.

4.5 mm, Velero 5828; in toto view with pleopeds removed.
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

oRU3

FIGURE 46.—Tiron tropakis, new species, holotype, female, 4.9 mm, Velero 6688; y=male,

4.5 mm, Velero 5828; wzzfemale, 3.9 mm, Velero 5775;/2=ramus.
NUMBER 12 4 89

with medium and large dorsal teeth, pleonite 6 with

posterodorsal spine, no tooth; uropods 1-2 weakly

spinose, uropod 3 with normally elongate, lanceolate

rami; telson with numerous dorsal spines arranged

in medial row on each lobe.

Male.—Pleonites 4-5 higher than in female, teeth

slightly better developed, pleonite 6 with weak dorsal

hump; antennae elongate, base of primary flagellum

on antenna 1 conjoint; telson with only 2 subapical

spines on each lobe.

HOLOTYPE.—AHF 663, female, 4.9 mm.

TYPE-LOCALITY.—Velero 6688, California, 34° 24'

00"N, 119°50'45"W, 9 m, 3 December 1959.

RELATIONSHIP.—This species resembles T. biocellata in its general appearance except for the

terminal ends of the pereopods which have shortened

sixth articles with prehensoid condition. In this respect T. tropakis resembles T. australis Stebbing

(1908) but differs from that species in the absence

of a mandibular palp, the presence of 2 discrete

lateral accessory ommatidia instead of a small bundle

of several, and apparently the structure of articles 6

and 7 of the pereopods differs but Stebbing does not

clearly draw the condition. Tiron brevidactylus

(Pillai, 1957) also lacks a mandibular palp and has

relatively stout and shortened pereopod apices but

not as strongly modified as in T. tropakis. No accessory eyes are mentioned for T. brevidactylus, apparently the inner lobe of maxilla 1 is small and

without setae, the inner lobe of maxilla 2 has no

medial setae, the posterior lobes of coxae 5-6 are

small, article 2 of pereopod 4 is pear-shaped and

pleonites 1-3 have a distinctly marked middorsal

point almost developed into a tooth.

Tiron thompsoni Walker (1904) has peculiar outer

plates of the maxillipeds, a supernumerary dactylus

on the pereopods, a stout hook-tooth on dactyls of

the gnathopods, less strongly prehensile pereopods

and dorsal points on pleonites 1-3.

Apparently both T. altifrons Reid (1951) and T.

intermedia Reid (1951) have the accessory eye small

and formed of several ommatidia closely compacted,

and article 2 of pereopod 5 is either fully hemispherical (altifrons) or bears a posterodorsal extended wing

marked below by a sinuation (intermedia).

Adults between 5-6 mm lose serrations and points

on pleonal epimera. The holotype has exceptionally

large accessory ommatidia; generally they are as

small as on T. biocellata.

MATERIAL.—Pacific Ocean, Peru: Velero 395

(200+); California: 4836 (1), 4861 (1), 5346 (2),

5614B (1), 5617 (1), 5631 (4), 5756 (1), 5768B

(5), 5775 (6), 5828 (1), 5880 (7), 6205 (1), 6298

(2), 6300 (2), 6395 (1), 6409 (1), 6688 (1) (see

Anonymous, 1965, for Velero data); USNM Ace.

125734, off Balboa, California, February-March 1933,

7-15 fms, G. E. MacGinitie (1); USNM 151/571-

152/584, Laguna Beach, California, Hilton (1);

Caribbean, Venezuela, Velero Atlantic samples, A27

(2), 28 (1), 32 (19), 42 (1); Tampa Bay, Florida,

USNM (1) (no further data); North Carolina, Albatross III, Sta. 19-5, No. 8 amphipod, 24 May 1949,

vertical haul from 10 fms, 34*05^, 77°12/W, Don

W. H. Sutcliffe, Jr. (1); USNM Ace. 183598, Fish

Hawk Sta. 8895, Bottom Net, off Cape Henry,

Virginia, Light Ship, 16.5 fms, 21 October 1920 (1);

USNM D4-41 (1) (data unavailable).

California to Peru, Virginia to Venezuela, 3-157 m.

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Appendix

(STATION LIST)

Samples collected by RV Vema and RV Eltanin.

See Anonymous (1965) for data of samples collected

by RV Velero IV. See J. L. Barnard (1971) for data

of Oregon material collected by Dr. A. G. Carey, Jr.

Eltanin:

50, off southern Peru, 16°12'S, 74°41'W, 2599-2858

m, 15 June 1962.

75, off Chile, 31°10'S, 71°56'W, 1932-3142 m, 24

June 1962.

257, Drake Passage, 61°09'S, 67°51'W, 3867-4086 m.

350, east of Tierra del Fuego, 55°03'S, 58°57'W,

2452 m, 4 December 1962.

Vema:

14-58, west Mediterranean Sea, off Oran, 36°10'N,

01°35'W, 2070 m, 31 July 1958.

15-13, off Caribbean Columbia, ll°30'N, 75°50'W,

2875-2944 m, 8 November 1958.

15-20, off Caribbean Panama, 09°46.5'N, 79°37.5'W,

825-860 m, 10 November 1958.

15-38, off Pacific Colombia, 05°00'N, 79°04/W,

3023-3251 m, 15 November 1958.

15-46, off Pacific Costa Rica, 09°22'N, 89°33'W,

3517-3528 m, 20 November 1958.

15-49, off Pacific Costa Rica, 09°24'N, 89°27'W,

3563 m, 22 November 1958.

15-51, off Pacific Costa Rica, 09°15'N, 89°29'W,

3514-3554 m, 23 November 1958.

15-52, off Pacific Costa Rica, 09°20.5'N, 89°39'W,

3475-3497 m, 23 November 1958.

15-53, off Pacific Costa Rica, 09°23'N, 89°32'W,

3545 m, 23 November 1958.

15-55, off Pacific Nicaragua, 12°45'N, 88°38'W,

3777-3950 m, 11 November 1958.

15-58, off Pacific Nicaragua, 12°11'N, 89°34'W,

5680-5690 m, 27 November 1958.

15-59, off Pacific Panama, 07°05'N, 85°55'W, 2853-

2855 m, 30 November 1958.

15-62, off Ecuador, 01°30'S, 82°19'W, 1363-1369 m,

3 December 1958.

15-131, east of Bahia Blanca, Argentina, 40°14.6'S,

55°24.7'W, 1475 m, 3 April 1959.

15-153, Bahamas Islands, 25°01.5'N, 77°47'W, 2357-

2370 m, 21 June 1959.

92
Index

(One primary page reference to each taxon is cited.)

acanthurus, Tiron, 86

affinis, Syrrhoe, 52

alberti, Bruzeliopsis, 32

altifrons, Tiron, 84

anaticauda, Syrrhoites, 65

angustifrons, Synopia, 51

antarcticus, Tiron, 84

ascua, Bruzelia, 18

Astyra, 5

australis, Bruzelia, 18

Tiron, 84

Austrosyrrhoe, 16

crassipes, 17

fimbriatus, 17

ilergetes, 35

priscis, 44

rinconis, 17

septentrionalis, 17

Ptorpens, 35

bigarra, Garosyrrhoe, 34

Syrrhoites, 34

biocellata, Tiron, 84

bouvieri, Pseudotiron, 45

brevidactylus, Pseudotiron, 85

Tiron, 85

Bruzelia, 17

ascua, 18

australis, 18

cuspidata, 32

dentata, 50

diodon, 19

guayacura, 19

inlex, 21

pericu, 21

(?) popolocan, 24

poton, 27

serrata, 76

tuberculata, 29

typica, 29

Bruzeliopsis, 32

alberti, 32

cuspidata, 32

turba, 32

Calliopius, 6

caraibica, Synopia, 51

coas, Pseudotiron, 45

cohasseta, Syrrhoites, 65

columbiae, Syrrhoites, 65

comanita, Latacunga, 41

crassipes, Austrosyrrhoe, 17

crenulata, Syrrhoe, 52

cu, Syrrhoites, 69

cuspidata, Bruzelia, 32

Bruzeliopsis, 32

dentata, Bruzelia, 50

Stephobruzelia, 50

diodon, Bruzelia, 19

disjuncta, Garosyrrhoe, 34

dulcis, Syrrhoites, 72

fimbriatus, Austrosyrrhoe, 17

Syrrhoe, 17

Syrrhoites, 17

Gammarellus, 6

Garosyrrhoe, 33

bigarra, 34

disjuncta, 34

golens, Pseudotiron, 45

gracilis, Synopia, 51

guayacura, Bruzelia, 19

Ileraustroe, 34

ilergetes, 35

torpens, 35

ilergetes, Austrosyrrhoe, 35

Ileraustroe, 35

inlex, Bruzelia, 21

intermedia, Tiron, 85

Jeddo, 35

simplisyrrhis, 35

Kindia, 2

lorida, 72

sorpresa, 76

Latacunga, 35

comanita, 41

latacunga, 41

longicaudatus, Pseudotiron, 46

longifrons, Syrrhoe, 52

lorida, Kindia, 72

Syrrhoites, 72

Lysianassa spinifera, 86

Melphidippa, 5

Metoediceros, 5

nodulosa, Syrrhoe, 52

oluta, Syrrhoe, 52

orientalis, Synopia, 51

pacifica, Syrrhoites, 73

pantasma, Syrrhoites, 73

papyracea, Syrrhoe, 55

pericu, Bruzelia, 21

pervicax, Pseudotiron, 46

priscis, Austrosyrrhoe, 44

Priscosyrrhoe, 44

Priscosyrrhoe, 44

priscis, 44

popolocan, Bruzelia(?), 24

poton, Bruzelia, 27

Pseudotiron, 44

bouvieri, 45

brevidactylus, 85

coas, 45

golens, 45

longicaudatus, 46

pervicax, 46

psychrophila, Syrrhoe, 56

pusillus, Syrrhoites, 76

redox, Syrrhoites, 76

rinconis, Austrosyrrhoe, 17

scheeleana, Synopia, 51

semiserrata, Syrrhoe, 60

septentrionalis, Austrosyrrhoe, 1 7

serrata, Bruzelia, 76

Syrrhoites, 76

serrima, Syrrhoe, 60

silex, Syrrhoites, 76

simplisyrrhis, Jeddo, 35

sorpresa, Kindia, 76

Syrrhoites, 76

species A, Syrrhoites, 80

spinifera, Lysianassa, 86

spiniferum, Tiron, 86

Stephobruzelia, 50

dentata, 50

Synchelidium, 4, 5

93
94 SMITHSONIAN CONTRIBUTION TO ZOOLOGY

Synopia, 50

angustifrons, 51

caraibica, 51

gracilis, 51

orientalis, 51

scheeleana, 51

ultramarina, 51

variabilis, 52

Syrrhoe, 52

affinis, 52

crenulata, 52

fimbriatus, 17

longifrons, 52

nodulosa, 52

oluta, 52

papyracea, 55

psychrophila, 56

semiserrata, 60

serrima, 60

tuberculata, 64

Syrrhoites, 64

anaticauda, 65

bigarra, 34

cohasseta, 65

columbiae, 65

cu, 69

dulcis, 72

fimbriatus, 1 7

lorida, 72

pacifica, 73

pantasma, 73

pusilla. 76

redox, 76

semiserrata, 60

serrata, 76

silex, 76

sorpresa, 76

species A, 80

tenella, 79

terceris, 79

torpens, 35

trux, 79

walkeri, 79

tenella, Syrrhoites, 79

terceris, Syrrhoites, 79

Tessarops, 83

thompsoni, Tiron, 86

Tiron, 83

acanthurus, 86

altifrons, 84

antarcticus, 84

australis, 84

biocellata, 84

brevidactylus, 85

intermedia, 85

spiniferum, 86

thompsoni, 86

tropakis, 86

Ptorpens, Austrosyrrhoe, 35

Ileraustroe, 35

Syrrhoites, 35

tropakis, Tiron, 86

trux, Syrrhoites, 79

tuberculata, Bnizelia, 29

Syrrhoe, 64

turba, Bruzeliopsis, 32

typica, Bruzelia, 29

ultramarina, Synopia, 51

variabilis, Synopia, 52

Vemana, 5

Vitjaziana, 5

walkeri, Syrrhoites, 79

' U0 GOVERNMENT PRINTING OFFICE: 1972 > I8-J-3I7/24


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