MEMOIRES DU MUSEUM NATIONAL D'HISTOIRE NATURELLE

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3 3001 0001 §§§&#: MNHN, Paris

Dr Isabel Perez Farfante was for

20 years, until 1986, a Research Scientist

in the Systematics Laboratory of the

National Marine Fisheries Sendee, National

Oceanographic and Atmospheric Admin¬

istration, based at the National Museum of

Natural History, Smithsonian Institution,

Washington, D.C. She continues her

research as an Emeritus Research Scientist

with the National Marine Fisheries Service in

Miami, Florida. Concurrently, she was (and

still is) a Research Associate of the National

Museum of Natural History. Prior to 1961

she was Professor of Zoology at the

Universidad de La Habana and Director of

the Centro de Investigaciones Pesquesas de

Cuba. A scientist with broad biological

interests, the penaeoid shrimps of the world

have occupied the prime position in her

research.

Dr Brian Kensley has been a curator in

the Department of Invertebrate Zoology in the

National Museum of Natural History,

Smithsonian Institution, for almost two

decades. Prior to this, he was a research

scientist in Marine Biology at the South

African Museum, Cape Town for 11 years.

He has broad interests in the biology and

systematics of the Crustacea, but carries out

research primarily on the marine Decapoda

and Isopoda.

Source: MNHN, Paris

Penaeoid and Sergestoid Shrimps

and Prawns of the World

Keys and Diagnoses for the Families and Genera

BISL DU

,MUSf ir

RAr :S

* ,

BIBLDU

.MUSEUM

PARIS

* /

Source: MNHN. Paris

ISBN : 2-85653-510-0

ISSN : 1243-4442

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MEMOIRES DU MUSEUM NATIONAL D’HISTOIRE NATURE! I F

TOME 175

ZOOLOGIE

Penaeoid and Sergestoid Shrimps

and Prawns of the World

Keys and Diagnoses for the Families and Genera

Isabel PEREZ FARFANTE* & Brian KENSLEY**

Illustrations by Molly Kelly RYAN**

*NOAA/National Marine Fisheries Service

National Museum of Natural History

Smithsonian Institution

Washington, D.C. 20560

USA

**National Museum of Natural History

Smithsonian Institution

Washington, D.C. 20560

USA

EDITIONS

DU MUSEUM

PARIS

1997

Source: MNHN. Paris

CONTENTS

Abstract/Res um£. 9

Extended Abstract/Resume developpe. 11

Introduction . j 5

Notes on the Format of the Text and Illustrations . 16

Abbreviations . 17

Acknowledgements . 17

Glossary . 19

Systematic Section..

Suborder Dendrobranchiata .31

Superfamily Penaeoidea . 31

Family Aristeidae..

Family Benthesicymidae .56

Family Penaeidae.67

Family Sicyoniidae. 152

Family Solenoceridae . 157

Superfamily Sergestoidea . 182

Family Luciferidae . 183

Family Sergestidae . 185

References . 203

lNDE * . 225

Source:

ABSTRACT

P£rez Farfante, I. & Kensley, B. F., 1997. —Penaeoid and Sergestoid Shrimps and Prawns of the World.

Keys and Diagnoses for the Families and Genera. Mem. Mus. natn. Hist, nat., 175: 1-233. Paris ISBN: 2-85653-

510-0.

Keys for the determination of seven families, and 56 genera of Penaeoidean and Sergestoidcan shrimps are

provided, along with diagnoses of these taxa. The Penaeoidean families are: Aristeidae, Benthesicymidae, Penaeidae,

Sicyoniidae, and Solenoceridae; the Sergestoidean families are: Luciferidae and Sergestidae. Three new genera are

diagnosed: Austropenaeus gen. nov. (family Aristeidae), Megokris gen. nov. and Rimapenaeus gen. nov. (family

Penaeidae). Pelagopenaeus, (family Penaeidae), a name originally proposed by Burkenroad (1934), is given

nomenclatural availability. The earlier names Aristaeopsis Wood-Mason, 1891 (family Aristeidae) and

Trachysalambria Burkenroad, 1934 (family Penaeidae) are resurrected. The subgenera Farfantepenaeus ,

Fenneropenaeus, Litopenaeus , Marsupenaeus, and Melicertus are raised to full genera. In addition to diagnoses for

the families and genera, a section on synonymy for each taxon includes references that have contributed to our

understanding of the taxa. For each genus, a list of its currently recognized species and subspecies, along with

author and date of publication, and distributional records is given. These lists include about 500 species. A

reference is provided in the bibliography for every author and date mentioned in the text. An illustrated glossary

explains the terms used in the diagnoses. At least one species of each genus (often the type species) is illustrated,

along with figures of diagnostic features such as the petasma, thelycum, and seminal receptacles.

RESUME

P£rez Farf'ante, I. & Kensley, B. F., 1997. —Les crevettes peneides et sergestides du monde. Cles

d’identification et diagnoses des families et des genres. Mem. Mus. natn. Hist, nat., 175 : 1-233. Paris ISBN :

2-85653-510-0.

Des cles pour I'identification des sept families et des 56 genres de crevettes peneides et sergestides sont fournies

de meme que des diagnoses de ces divers taxons. Les Peneides comprennent cinq families : Aristeidae,

Benthesicymidae, Penaeidae, Sicyoniidae et Solenoceridae, les Sergestides deux : Luciferidae et Sergestidae. Trois

genres nouveaux sont decrits : Austropenaeus gen. nov. (famille Aristeidae), Megokris gen. nov. et Rimapenaeus

gen. nov. (famille Penaeidae). Pelagopenaeus (famille Penaeidae), un nom propose a l’origine par Burkenroad, est

validd du point de vue nomenclatorial. Les anciens noms Aristaeopsis Wood-Mason, 1891 (famille Aristeidae) et

Trachysalambria Burkenroad, 1934 (famille Penaeidae) sont ressuscites. Les sous-genres Farfantepenaeus ,

Fenneropenaeus, Litopenaeus, Marsupenaeus , et Melicertus sont eleves au rang de genre. En complement aux

diagnoses des families et des genres, les synonymies et les references des travaux ayant contribue a notre

connaissance des taxons consideres sont citees. Pour chaque genre, la liste des especes et sous-especes couramment

admises, avec leurs auteurs, la date de leur description et leur repartition geographique, est fournie. L'ensemble de

ces listes mentionne environ 500 especes. Les references completes de tous les travaux cites dans le texte sont

fournies en fin de travail. Un glossaire illustre explique les termes utilises dans les diagnoses. Au moins une espece

de chaque genre (souvent l'espece-type) est illustree en entier avec, en outre, des dessins representant des caracteres

distinctifs tels que le pdtasma, le thelycum et les vesicules seminales.

EXTENDED ABSTRACT

RESUME DEVELOPPE

This work presents a study of the taxon¬

omy of the penaeidean shrimps of the world.

This group of shrimps includes many of the

commercially important marine species of the

tropics and subtropics, as well as many of the

species that constitute a major link in the food

chain of the oceans. The Penaeidea are made

up of two superfamilies, the Penaeoidea

(families Aristeidae, Benthesicymidae,

Penaeidae, Sicyoniidae, and Solenoceridae),

and the Sergestoidea (families Luciferidae and

Sergestidae). The genera in each of these

families are listed in the table of the page 12.

Keys and diagnoses are provided for these

two superfamilies and seven families, and for

their 56 genera. For each of these genera, a

synonymy, the type species, its type locality,

and gender are given, along with a diagnosis,

branchial formula, and a list of the species in¬

cluded in the genus. References are included

in the synonymy if the work referred to con¬

tributes to an understanding of the diagnosis

of the genus, or includes records that signifi¬

cantly extend the geographical range of the

constituent species. The species and sub¬

species list of almost 500 names includes the

author and date, and geographical records.

Each genus is illustrated with a lateral view of

a whole animal (usually the type species),

petasma, thelycum, and seminal receptacles,

structures which exhibit important diagnostic

characters. Features used in the diagnoses are

defined in a glossary; some of these are also

illustrated. All references giving both author

and date in the text are provided in full in the

bibliography.

Three new genera are diagnosed:

Austropenaeus (family Aristeidae), Megokris

and Rimapenaeus (family Penaeidae).

Austropenaeus, with its type and only species

A. nitidus Barnard, 1947 (previously placed

in Plesiopenaeus) is characterized by having

a reduced podobranch on the third pereopod,

a modified ventral antennular flagellum ami

Ce travail porte sur la taxonomie des cre-

vettes peneides du monde, groupe renfermant

de nombreuses especes marines des regions

tropicales et subtropicales importantes du

point de vue commercial, ainsi que de

nombreuses especes jouant un role primordial

dans la chaine alimentaire des oceans.

Les Penaeidea comprennent deux super-

families, les Penaeoidea avec les families

Aristeidae, Benthesicymidae, Penaeidae,

Sicyoniidae et Solenoceridae et les

Sergestoidea.avec les families Luciferidae et

Sergestidae. Les genres compris dans

chacune de ces families sont indiques dans le

tableau de la page 12.

Des cles d'identification et des diagnoses

sont proposees pour ces deux superfamilies,

ces sept families et les 56 genres qui s'y trou-

vent. Pour chacun des genres, les syno-

nymes, l'espece-type, la localite-type et le

genre grammatical sont indiques, avec en

outre une diagnose, la formule branchiale et

une liste des especes comprises dans le genre.

Les references bibliographiques correspon¬

dent a des travaux qui, soit permettent une

meilleure comprehension du genre, soit indi-

quent de nouvelles recoltes qui elargissent de

maniere significative la repartition de certaines

des especes du genre. Les especes et sous-

especes citees sont pres de 500; pour chacune

sont indiques son auteur, la date de sa

description et sa repartition geographique.

Chaque genre est illustre par la vue laterale

d'une crevette entiere (habituellement

l'espece-type), le petasma, le thelycum et,

souvent, les receptacle seminaux, structures

qui fournissent des caracteres d'identification

importants. Les caracteres utilises dans les

diagnoses sont definis dans un glossaire et

beaucoup sont illustres. Toutes les references

mentionnees dans le texte sont citees in ex-

tenso a la fin du volume.

Trois nouveaux genres sont decrits :

l'un, Austropenaeus, appartenant a la famille

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

Family ARISTEIDAE

Aristaeomorpha

Aristaeopsis

Aristeus

Austropenaeus

Hemipenaeus

Hepomadus

Parahepomadus

Plesiopenaeus

Pseudaristeus

Family BENTHESICYMIDAE

Bentheogennema

Benthesicymus

Benthonectes

Gennadas

Family PENAE1DAE

Artemesia

Atypopenaeus

Farfantepenaeus

Fenneropenaeus

Superfamily PENAEOIDEA

Funchalia

Heteropenaeus

Litopenaeus

Macropetasma

Marsupenaeus

Megokris

Melicertus

Metapenaeopsis

Metapenaeus

Miyadiella

Pcirapenaeopsis

Parapenaeiis

Pelagopenaeus

Penaeopsis

Penaeus

Protrachypene

Rimapenaeus

Tanypenaeus

Trachypenaeopsis

Trachypenaeus

Trachysalambria

Xiphopenaeus

Family S1CYON1IDAE

Sicyonia

Family SOLENOCERIDAE

Cryptopenaeus

Gordonella

Hadropenaeus

Haliporoides

Haliporus

Hymenopenaeus

Mesopenaeus

Pleoticus

Solenocera

Superfamily SERGESTOIDEA

Family LUCIFERIDAE

Lucifer

Family SERGESTIDAE

Acetes

Peisos

Petalidium

Sergestes

Sergia

Sicyonella

Families and genera in the superfamilies Penaeoidea and Sergestoidea

Families et genres des superfamilies Penaeoidea et Sergestoidea

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

modified third maxilliped in the male, and

feebly developed carapace carinae. Megokris

is separated from Trachypenaeus s.l., and is

characterized by features of the thelycum, by

the seminal receptacles being connected

neither by a transverse groove nor by a

median pocket, and by the features of the

petasma. Rimapenaeus is also part of the

complex of genera previously known as

Trachypenaeus s.l., and is characterized by

features of the thelycum and especially of the

internal trilobed seminal receptacles, and by

the characters of the petasma.

Pelagopenaeus (family Penaeidae) a name

originally proposed by Burkenroad (1934),

but not valid, no type species having been

designated, is given nomenclatorial validity.

The generic names Aristaeopsis Wood-

Mason, 1891, and Trachysalambria Burken¬

road, 1934, are resurrected. The former, with

its type and only species A. edwardsiana, is

characterized by the absence of exopods on

the first to fifth pereopods, the presence of a

rounded middorsal carina on the second

abdominal somite, a sharp middorsal carina

on the third abdominal somite, a modified

antennal scaphocerite in the male, and by the

absence of a distal meral spine on the first and

second pereopods. Trachysalambria, one of

the complex of genera formerly under

Trachypenaeus s.l., is characterized by the

features of the thelycum and petasma, in

having bilobed seminal receptacles opening

into a common groove, and in having a short

longitudinal suture on the carapace.

Five subgenera of Penaeus are raised to

generic level, namely Farfantepenaeus,

Fenneropenaeus, Litopenaeus, Marsupenaeus

and Melicertus.

Aristeidae, les deux autres, Megokris et

Rimapenaeus, a la famille Penaeidae.

Austropenaeus, qui ne comprend actuelle-

ment qu'une seule espece, A. nitidus

Barnard, 1947 (precedemment placee dans le

genre Plesiopenaeus) se caracterise par une

podobranchie de taille reduite sur les

troisiemes pereiopodes, un flagelle anten-

nulaire ventral et des troisiemes maxillipedes

modifies chez le male, des carenes faiblement

marquees sur la carapace. Megokris se separe

des Trachypenaeus s.l. et s'en distingue par

certains caracteres du thelycum, les recep¬

tacles seminaux qui ne sont connectes entre

eux ni par un sillon transversal ni par une

poche mediane, et les caracteres du petasma.

Rimapenaeus fait egalement partie du

complexe Trachypenaeus s. 1. et se distingue

par la disposition du thelycum, des

receptacles seminaux trilobes et les caracteres

du petasma.

Pelagopenaeus, nom de genre propose par

Burkenroad en 1934 mais qui n’etait pas va-

lide aucune espece-type n’ayant ete designee,

est valide.

Les genres Aristaeopsis Wood Mason,

1891, et Trachysalambria Burkenroad, 1934,

sont ressuscites. Le premier, avec pour

unique espece A. edwardsiana, se caracterise

par l'absence d’exopodes sur les cinq paires

de pereiopodes, la presence d'une carene

dorsale arrondie sur le deuxieme segment

abdominal, aigue sur le troisieme, un scapho¬

cerite modifie chez le male, et l'absence d'une

epine sur le merus des premiers et seconds

pereiopodes. Trachysalambria, l'un des gen¬

res du complexe autrefois designe sous

l'appellation Trachypenaeus s.l., se distingue

par les caracteres du thelycum et du petasma,

des receptacles seminaux bilobes s'ouvrant

dans un sillon commun, une courte suture

longitudinale sur la carapace.

Cinq sous-genres de Penaeus sont eleves

au rang de genre. Ce sont: Farfantepenaeus,

Fenneropenaeus, Litopenaeus, Marsupenaeus

et Melicertus.

INTRODUCTION

Crustaceans including shrimps of various kinds have probably been a source of food for

humans from very early times. Within historical times reference is made to shrimps in ancient

Chinese and Japanese literature. The earliest known published description and figure of a penaeid

shrimp in Western literature is that of Melicertus kerathurus, referred to by Guillaume Rondelet

in his Libri de Piscibus Marinis of 1554 as "Squilla Crangone" (Holthuis, in litt. 1996). In more

modern scientific literature, the species now known as the American white shrimp, Litopenaeus

setiferus, was referred to by Seba (1759) as Astacus fluviatilis Americanus, and shortly

thereafter, as Cancer setiferus by Linnaeus (1767). By 1814, the Penaeoidea had been

recognized as a taxonomic grouping by Rafinesque-Schmaltz. Since then, the literature on many

aspects of the systematics and biology of this assemblage has grown enormously, not least

because of the economic importance of many of the constituent species (see Dall et al. , 1990, for

an extended taxonomic history of the Penaeoidea).

The term Dendrobranchiata, coined by Bate (1888) to distinguish a group of crustaceans from

the Trichobranchiata and the Phyllobranchiata, originally included the Normalia (Penaeidae and

Sergestidae) and the Aberrantia (Euphausiacea and Mysidacea). This division was based on

Bate's interpretation of gill structure.

In the present-day conception of the Dendrobranchiata, seven families, 56 genera, and some

500 described species are recognized. Two families, the Penaeidae and Sicyoniidae, contain

many of the shrimp species caught commercially in the tropics and subtropics around the world.

The commercial shrimp catch for the United States alone in 1996 was valued at almost

$463 million, representing about 14% of the total commercial fisheries catch, while the shrimp

imports for the same year totalled $2.5 billion. For several countries, shrimp exports constitute a

major source of foreign currency income. Given this economic significance, and the fact that

penaeoideans occur in a wide variety of shallow-water marine and estuarine habitats, knowledge

of the ecology, feeding, reproduction, life cycles, population dynamics, predation, and fisheries

potential has vastly increased over the last three decades. In the Gulf of Mexico, penaeidean

shrimps play an important role as the prey species for a variety of commercial fish species such

as Atlantic croaker, silver perch, southern flounder, sand seatrout, spotted seatrout, and

lizardfish (Minello and Zimmerman, 1984; Minello et al, 1989: 13; $heridan et al, 1984).

Further, the commercial aquaculture production of a range of species has given an added impetus

to research into these organisms. A thorough knowledge of the systematics of these shrimps is

clearly an essential prerequisite for wise management and exploitation, and for placing the great

body of biological knowledge in context.

As with many invertebrate groups, the morphology of the external and internal reproductive

structures plays a fundamental role in interpreting the systematics of the Dendrobranchiata. This

aspect of the biology of the group has received considerable attention, given the central role of

reproduction in our understanding of fecundity, life histories, and exploitation of the group. Our

own studies have led us to reinterpret especially thelycal structures, and in particular those of the

seminal receptacles, a number of which had not previously been identified and which are

described herein for the first time. Close attention to these latter structures has shown many

earlier works to have erred in inteipreting the nature of an 'open' and a 'closed' thelycum. The

importance of these structures in the evolutionary success of the group has thus led us to pay

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

particular attention to the characters to be found in the reproductive morphology. Brief notes on

the spermatophores of a number of genera are given. These structures, although having a similar

basic pattern for each genus, may exhibit variation among congeneric species, as, for example, in

the species of Litopenaeus (Perez Farfante, 1975).

The three predominantly deepwater families, the Aristeidae, Benthesicymidae, and Sergestidae

contain species that are either deep benthic dwellers or are members of the meso- and

bathypelagic fauna of the oceans. Given their often great abundance in the water column, these

forms play a major role in the food chain of the oceans, many being active predators, and in turn

are the prey of many predatory fishes such as the various species of tuna (Dragovich, 1969).

The species of the seventh family, the Luciferidae, are true planktonic forms. The relationship

of this family to the rest of the Dendrobranchiata, however, is unclear, and is based on traditional

usage rather than shared advanced characters.

In spite of the importance of the shallow and deepwater Dendrobranchiata, the taxonomy of

the group has been unusually fluid and sometimes confused, with some genera poorly defined.

This is particularly the case for several genera having their greatest diversity in the Indo-West

Pacific region, e.g. Parapenaeopsis, for which no modern revision is available. Milestones of

taxonomic significance for the penaeideans include the works of Bate (1888), Kubo (1949),

Burkenroad (1934a, 1934b, 1983), Yaldwyn (1957), Perez Farfante (1969, 1977a, 1980a,

1985), Crosnier (1978, 1985, 1987, 1989, 1991, 1994a), and Dall etal. (1990), and those of

Heldt (1938a, 1938b) and Hudinaga (1942) on reproduction. Given this plethora of sources,

we were prompted to draw together all relevant taxonomic works on the dendrobranchiates, and

to add some of our own observations and data, to produce a single comprehensive reference that

would serve as a basic tool in the systematics of the group. In this task we were inspired by the

works of Holthuis (1955a, 1993) on the Caridea, whose summations of taxonomic knowledge

have proved to be indispensible in understanding that other large group of shrimps.

We have not considered the fossil history of the dendrobranchiates in this work. The poor

quality of the available fossils, and their dubious taxonomy (often linked to modern taxa with no

basis in real shared characters), pursuaded us that the paleontological evidence added little to our

understanding of the extant forms.

As for larval forms, many of which have formal generic and specific designations but which

have not been linked to adult forms, we decided not to deal with these (unless the adult

connection has been made, in which case any taxonomic implications are of course taken into

consideration), again because their uncertain systematic position would add little to our

understanding of the adult systematics.

Notes on the Format of the Text and Illustrations

Superfamilies, families, genera within families, and lists of species are arranged alphabetically

for ease of use. For convenience of usage, extended synonymies giving abbreviated journal titles

are provided; the abbreviations for journals used are those of the World List of Scientific

Periodicals (Brown and Stratton, 1963). Abbreviations for non-journal references are of our

own devising, and are used consistently throughout. References are included that contribute to

the definition of a taxon; mere mention of the taxon or of a geographic record did not warrant

inclusion in the synonymy.

In the descriptions, the middorsal armature of the rostrum and carapace are consistently

referred to as teeth, not spines, in spite of the confusion over the use of these two terms. Where

carapace length is given, as in the legends to figures, this is measured from the orbital margin to

the mid-dorsal posterior margin of the carapace. The fourteen thoracic somites, and especially the

posterior four, are referred to by Roman numerals, a tradition that has been in use for more than

60 years.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

For the list of species given for each genus, every attempt has been made to survey all relevant

taxonomic literature. Nevertheless, there may well be species missing from the list, and

synonyms amongst the cited names, especially in the larger genera that have not recently been

revised.

In the geographical distribution for species, some new records have been included, most

representing significant range extensions, from either the National Museum of Natural History or

other collections; these are not indicated separately as new records. Geographic records are

generally given from west to east, starting with the Eastern Pacific Ocean.

Illustrations for each genus include a lateral view of the whole animal of a representative

species (wherever possible of the type species), as well as the female thelycum and the male

petasma. In a few cases, additional illustrations of important diagnostic features are included. The

figures illustrating the seminal receptacles, most of which include the openings for the entrance

and exit of the sperm, were drawn from specimens that had been cleared and stained in hot lactic

acid and chlorozol black (see Monod and Cals, 1970, for this technique).

In addition to giving full references in the Literature Cited section for all citations in the

synonymies, wherever a taxon with author and date is given, a full reference has been provided,

as is fitting in a work intended to be used as a taxonomic resource.

Abbreviations

cl - carapace length, tl - total length.

Abbreviations used in the branchial formula tables: r - reduced; s - small; v - vestigial;

lam - branchial lamella.

ACKNOWLEDGEMENTS

Material used in this study came from several sources in addition to the collections of the

National Museum of Natural History (NMNH), Smithsonian Institution. We are very grateful to

the following individuals and institutions for the loan or donation of material: Enrique Boschi,

INIDEP, Mar del Plata, Argentina; Alain Crosnier, ORSTOM Senior Scientist, who facilitated

many loans from the Museum National d'Histoire Naturelle, Paris; Ken-Ichi Hayashi, of

Shimonoseki University, Shimonoseki, Japan; Lipke Holthuis and C.H.J.M. Fransen of the

Rijksmuseum van Natuurlijke Historic, Leiden, The Netherlands; Ardis Johnson of the Museum

of Comparative Zoology, Harvard University, Cambridge, Massachusetts; J. Liu, Institute of

Oceanology, Academia Sinica, China; Spencer R. Luke, Scripps Institution of Oceanography,

La Jolla, California; Michelle van der Merwe of the South African Museum, Cape Town;

Paul Clark of The Natural History Museum, London; Torben Wolff of the Zoological

Museum, University of Copenhagen, Denmark.

Much of the impetus for this project came from the enthusiastic encouragement of colleagues

in the Department of Invertebrate Zoology, National Museum of Natural History, including

Fenner A.Chace, Jr., the late Horton H. Hobbs, Jr., Raymond B. Manning, and from

Austin B. Williams of the Systematics Laboratory of the National Marine Fisheries Service, at

the Smithsonian Institution. All of these contributed valuable discussions on points of

systematics, the interpretation of features, and the clear expression of concepts. Ray Bauer of the

University of Southwestern Louisiana, and Alain Crosnier of the Paris Museum were unstinting

in sharing ideas, reading drafts of sections of this work, and pointing out missed references.

Lipke B. Holthuis of the Nationaal Natuurhistorisch Museum, Leiden, with his usual

exemplary thoroughness, sifted through the manuscript and made suggestions and corrections,

and also pointed out many missed references and synonymies, especially those involving older

literature or complex taxonomic tangles, of which is he is undisputed master. To all of these, we

are extremely grateful for their support and generosity.

ISABEL P£*EZ FARFANTE & BRIAN KENSLEY

The National Marine Fisheries Service Systematics Laboratory provided general support to the

senior author, including advice and material resources furnished through laboratory directors

Daniel M. Cohen and Bruce B. Collette, and post-retirement funding through NOAA/NMFS

Contract 50-EANF-0-00035.

A few of the illustrations were taken from earlier works of the first author; these illustrations

were executed by Maria M. Di£guez, and are acknowledged as such. The bulk of the

illustrations, however, are the work of Molly Kelly Ryan, Scientific Illustrator in the Department

of Invertebrate Zoology, NMNH, Smithsonian Institution. Ms. Ryan's dogged devotion to this

project in the face of sometimes strident competition for her talents, has earned our undying

gratitude. Her superb renderings and attention to our demands for accuracy of detail speak to her

unflinching patience and unvarying pleasantness.

Alain Crosnier encouraged us to submit our manuscript to the Paris Museum, and when it

was finally accepted, undertook the detailed editing. During this process, because of his

knowledge of the penaeideans, he was able to expose several problems and errors and suggested

improvements well beyond the normal scope of an editor. We are truly indebted to him.

antennule

'v.v^vrr~

antenna

scaphocerite

cicatrix

pleuron

telson

third 4

maxilliped

\pleopods/

uro

mesial ramus

lateral ramus

tergum

dorsomedian

carina

dorsolateral

sulcus

pereopods —

— rostrum-carapace-abdomen

antennal flagellum

eye

Fig. 1. — Schematic drawing of a penaeidean shrimp in lateral view.

Source: MNHN, Paris

GLOSSARY

Plural form in parentheses.

Abdomen. That part of the body posterior to the cephalothorax, consisting of six body segments

or somites plus the telson; the "tail" (Fig. 1)

Accessory Lobe. Small lobe at base of median lobe, on anterior surface of petasma in Gennadas.

Adrostral Carina. Ridge flanking the rostrum, sometimes nearly reaching the posterior margin

of the carapace (Fig. 2B).

Adrostral Sulcus. Groove flanking the rostrum mesial to the adrostral carina, sometimes nearly

reaching the posterior margin of the carapace (Fig. 2B).

Antenna (antennae). More lateral of the two paired, usually flagellate appendages projecting

distally from the anterior end of the cephalothorax (Fig. 1).

Antennal Carina. Ridge extending posteriorly along dorsal extremity of antennal region, often

continuous with antennal spine (Fig. 2B).

Antennal Flagellum (Antennal Flagella). Multiarticulate, whiplike terminal part of the

antenna (Fig. 1).

Antennal Peduncle. Five basal segments of the antenna, from which the flagellum arises

distally.

Antennal Region. Area on the lateral face of the carapace posterior to and encompassing the

antennal spine.

Antennal Spine. Spine situated on the anterior margin of the carapace just ventral to the orbital

margin (Fig. 2A).

Antennular Flagellum (Antennular Flagella). Multiarticulate paired filaments (sometimes

flattened and lamellate) of the antennule (Fig. 3B).

Antennular Peduncle. Three basal segments of the antennule, from which the flagella arise

distally (Fig. 3B).

Antennule. More mesial of the two paired, usually flagellate appendages projecting from the

anterior end of the cephalothorax (Fig. 1, 3B).

Anterior Process. Anterior part of an elongate median protuberance lying on the penultimate

(XIII) thoracic sternite (Fig. 4A).

Anterolateral Carina. Longitudinal ridge extending along anterior pail of carapace, ventral to

gastro-orbital carina.

Appendix Interna (Appendices Internae). Slender lappet, sometimes rodlike, at the mesial base

of the endopod of the second pleopod (dorsal to the appendix masculina) of many males.

Appendix masculina (Appendices masculinae). Lappet, sometimes scalelike, at the mesial base

of the endopod of the second pleopod (ventral to the appendix interna if latter present) in some

males.

Arthrobranchia (Arthrobranchiae). Branchia (gill) attached to the joint area between the body

and the first podomere of the leg (Fig. 3C).

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Article. Any one of the subdivisions of an appendage segment.

Articular Membrane. Uncalcified integument at a joint, permitting movement of the

exoskeleton, as between the segments of a pereopod.

Basial Spine. Spine projecting from basis of a thoracic appendage.

Basis (Bases). Second podomere or segment, from the proximal end of a typically

7-segmented appendage (Fig. 3C).

Branchia (Branchiae). Respiratory organ (gill) associated with an appendage or with the body

wall.

Branchial Region. Area of the carapace overlying the branchial cavity.

Branchiocardiac Carina. Ridge extending along posterodorsal limit of branchiostegite

(Fig. 2A).

Branchiocardiac Sulcus. Groove extending along dorsal limit of branchiostegite, running

parallel to branchiocardiac carina.

Branchiostegal Carina. Longitudinal ridge extending along -anteroventral part of carapace,

usually continuous with branchiostegal spine.

Branchiostegal-hepatic Carina. Longitudinal ridge consisting of the fusion of the

branchiostegal and hepatic carinae.

Branchiostegal Spine. Short spine on or near the anterior margin of the carapace ventral to the

antennal spine and dorsal to the anteroventral angle of the carapace (Fig. 2A).

Branchiostegal Sulcus. Groove often accompanying branchiostegal carina, located on the

anteroventral part of the carapace.

Branchiostegite. Expanded ventro-lateral part of the carapace covering the gills (Fig. 2A).

Calathus. See Optic Calathus below.

Carapace. "Flead shield" cuticular structure arising from posterior margin of cephalon,

extending anteriorly and posteriorly, and covering the cephalothoracic somites of the body

(Fig. 1).

Carina (Carinae). A ridge or keel of the exoskeleton (Fig. 1, 2).

Carpus (Carpi). Fifth podomere from the proximal end of a typically 7-segmented appendage

(Fig. 3C).

Cephalothorax. Anterior part of the body consisting of the fused cephalon (head) and thorax,

bearing all the appendages except the pleopods and uropods.

Cervical Carina. Mesially transverse and laterally oblique ridge extending from the anterior

limit of the hepatic region towards middorsal line of carapace (Fig. 2A).

Cervical Sulcus. Mesially transverse and laterally oblique groove of the carapace extending

from the anterior limit of the hepatic region towards the midline of the carapace (Fig. 2B).

Chela (Chelae). Pincer formed by the two distal podomeres of a pereopod in which the movable

finger (dactyl) opposes a fixed finger formed by a distal extension of the propod (Fig. 3C).

Chelate. Appendage ending in chela (claw).

Cicatrix (Cicatrices). Longitudinally disposed ridge(s) often present on lateral part of sixth

abdominal somite (Fig. 1).

ISABEL P^REZ FARFANTE & BRIAN KENSLEY

Fig. 3.

A: EYE

basal segment

tubercle

ocular

distolateral spine

3rd

ventrolateral

flagellum

B: ANTENNULE

segments — 1 prosartema

antennular peduncle-

— Features of the eye (A), antennule (B), pereopod (C) and telson (D) used in penaeoid shrimp taxonomy.

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Cincinnuli (plural of Cincinnulus). Minute interlocking processes projecting from the

dorsomesial margins of the petasmal endopods (Fig. 4B).

Cornea. Faceted, usually pigmented portion of the eye (Fig. 3A).

Coxa (Coxae). First or proximal podomere of a typically 7-segmented appendage (Fig. 3C).

Coxal Spine. Spine projecting from the coxa of a thoracic appendage.

Dactyl. Terminal podomere of a typically 7-segmented appendage (Fig. 3C).

Dendrobranchiate Gill. One in which the paired primary branches are subdivided, sometimes

highly so.

Distal Fold. Distal pleat in the dorsolateral lobule of the petasma.

Distolateral Projection. Distolateral, ventrally inclined projection or spur of the basis of the

endopod of the male second pleopod.

Distomedian Projection. Distal, relatively narrow extension of the dorsomedian lobule of the

petasma.

Distoventral Projection. Outer distal flap articulating with distal extremity of ventrolateral

lobule of petasma in members of the genus Metapenaeopsis.

Dorsolateral Carina. Longitudinal ridge on dorsolateral region of carapace running dorsal to

orbital region.

Dorsolateral Lobule. Dorsal part of the lateral lobe of the petasma (Fig. 4B).

Dorsolateral Sulcus. Longitudinal groove sometimes present close to the dorsomedian line of

the sixth abdominal somite.

Dorsomedian Carina. Ridge extending along the middorsal line of the abdominal somites

(Fig. 1).

Dorsomedian Lobule. Mesial part of the median lobe of the petasma (Fig. 4B).

Endopod. Mesial ramus of a biramous appendage, especially one arising from the basis or from

the protopodite of the pleopod.

Endite. Lobe of several proximal podomeres of various appendages.

Epigastric Tooth. Tooth on the carapace situated above the gastric region behind the first

(posteriormost) rostral tooth (Fig. 2A).

Epipod. Lateral exite of the coxa of a thoracic appendage, sometimes branchial in function

(Fig. 3C).

Epistome. Transverse plate anterior to mouth area.

Exopod. Lateral ramus of a biramous appendage, arising from the basis, or from the protopodite

(Fig. 3C).

Eyestalk. Peduncle or unfaceted part of the eye supporting the cornea.

Flagellum (Flagella). Multiarticulate, usually whiplike terminal part of the antennule or antenna

(Fig. 3B).

Frontal Region. Anterior area of the carapace lying between the orbits and bounded posteriorly

by the gastric region.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Gastric Region. Principal median area of the carapace bounded anteriorly by the frontal and

orbital regions, and posteriorly by the cardiac region, and laterally by the branchial and hepatic

regions.

Gastrofrontal Carina. Short longitudinal ridge extending posteriorly from the ventral

extremity of the orbital region (Fig. 2B).

Gastrofrontal Sulcus. Short longitudinal depression accompanying the gastrofrontal carina

dorsally (Fig. 2B).

Gastrohepatic Gland. Digestive gland associated with the midgut, within the cephalothorax.

Gastro-orbital Carina. Short longitudinal ridge extending (often curving) anterodorsally from

the cervical sulcus towards the orbital region (Fig. 2B).

Genitalia. The external reproductive structures.

Hepatic Carina. Longitudinally or obliquely disposed ridge of variable length lying ventral to

the hepatic region, sometimes extending almost to the anterior margin of the carapace.

Hepatic Region. Paired anterolateral areas of the carapace bounded anteriorly by the antennal

region, posteriorly by the branchial region, and mesially by the gastric region.

Hepatic Spine. Lateral spine situated near the anterior margin of the hepatic region of the

carapace (Fig. 2A).

Hepatic Sulcus. Groove ventral to the hepatic region extending posteriorly, sometimes from

near the anterior margin of the carapace.

Incisor Process. Cutting process, often toothed or cusped, of the mandible.

Insemination. The act of placing or introducing sperm onto or into the thelycum or seminal

receptacles of the female.

Integument. Outer covering or exoskeleton.

Ischial Spine. Spine projecting from ischium or third segment of thoracic appendage.

Ischium (Ischia). Third podomere from the proximal end of a typically 7-segmented appendage

(Fig. 3C).

Labrum. Upper lip or unpaired structure arising anterior to the mouth and often covering it.

Lateral Lobe. One of the paired lateral parts, often folded, of the petasma (Fig. 4B).

Lateral Plate. One of the paired, adjacent flaps sometimes present on sternite XIV in females

(Fig. 4A).

Lateral Ramus (of uropod). Outer branch (not necessarily equivalent to the exopod) (Fig. 1).

Longitudinal Suture. Fine longitudinal line extending posteriorly just above the base of the

antennal spine.

Mandible. One of the heavily calcified jaws lying anterior to (beneath, in ventral view) to other

mouth parts.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

protuberance

anterior process

—posterior process

lateral

posterior protuberance

A: THELYCUM

4. — Some features of the external genitalia, thelycum (A) and petasma (B), used in the penaeoid shrimps

taxonomy.

Source

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Mandibular Palp. One- to three-segmented endopod attached laterally to the masticatory part of

the mandible.

Maxilla (Maxillae). Paired mouthpart appendages of the fourth and fifth cephalic somites.

Maxilliped. One of a pair of three sets of thoracic appendages, arising posterior to the primary

mouthparts. The two anterior pairs are often modified for feeding, while the third pair (Fig. 1)

is often pediform, resembling the pereopods.

Median Lobe. One of the paired dorsal parts, often folded, of the petasma (Fig. 4B).

Median Protuberance. Conspicuous elevation, sometimes platelike, arising from the

posteromedian part of the penultimate (XIII) thoracic sternite (Fig. 4A).

Median Sulcus. Dorsomedian groove on the carapace (Fig. 2B).

Merus (Meri). Fourth segment from the proximal end of a typically 7-segmented appendage

Mesial Ramus (of uropod). Inner branch (not necessarily equivalent to the endopod) (Fig. 1).

Mesial Tubercle. Strong conical to low, rounded protuberance on mesial surface of optic

calathus.

Ocular Plate. Median cephalic plate bearing the eyestalks laterally (Fig. 3A).

Ocular Scale. Scale-like structure located on basal segment of eyestalk (Fig. 3A).

Ocular Stylet. Spiniform process arising from ocular plate in the genus Sicyonia (Fig. 3A).

Optic Calathus. Terminal article of the eyestalk supporting, often embracing the cornea of the

eye (Fig. 3A).

Orbital Margin. Anterior border of the carapace, often contiguous to the eye.

Orbital Region. Paired areas on the carapace just posterior to the eyes.

Orbital Spine. Spine projecting from the ventral extremity of the orbital margin (Fig. 2A).

Orbito-antennal Sulcus. Longitudinal or oblique depression between the orbital margin and the

hepatic spine (Fig. 2A).

Organs of Pesta. Luminescent modified areas of the gastrohepatic glands, found within the

cephalothorax of some species of Sergestes s.l.

Palm. Portion of the chela proximal to the propodal finger (Fig. 3C).

Parapenaeid Spine. Spine projecting from the distomesial margin of the first antennular segment.

Pereopod. One of the five posterior paired appendages or legs of the cephalothorax (Fig. 1, 3C).

Petasma (Petasmata). The male genital structure consisting of the much enlarged and coupled

endopods of the first pair of pleopods. The presence or absence of a petasma, or in juveniles

the position of the first endopods (situated more distally in females than in males), is the

easiest means of distinguishing between the sexes in penaeidean shrimps (Fig. 1,4B).

Open Petasma. The lateral lobes are quite flexible, partially or entirely extended

laterally, with the ventral costae not or barely turned ventrally.

Source . MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Semi-open Petasma. The lateral lobes are flexible but folded, with the ventral costae

distinctly turned ventromesially, delimiting relatively ample space extending from

proximal to distal ends.

Semi-closed Petasma. The lateral lobes are rather flexible, markedly folded, supported

by strong ribs, with the ventral costae approaching rather closely, delimiting

moderately large space, narrowly open distally where usually overlapped by well

developed distomedian projections.

Closed Petasma. The lateral lobes are heavily sclerotized, sometimes making structure

virtually rigid, with the ventral costae situated ventromesially, almost abutting, and

delimiting a small, sometimes extremely so, space; lateral lobe usually produced

distally into lateral spouts or horns.

Photophore. Luminescent organ of the integument.

Phyllobranchiate Gill. A gill in which the branches are platelike, usually occurring in paired

series along the gill axis.

Pleopod. One of the biramous paired appendages typically arising ventrally from each of the

anterior five abdominal somites. In the shrimps, they are primarily swimming organs (Fig. 1).

Pleurobranchia (Pleurobranchiae). Gill attached to the body-wall (pleural membrane), dorsal

to the articulation of the appendage (Fig. 3C).

Pleuron (Pleura). One of the lateral flaps on each of the anterior five abdominal somites

(Fig. 1).

Podobranchia (Podobranchiae). Gill borne on the basal segment (coxa) of a thoracic appendage

(Fig. 3C).

Podomere. Any one of the segments of an appendage, such as a segment of a pereopod or

maxilliped.

Postantennal Spine. Spine located on anterolateral area of carapace on the posterior part of the

antennal region.

Postcervical Spine. Spine located immediately posterior to cervical carina.

Postcervical Sulcus. Subvertical carapace groove located posterior to cervical sulcus.

Posterior Process. Posterior part of an elongate median protuberance projecting caudally onto

the last (XIV) thoracic sternite (Fig. 4A).

Posterior Protuberance. Conspicuous elevation arising from the posteromedian part of the last

(XIV) thoracic sternite (Fig. 4A).

Posthepatic Carina. Ridge posterior to hepatic carina, extending onto lower branchiostegite

(Fig. 2A).

Postocular Sulcus. A small groove situated near the dorsal extremity of the orbital margin.

Postorbital Spine. Spine situated near the orbital margin posterior to the antennal spine

(Fig. 2A).

Postrostral Carina. Dorsomedian ridge extending posteriorly from the base of the rostrum,

sometimes nearly reaching the posterior margin of the carapace (Fig. 2A).

Processus uncifer. Mesial, well separated lobule, usually having a terminal hook, in the petasma

of sergestids.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Processus ventralis. Ventromesial lobule located on distal half of pars media of the petasma of

sergestids.

Propodus (Propodi). Sixth or penultimate segment of a typically 7-segmented appendage

(Fig. 3C).

Prosartema. Variable in shape, thin, sometimes scalelike process arising from the mesial base of

the first antennular segment, and extending distally (Fig. 3B).

Protocephalon. Anteriormost part of the body bearing eyes.

Pterygostomian Carina. Ridge running posterior to pterygostomian spine on antero-ventral part

of carapace.

Pterygostomian Region. Anteroventral area of the carapace.

Pterygostomian Spine. Marginal spine arising from the anteroventral angle or border of the

carapace.

Ramus (Rami). A branch of an appendage, e.g. the exopod or endopod.

Rostrum (Rostra). Anteromedian projection of the carapace between the eyes (Fig. 1).

Scaphocerite. Laterally rigid lamellate exopod of the antenna; the antennal scale (Fig. 1).

Segment. Division of an appendage.

Seminal Receptacles. Unpaired or paired bulbous or tubular sacs associated with the thelycum

for the storage of spermatophores or sperm, situated immediately dorsal to plate(s) of sternite

XIV, sometimes XIII and/or XII.

Unpaired Receptacles. Non-invaginated, opening either through an exposed median

longitudinal slit flanked by lateral plates of sternite XIV, or opening anterior to single

plate.

Paired Receptacles. Invaginated into the cephalothoracic cavity, opening through well

protected slits usually on anterior border of sternite XIV, or on anterior"part of XIII.

Somite. Each of the main divisions of the body.

Spermatophore. The sperm-carrying, variously complex mass, issuing from the male gonopore

during copulation.

Statocyst. Sensory organ of awareness of rotation and position located at base of first antenna.

Sternite. Ventral part of a thoracic or abdominal somite.

Sternum. Ventral surface of the cephalothorax or abdomen.

Stylocerite. Pointed scale arising from the lateral base of the first segment of the antennular

peduncle (Fig. 3B).

Subhepatic Sulcus. Groove located well ventral to the hepatic region of the carapace and the

hepatic spine.

Sublateral Sulcus. Groove located ventral to lateral carina of carapace.

Submarginal Carina. An almost longitudinal ridge extending between rigid, and membranous

part of the branchiostegite (Fig. 2A).

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Subrostral Sulcus. A longitudinal elongate groove extending along dorsal limit of the orbital

region.

Sulcus (Sulci). Groove.

Suprahepatic Spine. Spine arising from the edge of the cervical carina dorsal to the hepatic spine

(Fig. 2A).

Supraorbital Spine. Spine located posterior to the orbital margin of the carapace.

Suture. Either transverse or longitudinal, weakly sclerotized line or seam on the carapace.

Telson. Terminal unit of the abdomen bearing the anus (Fig. 1, 3D).

Tergum (Terga). Arched dorsal part of each of the anterior five abdominal somites (Fig. 1).

Thelycum (Thelyca). The female genitalia consisting of modifications of the posterior two, or

sometimes three thoracic sternites (XII-XIV) serving for the storage or transfer of the sperm,

usually in spermatophores, and often shielding seminal receptacles (Fig. 4A).

Open Thelycum. One in which the seminal receptacles are absent.

Closed Thelycum. One in which the seminal receptacles are present.

Thoracic Ridge. Highly sclerotized rib-like transverse structure across posterior margin of

sternite XIV.

Transverse Suture. Fine short vertical line extending dorsally from the ventral margin of the

carapace.

Trichobranchiate Gill. A gill in which the branches are fingerlike and project from a central

axis.

Uropod. Paired biramous appendage attached to the sixth abdominal somite, usually combining

with the telson to form a tailfan.

Ventral Costa. Ridge extending along the ventromesial margin of the ventrolateral lobule ot the

petasma (Fig. 4B).

Ventrolateral Lobule. Ventral part of the lateral lobe of the petasma (Fig. 4B).

Ventromedian Lobule. Lateral part of the median lobe of the petasma (Fig. 4B).

SYSTEMATIC SECTION

Superclass CRUSTACEA Pennant, 1777

Class MALACOSTRACA Latreille, 1806

Order DECAPODA Latreille, 1803

Suborder DENDROBRANCHI AT A Bate, 1888

Dendrobmnchiata Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 217. — Burkenroad, 1963a, Tulane

Stud. Geol., 2(1): 2. — GLAESSNER, 1969, Treat. Invert. Paleont., R, Arthropoda 4(2): R446. —

Burkenroad, 1981, Trans. San Diego Nat. Hist. Soc., 19(17): 251. — Bowman and Abele, 1983, Biol.

Crust., 1:21. — BURKENROAD, 1983, Crust. Issues, 1: 279. — FELGENHAUER and abele, 1983, Crust.

Issues, 1: 304. — Dall el ai, 1990, Adv. Mar. Biol., 27: 57.

Diagnosis. — Protocephalon consisting of ocular plate and epistomial region. Epistome

divided by membranous invagination. Lateral epistomial bars present anterior to labrum. Gastric

mill well developed, with strong median tooth and well developed lateral teeth. Third maxilliped

pediform, of 7 segments. First three pairs of pereopods usually chelate (except in some

Sergestoidea), third pair never unusually robust. Pleura of second abdominal somite not

overlapping those of first. Abdominal somites locked to each other by midlateral hinges, usually

exposed on anterior margin of second, third, fifth, and sixth somites, those of fourth very low

and hidden. Gills dendrobranchiate. First pair of pleopods in males with endopods considerably

modified, forming petasma; in females, much reduced or lacking. Eggs released directly into

water, not carried by females, (except in Lucifer, where eggs retained for short period on third

pereopods), hatching as nauplii or protozoeae.

Key to the Superfamilies of the Dendrobranchiata

1. Some thoracic somites with at least 3 branchiae on each side. At least

11 branchiae present on each side .PENAEOIDEA

— Never more than 2 branchiae per thoracic somite. Never more than 7 or

8 branchiae per side . SERGESTOIDEA

Superfamily PENAEOIDEA Rafinesque-Schmaltz, 1815

Penaeoidea Rafinesque-Schmaltz, 1815, Anal. Nat. Tabl. Univers, : 98. — Glaessner, 1969, Treat. Invert.

Paleont., R, Arthropoda, 4(2): R447. — BURKENROAD, 1983, Crust. Issues, 1: 280, 281, 285.

Diagnosis. — All five pairs of pereopods well developed. Pleurobranchia present at least on

somite IX (that of third maxilliped). Some somites with at least three branchiae on each side.

Total number of well developed gills on each side at least 11.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Includes five families: Aristeidae Wood-Mason, 1891; Benthesicymidae Wood-Mason, 1891;

Penaeidae Rafinesque-Schmaltz, 1815; Sicyoniidae Ortmann, 1898; Solenoceridae Wood-

Mason, 1891.

Key to the Families of the Superfamily PENAEOIDEA

1. Postorbital spine present. SOLENOCERIDAE

— Postorbital spine absent. 2

2. Integument rigid and stony; third to fifth pleopods uniramous, lacking

endopods . SICYONIIDAE

— Integument never stony; third to fifth pleopods biramous . 3

3. 1 or 2 (occasionally 3) rostral/postrostral teeth. BENTHESICYMIDAE

— More than 2 rostral/postrostral teeth. 4

4. Prosartema well developed .PENAEIDAE

— Prosartema reduced to setose boss . ARISTEIDAE

Family ARISTEIDAE Wood-Mason, 1891

Aristaeina Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 278.

Aristaeinae - ALCOCK, 1901, Descr. Cat. Indian Deep-Sea Crust., : 27. — RAMADAN, 1938. Scient. Rep. John

Murray Exped., 5(3): 36. — KUBO, 1949, J. Tokyo Coll. Fish., 36(1): 193.

Aristeinae - Bouvier, 1908a, Result. Camp, scient. Prince Albert I, 33: 6, 13. — Balss, 1957. Bronn’s Kl.

Ordn. Tierreichs. 5(1)7(12): 1516. — Crosnier, 1978, Faune Madagascar, 46: 14. — Burkenroad, 1983,

Crust. Issues, 1: 281, 286.

Placed on the Official List of Family-Group Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Bull. zool. Norn., 25(4/5): 141.

Aristeae - Bouvier, 1908a, Result. Camp, scient. Prince Albert I, 33: 6, 51. — Balss, 1957, Bronn's Kl. Ordn.

Tierreichs, 5(1)7(12): 1517.

Penaeidae - BARNARD, 1950, Ann. S. Afr. Mus., 38: 580 [part],

Aristeidae - CROSNIER, 1978, Faune Madagascar, 46: 14. — Perez Farfante, 1978, FAO Sp. Ident. Sh„ 6: 2.

— Burkenroad, 1983, Crust. Issues, 1: 281. — De Freitas, 1985, Investl. Rep. oceanogr. Res. Inst.,

Durban, 57: 3. — PEREZ FARFANTE, 1988, NOAA Tech. Rep. NMFS, 64: iii, 6. — SQUIRES, 1990, Can.

Bull. Fish. Aq. Sci., 221: 20.

Aristaeidae - GREY et al. , 1983, Guide Austr. Prawns, : 13, 14. — Dall etai, 1990, Adv. Mar. Biol., 27: 58.

Diagnosis. — Rostrum usually elongate in females, sexually dimorphic in several genera,

being elongate in females and juvenile males, short in adult males; usually with only 3 dorsal

rostral/postrostral teeth (more only in Aristaeomorpha)\ lacking ventral teeth. Carapace lacking

postorbital and pterygostomian spines; antennal and branchiostegal spines always present;

postantennal spine rarely present and hepatic spines often lacking; cervical and postcervical sulci

sometimes present, most often reaching dorsal midline, or almost absent and visible only

laterally. Third abdominal somite sometimes carinate, fourth to sixth somites always carinate.

Telson apically acute, bearing 3 or 4 pairs of movable lateral spines.

Eye with optic calathus bearing mesial tubercle; ocular scale and styliform projection lacking

Antennule with prosartema reduced to setose boss; flagella unequal, dorsal flagellum short,

flattened for most of its length, inserted proximally on third segment; ventral flagellum sexually

dimorphic in some genera. Palp of first maxilla consisting of single curved article. Exopods

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

present on all maxillipeds, present or absent on pereopods. Third through fifth pleopods

biramous. Pleurobranchia on somites IX to XIV; one arthrobranchia on somite VII, two on

somites VIII through XIII; podobranchia on second maxilliped through second or third

pereopods; epipod present on maxillipeds and first to third or fourth pereopods.

Petasma open; ventral costa usually projecting free for variable fraction of its length, often

extending as far as distal margin of dorsolateral lobule. Second pleopod of male bearing appendix

masculina and appendix interna, lacking distolateral projection. Thelycum open; sternite XIV

with deep concavity, sternite XIII with variously shaped shieldlike median protuberance.

Key to the Genera of the Family ARISTEIDAE

1. Three rostral/postrostral teeth present . 2

— More than 3 rostral/postrostral teeth present. Aristaeomorpha

2. Postantennal spine absent . 3

— Postantennal spine present . Parahepomadus

3. Hepatic spine absent . 4

— Hepatic spine present . Hepomadus

4. Exopods absent from first to fifth pereopods . 5

— Exopods present on first to fifth pereopods . 7

5. Distal movable meral spine present on first and second pereopods;

Scaphocerite in male not modified.. • • • .. 6

— Meral spine lacking on first and second pereopods; Scaphocerite in male

modified, distally elongate . Aristaeopsis

6. Postcervical sulcus present . Pseudaristeus

— Postcervical sulcus absent. Aristeus

7. Fourth to sixth abdominal somites dorsally carinate; Epipod on fourth

pereopod large . ®

— Third to sixth abdominal somites dorsally carinate; Epipod on fourth

pereopod rudimentary. Hemipenaeus

8. Carapace ridges and carinae weak; Podobranchia on third pereopod

rudimentary; Ventral antennular flagellum in male modified; Dactyl of third

maxilliped in male modified . Austropenaeus

— Carapace with ridges and carinae moderately strong to strong;

Podobranchia on third pereopod large; Ventral antennular flagellum in male

not modified; Dactyl of third maxilliped in male not modified.

. Plesiopenaeus

Genus Aristaeomorpha Wood-Mason, 1891

Figures 5-6

Penaeus - Risso, 1827, Hist. Nat. Prin. Prod. Europe Merid.. : 69 [part], — H. Milne Edwards, 1837, Hist. Nat.

Crust., 2: 418 [part].

Aristeus - Bate, 1881. Ann. Mag. nat. Hist., (5)8: 189; 1888, Rep. scient. Results Voy. Challenger, 24: 317

[part].

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Aristaeomorpha Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 286. — Anderson and Lindner, 1943, Trans.

Am. Fish. Soc., 73: 301. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 200. — Crosnier, 1978, Faune

Madagascar, 46: 52. — De Freitas, 1985, Investl. Rep. oceanogr. Res. Inst., Durban, 57: 15.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1804, Bull. zool. Nom., 25(4/5): 138.

Aristeomorpha - Senna, 1903, Boll. Soc. ent. ital., 34: 268. — Bouvier, 1908a, Result. Camp, scient. Prince

Albert I, 33: 52. — Ramadan, 1938, Scient. Rep. John Murray Exped., 5(3): 53. — Burkenroad, 1983,

Crust. Issues, 1: 281.

Fig. 5. — Aristaeomorpha foliacea, 9 52.5 mm cl, south of Soto la Marina, northeast Mexico. Lateral view.

Scale = 10 mm.

Type Species: By original designation, Aristeus rostridentatus Bate, 1881, Ann. Mag. nat.

Hist., (5)8: 189 [= Aristaeomorpha foliacea (Risso, 1827)].

Type Locality: Off Matuku, Fiji, 19°09'35"S, 179°41'50"E, 549 m.

Gender: Feminine.

Diagnosis. — Integument pubescent, with small scattered photophores on ventrum of

cephalothorax and abdomen, and on antennal scale and on lateral ramus of uropod, and on most

other appendages. Rostrum elongate in females and juvenile males, reaching well beyond

scaphocerite; short in adult males, not reaching third antennular peduncle segment, bearing 3-7

dorsal teeth along its entire length, followed by 2 postrostral teeth. Carapace with antennal and

branchiostegal spines on anterior margin; orbital, pterygostomian, and postantennal spines

absent; hepatic spine well developed; postrostral carina reaching about two-thirds of carapace;

cervical sulcus demarked ventrolaterally, becoming obsolete towards dorsal midline; postcervical

sulcus faint; branchiostegal carina sharp anteriorly; hepatic carina well marked, accompanying

shallow but well defined sulcus; branchiocardiac carina strong, almost reaching posterior margin

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

of carapace. First and second abdominal somites dorsally rounded, third to sixth somites dorsally

carinate, carina of third somite extending only along posterior two-thirds, almost indistinct, four

posterior somites ending in sharp spine. Telson bearing 4 pairs of movable lateral spines in

posterior half.

Fig. 6. — Aristaeomorpha foliacea: A, 9 52.5 mm cl, south of Soto la Marina, northeast Mexico. Thelycum. —

B-C, 3 40.0 mm cl, ibidem. B. Petasma, dorsal view. C, idem, ventral view. Scales = 1 mm.

ISABEL P£REZ FARFANTE & BRIAN KENSLEY

Eye with cornea well developed, optic calathus bearing minute mesial tubercle mesio-

distally. Dorsal antennular flagellum short, flattened for most of its length; ventral flagellum

elongate, slender, terete, not modified in male. Scaphocerite with lateral marginal ridge ending

in short spine at about three-fourths length of lamella. Pereopods lacking exopods; ischial

and meral spines absent; fourth and fifth pereopods slender. Third pereopod bearing

podobranchia.

Petasma with dorsolateral lobule roughly trapezoidal, conspicuously surpassing median lobe-

ventral costa as long as dorsolateral lobule, attached along almost entire length, with tip turning

slightly mesially. Thelycum open, with shieldlike median protuberance on sternite XIII directed

anteroventrally; sternite XIV bearing short rounded median prominence.

The formation and structure of the complex spermatophore of A. foliacea were described bv

Titmcci ( 1 QQ*7\ J

Distribution of branchiae, epipods, and exopods in the genus Aristaeomorpha.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

s+1

Podobranchiae

Epipods

Exopods

m SP f CI ^ S - — Aristaeomorpha foliacea (Risso, 1827). Western Atlantic Ocean, Massachusetts to

Florida, Gulf of Mexico, Caribbean Sea to Venezuela; Eastern Atlantic Ocean. Bay of Biscay to

Western Sahara; Azores; Madeira; Canary Islands; Mediterranean; off east coast of South Africa-

Mozambique, East Africa; Madagascar; Reunion; Maidive Islands; Sri Lanka; Indonesia!

Philippines Taiwan; Japan, Western Australia, New South Wales, Victoria, Australia; New

Caledonia; New Zealand; Wallis and Futuna Islands; Fiji.

Aristaeomorpha woodmasoni Caiman, 1925. Arabian Sea; Bay of Bengal; Andaman Sea.

Genus Aristaeopsis Wood-Mason, 1891

Figures 7-8

/> T88''[pJt] HNSON ’ 186? ’ Pr ° C ' Z0 ° L S0C ‘ L ° nd -’ ' 867: 897 [pan] ' - BATE ’ 1881 ’ Ann - Ma §- na '- Hist., (5)8:

Aristeus - Miers, 1878, Proc. zool. Soc. Lond., 1878: 308 [part]

AriS SZ^ d '^T’ T’ Ann ' Ma f Q (6)8: 281 - B °™e r , 1908a - R-suI,. Camp, scent.

Faune Madagascar, 46: 86 . ^ ,936b ’ BuU - B,n S ham oceano S r - Co “- 5(2): 94. - Crosn.er, 1978,

PleS l 909 n UL Mns N ;^ 95 ’ 7 M T» MUS - fr^r 001 - Harv -' 18: 199 [part| - - A - M,lne Edwards and Bouvier,

5?2°) 9, 94 [narH S P ‘ Z °° ' tifo ’c 7 ' 2 °° [paitJ - _ Burkenr °™- 1936b, Bull. Bingham occanogr. Coll,

Ann S Afr tiluT o ’ n' 6 "'' RCP ' S? MUrf3y Exped - 5 < 3 ) : 49 [part]. - Barnard, 1950,

Faune Madagascar - 46:85 [par,] - - Squ,res - l99 °-

Arises (Plesiopenaeus) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 35 [part]

Ansiaeus Anstaeopsu) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 40 [part] '

Ansteopsis - De Man, 1911, Siboga Exped., 39a: 6. lF J

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Fig. 7. — Aristaeopsis edwardsiana, 9 54 mm cl, Guadeloupe Passage, Caribbean Sea. Lateral view.

Scale = 10 mm.

Type Species: By original designation, Penaeus edwardsianus Johnson, 1867, Proc. zool.

Soc. Lond., 1867: 897.

Type Locality: Off Madeira, Northeast Atlantic Ocean.

Gender: Feminine.

Diagnosis. — Integument glabrous. Rostrum moderately elongate in females and juvenile

males, reaching beyond apex of scaphocerite; somewhat shorter in adult males, not reaching third

antennular peduncle segment, bearing 2 basal teeth followed by single postrostral tooth. Carapace

with antennal and branchiostegal spines on anterior margin, each supported by strong carina;

orbital, postantennal, and hepatic spines absent. Cervical sulcus ventrally deep, rather weak

dorsally, almost reaching dorsal midline; postcervical sulcus absent; postrostral carina reaching

posteriorly about three-fourths length of carapace; gastro-orbital, hepatic, and branchiocardiac

carina strong. First abdominal somite dorsally rounded; second somite with barely discernible

rounded middorsal carina in posterior half; third to sixth somites with sharp carina, ending

posteriorly in short spine. Telson pointed, with 4 pairs of small lateral movable spines in

posterior half.

Eye with cornea slightly flattened; calathus moderately short, with strong mesial tubercle

distally. Dorsal antennular flagellum short, flattened; ventral antennular flagellum elongate, terete,

not modified in adult male. Scaphocerite in adult female with blade apically acute-angled, lateral

marginal ridge ending in short spine at about four-fifths length; in adult male, lamella of

scaphocerite produced distally into elongate slender structure, ovate in cross-section, subequal in

length to proximal broad part; in female, lateral ridge falling short of distal margin of lamella.

Pereopods lacking exopods; first and second pereopods lacking movable distal meral and ischial

spines. Fourth and fifth pereopods more slender, especially distally, than preceding pereopods.

Third pereopod bearing podobranchia.

ISABEL PgREZ FARFANTE & BRIAN KENSLEY

Petasma with dorsolateral lobule reaching as far as median lobe; ventral costa slightly

overreaching distal margin of dorsolateral lobule, attached along entire length to adjacent flexible

part of ventrolateral lobule; its tip becoming mesially-directed conspicuous hook. Thelycum

open, with cup-like hollow formed by anteriorly acute median protuberance on sternite XIII and

strongly raised, roughly rectangular, anteriorly rounded median prominence flanked anteriorly by

strong rounded lateral flaps on stemite XIV.

Distribution of branchiae, epipods, and exopods in the genus Aristaeopsis.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

s+1

Podobranchiae

Epipods

Exopods

Fig. 8. — Aristaeopsis edwardsiana: A, 2 72 mm cl, Gulf of Mexico.

Passage, Caribbean Sea. Petasma, dorsal view. Scales = 1 mm.

Thelycum. — B, cJ 54 mm

cl, Guadeloupe

Remarks. — The genus Aristaeopsis was originally created for the species Penaeus

197RWW? f*’ I >Ut W l S 3ter s y non >' mised Wlth Plesiopenaeus (Burkenroad, 1936b; Crosnier

wolf ’ f however, separate this species from typical Plesiopenaeus (P armatus) as

Z a val[d°Snus^T^Tr7 ,/, ' /M ' V and thus necessitate the recognition of Aristaeopsis

as a valid genus. These features are: the absence of exopods on the first to fifth pereoDods- the

presence of a blunt, rounded middorsal carina on the second abdominal somite; the presence of a

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

sharp middorsal carina on the third abdominal somite; the apparently unique modification of the

antennal scaphocerite in the adult male; the absence of a distal meral spine on the first and second

pereopods.

Species. —Aristaeopsis edwardsiana (Johnson, 1867). Western Atlantic Ocean from Grand

Banks to Gulf of Mexico, Caribbean Sea to French Guiana; Bermuda; eastern Atlantic Ocean

from Azores, Madeira, Canary Islands, Portugal, Morocco, Western Sahara, to South Africa;

East Coast of Africa; Madagascar; Arabian Sea; central Indian Ocean; Bay of Bengal; Andaman

Sea; Indonesia; Japan; South China Sea; New South Wales, Australia; Wallis and Futuna

Islands.

Genus Aristeus Duvernoy, 1840

Figures 9-10

Penaeus - Risso, 1816, Hist. nat. Crust., Nice, : 96.

Aristeus Duvernoy, 1840, C. r. hebd. Seanc. Acad. Sci., Paris, 11: 217. — Bate, 1881, Ann. Mag. nat. Hist.,

(5)8: 171, 187; 1888, Rep. scient. Results Voy. Challenger, 24: 228, 240, 309. — Senna, 1903, Boll. Soc.

ent. ital., 34: 261. — Bouvier, 1908a, Result. Camp, scient. Prince Albert 1, 33: 69. — De Man, 1911,

Siboga Exped., 39a: 27. — Ramadan, 1938, Scient. Rep. John Murray Exped., 5(3): 36. — Crosnier, 1978,

Faune Madagascar, 46: 60. — Burkenroad, 1983, Crust. Issues, 1: 281. — De Freitas, 1985, Investl. Rep.

oceanogr. Res. Inst., Durban, 57: 3.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 805, Bull. zool. Nom., 25(4/5): 138.

Aristaeus - Faxon, 1895, Mem. Mus. comp. Zool. Harv., 18: 194, 197. — Alcock, 1901, Descr. Cat. Indian

Deep-Sea Crust., : 27. — Anderson and Lindner, 1943, Trans. Am. Fish. Soc.. 73: 300. — Kubo, 1949, J. Tokyo

Coll. Fish., 36(1): 194.

Aristaeus (Aristaeus) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 29.

Type Species: By original designation, Penaeus antennatus Risso, 1816, Hist. nat. Crust.,

Nice: 96.

Type Locality: Mediterranean Sea off Nice.

Gender: Masculine.

Diagnosis. — Integument glabrous or pubescent; photophores present on pereopods.

Rostrum elongate in females and juvenile males, reaching well beyond antennular peduncle; short

in adult males, not reaching end of antennular peduncle; bearing 2 dorsal teeth near base followed

by single postrostral tooth. Carapace with antennal and branchiostegal spines on anterior margin,

both supported by more or less well marked short carina; orbital, postantennal, and hepatic

spines absent; postrostral carina variable, reaching as far as midlength of carapace in some

species; cervical sulcus indistinct or only ventrolaterally demarked; hepatic sulcus often taint or

indistinct; branchiocardiac sulcus well marked; branchiocardiac carina low. First abdominal

somite dorsally rounded; second and third somites dorsally rounded or with very faint rounded

carina; fourth to sixth somites dorsally carinate, carina of fourth somite extending only along

posterior two-thirds. Telson bearing 4 pairs of movable lateral spines; apex acute, slender.

Eye with well developed cornea; optic calathus bearing large mesiodistal tubercle. Dorsal

antennular flagellum short, flattened for most of its length; ventral antennular flagellum elongate,

slender, terete, with proximal sinuosity in male. Scaphocerite lamella distally broadly rounded,

lateral marginal ridge with spine at about distal four-fifths of length. Pereopods lacking exopods;

first and second pereopods, sometimes third, with movable meral spine; fourth and fifth

pereopods slender. Third to fifth pereopod lacking podobranchia.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Fig. 9.

Aristeus antennatus , $ 50 mm cl, Indian Ocean off South Africa. Lateral view. Scale = 10 mir

ven^rnm^i „ W 'i h K b i° ad ’ y rounded dorsolateral lobule, slightly to considerably surpassing

? r ed an 1 t obul ^ ; co u sta unattached to adjacent flexible part for variable fraction of it!

sSe XmTtem^P XTV i nt ° Jl°° k ' Thel f Um ,°P en ’ with shieldlike median protuberance on

stemite XIII, stemite XIV broadly rectangular, slightly hollowed

descriwTfSsX™ SpCrmat0ph0re ofA

Distribution of branchiae, epipods, and exopods in the genus Aristeus.

Somite

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

vn vm ix x

(Mxpl) (Mxp2) (Mxp3) (PI)

1 1

1 s+1 2 2

1 1 1

1 1 1 1

1 1 1 -

(P2)

(P3)

(P4)

XIV

(P5)

Aristeus alcocki Ramadan, 1938. Gulf of Aden; Arabian Sea; India; Bay of

Species.

Bengal.

Aristeus antennatus (Risso, 1816). Eastern Atlantic Ocean from Portugal to Cane VerHe

Rtnt“ve ^ ^ Mozambi ^ Zanzibar; Madagascar;

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Aristeus antillensis A. Milne Edwards and Bouvier, 1909. Delaware to Florida; Gulf of

Mexico; Caribbean Sea to French Guiana.

Aristeus mabahissae Ramadan, 1938. Madagascar; Maidive Islands; Saya de Malha Bank;

Indonesia; Japan; New Caledonia; Wallis and Futuna Islands.

Aristeus pallidicauda Komai, 1993. Japan.

Aristeus semidentatus Bate, 1881. Madagascar; Arabian Sea; India; Malaysia; Indonesia; New

South Wales, Australia; Kermadec Islands; Hawaii.

Aristeus varidens Holthuis, 1952a. Eastern Atlantic Ocean from Western Sahara to Namibia.

Aristeus virilis (Bate, 1881). East coast of South Africa; Mozambique; Zanzibar; Madagascar;

Reunion; Andaman Islands; Indonesia; Philippines; Taiwan; Japan; New South Wales, Australia;

Fiji; New Caledonia; New Hebrides.

Fig 10. — Aristeus antenna,us: A, J 33 mm cl, off Sicily, Mediterranean. Petasma, ventral view. Scale

1 mm . — B, 2 50 mm cl, Indian Ocean off South Africa. Thelycum. Scale - 5 mm.

Genus Austropenaeus new genus

Figures 11-12

Plesiopenaeus non Bate - Barnard, 1947, Ann. Mag. nat. Hist., (11)13: 383; 1950, Ann. S. Afr. Mus., 38: 622

[part]. — CROSNIER, 1978, Faune Madagascar, 46: 9, 88-90.

Type Species: By present designation, Plesiopenaeus nitidus Barnard, 1947, Ann. Mag. nat.

Hist., (11)13: 383.

Type Locality: Off Cape Point, South Africa, 870-1153 m.

Gender: Masculine.

Diagnosis - Adult males markedly smaller than adult females. Integument glabrous.

Rostrum similar in males and females, up to 1.5 times longer than carapace in adults, bearing

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

2 basal teeth followed by single postrostral tooth. Carapace with antennal and branchiostegal

spines on anterior margin, latter with fairly strong supporting carina; orbital, postantennal, and

hepatic spines absent; postrostral carina not quite reaching midlength of carapace; cervical sulcus

only distinguishable ventrolaterally; hepatic sulcus weak; branchiocardiac carina weak. First and

second abdominal somites dorsally rounded; third somite posteriorly weakly carinate, fourth to

sixth somites strongly carinate, third to sixth somites each ending in small middorsal spine.

Telson acute, with 3 pairs of movable lateral spines in posterior half.

Ridge off Namibia. Lateral view. Scale = 10 mm.

Fig. 11. — Austropenaeus nitidus , $ 28 mm cl, Walvis

Eye with cornea slightly flattened; calathus with strong mesiodistal tubercle. Dorsal antennular

flagellum short, reaching apex of scaphocente, flattened for most of its length; ventral antennular

flagellum elongate, terete, in male having proximal sinuosity formed by several dilated articles at

leve of apex of dorsal antennular flagellum. Scaphocerite with lateral marginal ridge ending in

short spme at about four-fifths length of blade. Third maxilliped of adult male with dactyl shorter

and basally thicker than in adult female. First to fifth pereopods each having rudimentary exopod-

first to third pereopods each with distal movable meral spine; fourth and fifth pereopods more

slender than preceding ones. Third pereopod bearing rudimentary podobranchia.

Distribution of branchiae, epipods, and exopods in the genus Austropenaeus.

Somite

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

vn VEI IX

(Mxpl) (Mxp2) (Mxp3)

s s+1 2

1 1

1 1 1

(PI)

(P2)

XII

(P3)

(P4)

XIV

(P5)

Petasma with dorsolateral lobule produced into elliptical

surpassing ventromedian lobule; ventral costa distally expanded

narrow process, conspicuously

into subcircular lobe. Thelycum

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

open, with apically pointed shieldlike median protuberance on sternite XIII; sternite XIV with

low anteriorly pointed median prominence and paired short anterolateral ridges.

Fig. 12. - Austropenaeus nitidus-. A, d 22 mm cl. Walvis Ridge off Namibia. Petasma, dorsal view

B, 2 29 mm cl, Walvis Ridge off Namibia. Thelycum. Scales = 1 mm.

Remarks. — The anomalous position of Plesiopenaeus nitidus Barnard within the definition

of typical Plesiopenaeus , was pointed out by Crosnier (1978: 88 table 11). At least two

characters used in the latter table, however, were erroneous, due to the lack of material on which

to base conclusions. Examination of specimens reveals at least four

this species both from Plesiopenaeus and Aristaeopsis: the podobranch of the third pereopod is

reduced; the ventral antennular flagellum in the male is modified; the third maxilhped in the adult

male is modified; the carinae of the carapace are very feebly developed. In addition, the first to

third pereopods each bear a distal movable meral spine. , ,,

The modification of the ventral antennular flagellum in the male of A. nitidus resembles the

condition found in the genus Aristeus. The latter, however,

evestalk an unflattened cornea, lacks exopods on the pereopods, a podobranch on the third

pereopod, and an epipod on the fourth. Because of these differences, A. nitidus cannot be

included in the genus Aristeus.

Species. —Austropenaeus nitidus (Barnard, 1947) South-east Atlantic Ocean off Cape of

Good Hope; South Atlantic Ocean, Valdivia Bank; off Natal, South Africa, off St. Paul and

Amsterdam Islands; Bass Strait, Australia.

Genus Hemipenaeus Bate, 1881

Figures 13-14

Hemipenaeus Bate, 1881, Ann. Mag. nat. Hist., (5)8: 171 186; 1888, Rep. Results Vff-CMtajgO*

299. — De Man, 1911, Siboga Exped., 39a: 23 [part], — Burkenroad, 1936b, Bull. Bingham oceanogr.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Coll., 5(2): 90. — Ramadan, 1938, Scient. Rep. John Murray Exped., 5(3): 47 [part], — Anderson and

Lindner, 1943, Trans. Am. Fish. Soc., 73: 300. — Roberts and Pequegnat, 1970, Texas A&M Univ.

Oceanogr. Stud., 1: 43. — Crosnier, 1978, Faune Madagascar, 46: 74.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1813, Bull. zool. Norn., 25(4/5): 139.

Hemipeneus - Faxon, 1895, Mem. Mus. comp. Zool. Harv., 18: 198. — Burkenroad, 1983, Crust Issues

1:282.

Aristaeus (Hemipeneus ) - Alcock, 1901, Descr. Cat. Indian Deep- Sea Crust.,: 31 [part],

Hemypenaeus - Kikuchi and Nemoto, 1986, Bull, biogeogr. Soc. Japan, 41(7): 52.

Type Species: By subsequent designation, by Faxon, 1895, Hemipenaeus spinidorsalis Bate,

1881, Ann. Mag. nat. Hist., (5)8: 186.

Type Locality: South Atlantic Ocean off Tristan da Cunha, 35°41'S, 20°55'W, 3477 m.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum either moderately elongate in females and short

in males, or short in both sexes, bearing 2, occasionally 3 basal dorsal teeth, followed by single

postrostral tooth. Carapace with antennal and branchiostegal spines, latter continuous with sharp

carina; orbital, postantennal, pterygostomian, and hepatic spines absent; postrostral carina

reaching to midlength or almost to posterior margin of carapace; cervical sulcus either almost

reaching middorsal line, or only demarked ventrally, if at all; postcervical sulcus present or

absent; hepatic sulcus strong; branchiostegal carina well marked; branchiocardiac carina and

sulcus strong. First and second abdominal somites dorsally rounded, third to sixth somites

dorsally carinate, third somite armed with spiniform process arising from posterior third of

F 'Vi - Hemi P en “ e “ s spinidorsalis: A, 9 65.5 mm cl, off Galapagos, Ecuador. Lateral view.

B, o 44 mm cl., ibidem. Rostrum. Scale = 10 mm.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

carina; sixth somite with small middorsal spine on posterior margin. Telson bearing 4 pairs of

small movable lateral spines in about posterior half; apex acute.

Eye with cornea well developed, somewhat dorsoventrally flattened; calathus with strong

tubercle at about midpoint of mesial margin. Dorsal antennular flagellum short, flattened for most

of its length; ventral flagellum elongate, slender, terete, not modified in mature male.

Scaphocerite with apex of blade rounded, lateral marginal ridge ending in spine at about distal

fourth of length of lamella. First to fifth pereopods each with reduced exopod; men of pereopods

lacking distal movable spine; fourth and fifth pereopods very slender. Third pereopod bearing

podobranchia; rudimentary epipod on fourth pereopod.

FlG . 14. - Hemipenaeus spinidorsalis: A, 3 44 mm cl, off Galapagos, Ecuador. Petasma, ventral v.ew of right

half. Scale = 1 mm. — B, 9 65.5 mm cl, off Galapagos, Ecuador. Thelycum. Scale - 5 mm.

Petasma with distal margin of dorsolateral lobule oblique, mesially 1 reachin as far as

ventromedian lobule; ventral costa free distally for about half or one-third of its lei }g th -Thclycun

open, with broad anteriorly acute median protuberance on stermte XIII, stermte XIV broad,

short, roughly rectangular.

Distribution of branchiae, epipods, and exopods in the genus Hemipenaeus.

Somite

(Mxpl)

vrn

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

s+1

Podobranchiae

Epipods

Exopods

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Species. — Hemipenaeus carpenteri Wood-Mason, 1891. Eastern Pacific Ocean off Gulf of

Panama; Galapagos Islands; western Atlantic Ocean off Bahamas; Gulf of Mexico; Caribbean;

Madagascar; Reunion; Arabian Sea; central Indian Ocean; Bay of Bengal; Java Sea; Japan; Wallis

and Futuna Islands.

Hemipenaeus spinidorsalis Bate, 1881. Eastern Pacific Ocean off Gulf of Panama; Galapagos

Islands; South Atlantic Ocean off Tristan da Cunha; Madagascar; central Indian Ocean;

Philippines; north-west Pacific Ocean.

Genus Hepomadus Bate, 1881

Figures 15-16

Hepomadus Bate, 1881, Ann. Mag. nat. Hist., (5)8: 171, 189; 1888, Rep. scient. Results Voy. Challenger, 24:

319. — Bouvier, 1908a, Rdsult. Camp, scient. Prince Albert I, 33: 56. — A. Milne Edwards and Bouvier,

1909, Mem. Mus. comp. Zool. Harv., 27: 194. — Burkenroad, 1936b, Bull. Bingham oceanogr. Coll., 5(2):

86. — Ramadan, 1938, Scient. Rep. John Murray Exped., 5(3): 55. — Anderson and Lindner, 1943, Trans.

Am. Fish. Soc., 73: 300. — P£rez Farfante, 1973, Fish. Bull. U.S., 71(2): 441. — Crosnier, 1978, Faune

Madagascar, 46: 47. — Burkenroad, 1983, Crust. Issues, 1: 281.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1814, Bull. zool. Norn., 25(4/5): 139.

Aristaeus (Hepomadus) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 42.

Fig. 15. — Hepomadus tener, 9 51 mm cl, off Mobile, Alabama. Lateral view. Scale = 10 mm (From

PfiREZ Farfante, 1973).

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Type Species: By subsequent designation, by Fowler, 1912, Hepomadus glacialis Bate,

1881, Ann. Mag. nat. Hist., (5)8: 190.

Type Locality: Off Yokohama, 34°37'N, 140°32'E, 3431 m.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostral length in young females and apparently, in all

males, not exceeding 0.7 times that of carapace, in larger females length increasing to as much as

1.25 carapace length, reaching beyond distal end of antennular peduncle by half its length,

bearing 2 dorsal teeth near base, followed by single postrostral tooth. Carapace with postrostral

carina almost reaching posterior margin; antennal and marginal branchiostegal spines present;

orbital postantennal, and pterygostomian spines absent; hepatic spine present; cervical sulcus

well marked, reaching middorsal line; postcervical sulcus faint; branchiostegal carina anteriorly

sharp, posteriorly weak; hepatic sulcus distinct; branchiocardiac carina strong, accompanying

sulcus deep. First and second abdominal somites dorsally rounded; third to sixth somites dorsally

carinate, third somite carinate in posterior two-thirds, ending posteriorly in spine. Telson with

4 pairs of lateral movable spines; apex slender, acute. .

Eye with cornea well developed; calathus with proximal tubercle on mesial margin. Dorsal

antennular flagellum short, flattened for most of its length; ventral flagellum elongate, slender

terete not modified in mature male. Scaphocerite blade distally rounded, rigid lateral margin with

tooth’at distal 5/6 of length. First to fifth pereopods each with exopod reduced or absent; merus

of first and second pereopods with distal movable spine; fourth and fifth pereopods slender.

Third pereopod lacking podobranchia; somite XIV lacking arthrobranchia.

Fig. 16. — Hepomadus tener. A-B. <J 47 mm cl, southeast of Cabo Rojo, Veracruz, Mexico. A Petasma dorsal

view. B, Idem , ventral view. - C, $ 47 mm cl, off Back Bay, Virginia. Thelycum. Scales = 1 mm.

(All drawings from PCrez Farfante, 1973).

Petasma with dorsomedian lobule distally flexed laterally; dorsolateral lobule reaching or

barely surpassing median lobe; about distal half of ventral costa free from adjacent flexible part of

ventrolateral lobule, its tip curving laterally, forming sinuous projection. Thelycum open, with

large roughly lanceolate median protuberance on stemite XIII; stermte XIV subrectangular,

weakly convex.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Distribution of branchiae, epipods, and exopods in the genus Hepomadus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

r+1

Podobranchiae

Epipods

Exopods

Species. — Hepomadus glacialis Bate, 1881. Japan.

Hepomadus inermis Bate, 1881. South-central Pacific Ocean.

Hepomadus tener Smith, 1884. Western Atlantic Ocean; Gulf of Mexico; eastern Caribbean

Sea; Sargasso Sea; Azores, north-eastern Atlantic Ocean; Cape Verde Islands, Zanzibar;

Madagascar; Reunion; central Indian Ocean; Bay of Bengal; Wallis and Futuna Islands.

Genus Parahepomadus Crosnier, 1978

Figures 17-18

Parahepomadus Crosnier, 1978, Faune Madagascar, 46: 47.

F,G 'J 7 o - P “ rahep °" ,ad us vaubani: A. <J 58 mm cl. 16 km northeast of North Island, Indonesia, lateral view. -

B, V 13.5 mm cl, rosiral region. Scale for both drawings = 10 mm. (B from Crosnier, 1978).

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Type Species: By original designation, Parahepomadus vaubani Crosnier, 1978, Faune

Madagascar, 46: 48, figs 20-22.

Type Locality: Northwest and west coast of Madagascar, 880-1200 m.

Gender: Masculine.

Diagnosis. — Integument pubescent. Rostrum elongate in females and young males, reaching

well beyond antennular peduncle; relatively short in adult males, reaching distal end of second

segment of antennular peduncle; armed with 2 dorsal teeth at base, followed by single postrostral

tooth. Carapace with antennal, postantennal, and branchiostegal spines, latter supported by

elongate carina. Orbital, hepatic, and pterygostomian spines absent. Postrostral carina reaching to

about midlength of carapace. Cervical sulcus only ventrally visible; hepatic sulcus well marked,

continuous with branchiocardiac sulcus. First to third abdominal somites dorsally rounded,

fourth to sixth somites dorsally carinate. Telson apically acute, with 4 pairs of movable spines in

posterior fourth. . , , . , „ ,

Eye with well developed cornea; calathus with small conical tubercle on mesial margin. Dorsal

antennular flagellum short, flattened for most of its length; ventral flagellum elongate, filiform,

terete, with distinct proximal sinuosity formed by several dilated articles in male. Scaphocente

with lateral marginal ridge ending in short spine at about five-sixths length of blade; distal margin

of latter oblique-truncate. First to fifth pereopods lacking exopods; first and second pereopods

with subdistal movable spine on merus; fourth and fifth pereopods more slender than preceding

ones. Third pereopod lacking podobranchia.

Fig. 18. - Parahepomadus vaubani: A, <J 55 mm cl, off Madagascar. Petasma, ventral view of right half. Scale -

5 mm- _ b, 9 75.5 mm cl, off Madagascar. Thelycum. Scale = 10 mm. (Both drawings from Crosnier,

1978).

Petasma with dorsolateral lobule distally rounded, slightly surpassing median lobe, ventral

costa free from adjacent flexible part of ventrolateral lobule for one-third of its length. Thelycum

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

open, with large lanceolate median protuberance on sternite XIII; sternite XIV anteriorly with

narrow rugose area, broad posterior area flattened.

Distribution of branchiae, epipods, and exopods in the genus Parahepomadus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(Pi)

(P2)

(P3)

(P4)

xrv

(P5)

Pleurobranchiae

Arthrobranchiae

s+1

Podobranchiae

Epipods

Exopods

Species. — Parahepomadus vaubani Crosnier, 1978. Off Madagascar; Indonesia;

Philippines.

Genus Plesiopenaeus Bate, 1881

Figures 19-20

Aristeus - Bate, 1881, Ann. Mag. nat. Hist., (5)8: 187 [part]; 1888, Rep. scient. Results Voy. Challenger, 24-

309 [part].

Fig. 19. — Plesiopenaeus armatus , 2

Scale = 10 mm.

78 mm cl, North Atlantic Ocean off North Carolina. Lateral view

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Aristaeus - Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 278 [part].

Plesiopenaeus Bate, 1881, Ann. Mag. nat. Hist., (5)8: 188. — Bouvier, 1908a, Result. Camp, scient. Prince

Albert I, 33: 63 [part]. — Burkenroad, 1936b, Bull. Bingham oceanogr. Coll., 5(2): 94. — Ramadan, 1938,

Scient. Rep. John Murray Exped., 5(3): 49 [part], — Anderson and Lindner, 1943, Trans. Am. Fish. Soc. 73:

301. — Barnard, 1950, Ann. S. Afr. Mus., 38: 621. — Crosnier, 1978, Faune Madagascar, 46: 85. — De

Freitas, 1985, Investl. Rep. oceanogr. Res. Inst., Durban, 57: 20. — Squires, 1990, Can. Bull. Fish. Aq.

Sci., 221: 41.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1823, Bull. zool. Nom., 25(4/5): 139.

Plesiopeneus - Faxon, 1895, Mem. Mus. comp. Zool. Harv., 18: 199. — Burkenroad, 1983, Crust. Issues,

1: 281.

Aristeopsis - Bouvier, 1908a, Result. Camp, scient. Prince Albert I, 33: 61. — A. Milne Edwards and Bouvier,

1909, Mem. Mus. comp. Zool. Harv., 27: 197.

Type Species: By subsequent designation, by Faxon, 1895, Aristeus armatus Bate, 1881,

Ann. Mag. nat. Hist., (5)8: 188.

Type Locality not restricted, seven syntypes from following localities: Off Buenos Aires,

Argentina, 35°39'S, 50°47'W, 3477 m; South Atlantic Ocean off Tristan da Cunha, 35°41'S,

20°55'W, 3477 m; Torres Strait, 12°08'S, 145°10'E, 2562 m; Off Philippines, 5°47'N,

124°01'E, 3750 m; Off Yokohama, Japan, 34°37'N, 140°32’E, 3430 m; Central North Pacific

Ocean, 36°10'N, 178°00'E, 3750 m; Off Low Archipelago, Pacific Ocean, 13°28'S, 149°30'W,

4300 m.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum elongate, reaching beyond antennular peduncle

by as much as one and one-third its length in adult females, short in adult males, bearing 2 basal

dorsal teeth, followed by single postrostral tooth. Carapace with antennal and branchiostegal

spines present on anterior margin; orbital, postantennal, pterygostomian, and hepatic spines

absent. Cervical sulcus either almost reaching middorsal line, or only demarked laterally; gastro-

orbital, antennal, hepatic, and branchiocardiac carinae well marked; postrostral carina reaching at

least to midlength of carapace. First and second abdominal somites dorsally rounded; posterior

half of third abdominal somite and fourth to sixth somites dorsally carinate. Telson with 4 pairs

of movable lateral spines in posterior half; apex acute.

Eye with cornea slightly flattened; calathus moderately elongate, with strong tubercle at about

midlength of mesial margin. Dorsal antennular flagellum flattened for most of its length; ventral

antennular flagellum elongate, slender, terete, not modified in adult male. Scaphocerite with

lateral marginal ridge ending in short spine at about distal five-sixths length of lamella. First to

fifth pereopods each with rudimentary exopod; first pereopod, or first and second pereopods

with distal movable meral spine. Fourth and fifth pereopods more slender than preceding ones.

Third pereopod with well developed podobranchia; large epipod on fourth pereopod.

Distribution of branchiae, epipods, and exopods in the genus Plesiopenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

Xffl

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

s+1

Podobranchiae

Epipods

Exopods

ISABEL P£REZ FARFANTE & BRIAN KENSLEY

Petasma with distal margin of dorsolateral lobule oblique, mesially reaching as far as median

lobe; ventral costa free distally, its rounded or subacute apical part directed laterally. Thelycum

open, with shield-like median protuberance on stemite XIII anteriorly acute; stemite XIV roughly

rectangular.

Fig. 20. — Plesiopenaeus armatus: A, <J 74 mm cl, North Atlantic Ocean off North Carolina. Petasma. — B, $

78 mm cl, North Atlantic Ocean off North Carolina. Thelycum. Scales = 5 mm.

Species. — Plesiopenaeus armatus (Bate, 1881). Off north-west coast of U.S.A.; Western

Atlantic Ocean; Gulf of Mexico; eastern Atlantic Ocean off Azores; Madeira; Canary Islands-

Cape Verde Islands; Tristan da Cunha; Zanzibar; Madagascar; Maidive Islands; Chagos

Archipelago; Bay of Bengal; Philippines; Japan; north-west Pacific Ocean; Tuamotu Islands-

Wallis and Futuna Islands; Hawaii.

Plesiopenaeus coruscans (Wood-Mason, 1891). Gulf of Mexico; Bahamas; Madagascar-

Arabian Sea; Bay of Bengal; Andaman Islands.

Genus Pseudaristeus Crosnier, 1978

Figures 21-23

Hemipenaeus - Baw, 1881, Ann. Mag. nat. Hist., (5)8: 186 [part], - Ramadan, 1938, Scient. Rep. John Murray

bxped., 5(3): 47 [part]. J

Aristaeus - Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 278 [part].

Aristaeus (Hemipeneus) - Alcock, 1901, Descr. Cat. Indian Deep- Sea Crust.,: 31 [part].

Pseudansteus _Crosnier ,1978, Faune Madagascar, 46: 81. — De Freitas, 1985, Investl. Rep. oceanogr. Res. Inst

Durban, 57: 1 1. — P£rez Farfante, 1987, Fish. Bull., U. S., 85(2): 312. 6

Type Species: By original designation, Aristeus crassipes Wood-Mason, 1891, Ann. Mag.

nat. riist., ( 0 ) 8 ; 281.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Type Locality: Andaman Sea, 741 m.

Gender: Masculine.

Diagnosis. — Integument pubescent. Rostrum elongate in juvenile males and adult females,

reaching well beyond scaphocerite; fairly short in most adult males, not reaching apex of

scaphocerite; bearing 2 dorsal basal teeth, followed by single postrostral tooth. Carapace with

antennal and marginal branchiostegal spines; orbital, postantennal, pterygostomian, and hepatic

spines absent; postrostral carina, hepatic sulcus and cervical and postcervical sulci well

demarked, latter reaching or almost reaching middorsal line; branchiostegal-hepatic carina strong;

branchiocardiac carina and sulcus well marked. First to third abdominal somites dorsally

rounded, fourth to sixth somites dorsally carinate; fourth somite carinate only in posterior two-

thirds. Telson with slender acute apex, with 4 pairs of movable lateral spines in posterior half.

Eye with cornea well developed, dorsoventrally flattened; calathus with small tubercle on

mesial margin. Dorsal antennular flagellum short, flattened for most of its length, ventral

Fig. 21. — Pseudaristeus kathleenae , holotype, 6 23.5 mm cl, Sulawesi, Indonesia. Lateral view. Scale = 6 mm.

(From P£rez Farfante, 1987).

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

flagellum elongate terete, with proximal sinuosity in males of P. gracilis only. Scaphocerite with

lamella distal to lateral spine broadly rounded. First to fifth pereopods lacking exopods- first and

second pereopods with subdistal movable spine on meri; fourth and fifth pereopods more slender

than preceding ones. Third pereopod lacking podobranchia.

F,G »L A 5iS *"*?■* h0l0 Wt’ * 23 5 * Sulawesi. Monesia: A, Petasma, dorsolateral

nlmn!]? ! r| B j W ?' Venra view - C ’ Appendlx masculina, appendix interna, and basal sclerite of

Fir P 2? n p 2en p° P0 5 d ° rSa V ' eW 3nd Ventral View - Scales = 1 mm - (From P£REZ Farfante, 1987).

F'o. 22 D-E. - Pseudansteus crass,pes, 6 26.5 mm cl, South of Cape Comorin, India: D, Petasma dorsal view

E, Idem, ventral view. Scale = I mm. (From P£rez Farfante 1987) ’ aorsai view.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Petasma with distal margin of dorsolateral lobule slightly to strongly oblique, mesially

reaching or slightly surpassing median lobe; ventral costa distally free for about half its length

and turning slightly to strongly mesially. Thelycum open, with large lanceolate median

protuberance on sternite XIII; sternite XIV forming broad plate produced into anterolateral

hoods, delimiting anterior depression flanked by rugose area.

Distribution of branchiae, epipods, and exopods in the genus Pseudaristeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

XII

(P3)

(P4)

xrv

(P5)

Pleurobranchiae

Arthrobranchiae

r+1

Podobranchiae

Epipods

Exopods

■

Fig. 23. — A, Pseudaristeus kathleenae , 2 47 mm cl, Lagonoy Gulf, east of southern Luzon, Philippines.

Thelycum. — B, Pseudaristeus crassipes, 2 36 mm cl, Straits of Makassar, Indonesia. Thelycum. Scales =

2 mm. (All drawings from P£rez Farfante, 1987).

Species. — Pseudaristeus crassipes (Wood-Mason, 1891). Gulf of Aden; central Indian

Ocean; India; Sri Lanka; Bay of Bengal; Indonesia.

Pseudaristeus gracilis (Bate, 1888). Philippines.

Pseudaristeus kathleenae Perez Farfante, 1987. Gulf of Aden; India; Indonesia; Philippines.

Pseudaristeus protensus Perez Farfante, 1987. Arabian Sea; Bay of Bengal.

Pseudaristeus sibogae (De Man, 1911). Natal, South Africa; Madagascar; Saya de Malha

Bank; Indonesia.

Pseudaristeus speciosus (Bate, 1881). Argentina Basin.

ISABEL P6REZ FARFANTE & BRIAN KENSLEY

Family BENTHESICYMIDAE Wood-Mason, 1891

Benthesicymina Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 286.

Benthesicymae - Bouvier, 1908a, Result. Camp, scicnt. Prince Albert I, 33: 16. — Burkenroad, 1936b, Bull.

Bingham Oceanogr. Coll., 5(2): 15. — Anderson and Lindner, 1943, Trans. Am. Fish. Soc., 73: 290._

Balss, 1957, Bronn's Kl. Ordn. Tierreichs, 5(1)7(12): 1517. — Tirmizi, 1960, Scient. Rep. John Murray

Exped., 10(7): 321. — Roberts and Pequegnat, 1970, Texas A&M Univ. Oceanogr. Stud., 1: 32

Penaeidae - Barnard, 1950, Ann. S. Afr. Mus., 38: 580 [part],

Benthesicyminae - Crosnier, 1978, Faune Madagascar, 46: 14. — Hayashi, 1992, Dendr. Crust. Jap. Waters : 36.

Placed on the Official List of Family-Group Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Bull. zool. Norn., 25(4/5): 141.

Benthesicymini - Burkenroad, 1983, Crust. Issues, 1: 281, 286.

Diagnosis. — Integument thin, soft, flexible. Rostrum short, not reaching beyond eyes,

laterally compressed, dorsal rostral/postrostral teeth no more than 3, usually 2 or less, ventrally

unarmed. Carapace with branchiostegal spine; hepatic spine present or absent; postorbital and

postantennal spines lacking; cervical and postcervical sulcus reaching middorsal line;

branchiocardiac and hepatic sulci usually well defined. Abdominal somites variously carinate

occasionally ending in posterior spine. Telson bearing 1 to 4 pairs of lateral movable spines; apex

usually truncate, sometimes acute.

Eye with optic calathus bearing mesial tubercle; ocular scale and styliform projection lacking

Antennule with prosartema usually represented by tuft of setae; with 2 elongate filiform flagella.

Second pleopod of males with appendix masculina and appendix interna, lacking distolateral

projection. Third through fifth pleopods biramous. Exopods on first to third maxillipeds, present

or absent on first to fifth pereopods. Pleurobranchia present on somites IX to XIV; one

arthrobranchia on somite VII, two on somites VIII to XIII; podobranchia on second and third

maxillipeds and first to third pereopods, but only on second maxilliped in Gennadas-, epipod

present on second maxilliped to fourth or fifth pereopod.

Petasma open, generally broadly lamellar, with flexible part of ventrolateral lobule attached to

dorsolateral for much of, or for its entire length; ventral costa entirely attached. Thelycum open or

sometimes closed; in latter case having shallow seminal receptacles formed by sternal

invaginations between sternites XII and XIII at base of third pereopods.

Key to the Genera of the Family BENTHESICYMIDAE

1. Podobranchia present on second maxilliped through third pereopod. Telson

armed with more than one pair of movable lateral spines. 2

Podobranchia present only on second maxilliped. Telson armed with only

one pair of movable lateral spines . Gennadas

2. Telson apically pointed. Several abdominal somites dorsally carinate .... 3

Telson apically truncate. Only sixth abdominal somite dorsally carinate.

. Bentheogennema

3. Dactyls of fourth and fifth pereopods not multiarticulate or markedly

elongate. Hepatic spine present or absent. Benthesicymus

— Dactyls of fourth and fifth pereopods multiarticulate, markedly slender and

elongate. Hepatic spine present . Benthonectes

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Genus Bentheogennema Burkenroad, 1936

Figures 24-25

Bentheogennema Burkenroad, 1936b, Bull. Bingham oceanogr. Coll., 5(2): 56. — Anderson and Lindner, 1943,

Trans. Am. Fish. Soc., 73: 295. — Tirmizi, 1960, Scient. Rep. John Murray Exped., 10(7): 33/. —

Crosnier, 1978, Faune Madagascar, 46: 28. — Burkenroad, 1983, Crust. Issues, 1: 282. — Hayashi, 1992,

Dendr. Crust. Jap. Waters,: 57.

Gennadas - Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 343 [part].

Type Species: By original designation, Gennadas intermedius Bate, 1888, Rep. scient. Results

Voy. Challenger, 24: 343.

Type Locality: North Atlantic Ocean, off Sierra Leone, 1°47'N, 24°26'W, 3385 m; North

Atlantic Ocean between Bermuda and Azores, surface; South-east Atlantic Ocean, 35 59 S,

1°34'E, surface.

Gender: Feminine.

Fig. 24. — Bentheogennema intermedia , composite from several female specimens. Northwest Atlantic Ocean.

Lateral view. Scale = 5 mm.

Diagnosis. — Integument soft, thin. Rostrum short, triangular, not reaching cornea of eye,

apically acute, with one dorsal tooth; one postrostral tooth present in some species. Cara P a ^

lacking^hepatic spine; small marginal branchiostegal spine present on anteroventral emargination

of carapace- hepatic, suprahepatic, cervical, postcervical, and branchiocardiac sulci well parked

often further emphasized by carinae. First to fifth abdominal somites dorsally rounded sixth

somite dorsally carinate. Telson lacking, or with 1-4 pairs of movable lateral spines (usually with

2, rarely 1 or none); apically truncate.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Eye with cornea poorly-pigmented, barely wider than optic calathus; ocellus distinct; strong

conical mesial tubercle at about midlength of optic calathus. Exopod of first maxilliped distally

rounded, not segmented. Dactyli of fourth and fifth pereopods slender, consisting of single

segment. Single podobranchia on second and third maxilliped and first three pereopods.

Petasma broad, with external, median, and internal lobes distally variously developed

Thelycum closed, with triangular plate on sternite XII, broader plates on sternites XIII and XIV;

apertures of small seminal receptacles situated laterally between sternites XII and XIII.

Distribution of branchiae, epipods, and exopods in the genus Bentheogennema.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

XIII

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

FlG ' 25 n — B ‘ nthe °8? nn ema intermedia : A, 16 mm cl, Northwest Atlantic Ocean. Petasma, dorsal view

— Y 14 mm cl, Northwest Atlantic Ocean. Thelycum. Scales = 1 mm.

Species. — Bentheogennema borealis (Rathbun, 1902). North-west Pacific Ocean; Japan-

Bering Strait to Baja California. F

Cmom\& gennema burkenroadl Kr y§ ier and Wasmer, 1975. North-east Pacific Ocean; off Baja

rJZ u 1888) - 0ff Baja California ; western Atlantic Ocean from

nf h w Brazi1 ’ Gulf of Mexico; Bahamas; Gabon; Congo; south-east Atlantic Ocean off Cape

weS'Pacific'Ocean- Hawaif A&Ka; Zanzibar; Mada g ascar ; Arabian Sea; Maidive Islands; north-

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Bentheogennema pasithea (De Man, 1907). Off southern California, U.S.A. and Baja

California; East coast of Africa; Madagascar; Seychelles; India; Indonesia; Taiwan; Caroline

Islands; Gilbert Islands.

Bentheogennema stephenseni Burkenroad, 1940. Off Baja California; central Indian Ocean,

off Kermadec Islands and New Zealand, south-eastern Pacific Ocean.

Genus Benthesicymus Bate, 1881

Figures 26-27

Benthesicymus Bate, 1881, Ann. Mag. nat. Hist., (5)8: 171, 190; 1888, Rep. scient. Results Voy. Challenger,

24 : 326 . — Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 42. — De Man, 1911, Siboga Exped., 39a.

13 — Burkenroad, 1936b, Bull. Bingham Oceanogr, Coll., 5(2): 23. — Anderson and Lindner, 1943,

Trans. Am. Fish. Soc., 73: 296. — Tirmizi, 1960, Scieni. Rep. John Murray Exped., 10(7): 322^ —

Crosnier, 1978, Faune Madagascar, 46: 15. — Burkenroad, 1983, Crust. Issues, 1: 282. — Squires. 1990,

Can. Bull. Fish. Aq. Sci., 221: 21. — Kikuchi and Nemoto, 1991, J. Crust. Biol., 11(1): 64. — Hayashi,

1992, Dendr. Crust. Jap. Waters,: 37. , „ . . „ . . ,

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1808. Bull. zool. Norn., 25(4/5): 138.

Benthoecetes Smith. 1884, Rep. U. S. Commnr Fish., 1882: 391. Type species: By monotypy, Benthesicymus

bartletti Smith, 1882, Bull. Mus. comp. Zool. Harv., 10: 82. Type locality: East coast of United States, 1076-

1570 m.

Gennadas - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 46 [part].

Fig. 26. - Benthesicymus bartletti , $ 26 mm cl, North Atlantic Ocean off U.S.A. Lateral view. Scale = 10 mm.

Source: MNHN. Paris

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Type Species: By subsequent designation, by Bate, 1888, Rep. scient. Results Voy.

Challenger, 24: 329, Benthesicymus crenatus Bate, 1881, Ann. Mag. nat. Hist., (5)8: 190.

Type Locality: Mid-Pacific Ocean off Low Archipelago, 3°48'S, 152°56'W, 4750 m; Mid-

Pacific Ocean, 13°28’S, 149°30'W, 4300 m.

Gender: Masculine.

Diagnosis. — Integument thin to fairly firm. Rostrum short, falling short of, to barely

overreaching, cornea of eye, triangular, apically acute, with 2 dorsal teeth. Carapace with

marginal branchiostegal spine present, often supported by short carina; hepatic spine present or

absent; cervical, postcervical, hepatic, and branchiocardiac sulci well marked. First to third

abdominal somites dorsally rounded; posterior half of fourth somite, and fifth and sixth somites

dorsally carinate. Telson with 4 pairs of movable lateral spines; apex acute.

Cornea of eye moderately well pigmented, slightly wider than eyestalk; conical mesiodorsal

process at about midlength of eyestalk. Exopod of first maxilliped distally gently tapering to tip,

or narrowing abruptly to segmented distal part. Dactyli of fourth and fifth pereopods slender,’

uniarticulate. Podobranchia on second maxilliped to third pereopod, one small arthrobranchia on

first maxilliped and 2 on second maxilliped to fourth pereopod.

Petasma with dorsolateral lobule broad, distally sometimes bilobed, longer than ventrolateral

lobule. Thelycum either lacking seminal receptacles, or with median plate of sternite XIII broad,

having short submedian anterior lobes enclosing shallow seminal receptacles between sternites of

somites XII and XIII.

Distribution of branchiae, epipods, and exopods in the genus Benthesicymus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

Xffl

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

s or r

S or r

s or r

S or r

Species. — Benthesicymus altus Bate, 1881. California, U.S.A.; Gulf of Panama; Galapagos

Islands; Tristan da Cunha; Madagascar; Comoro Islands; Reunion; Arabian Sea; Maidive Islands-

Philippines; Japan; Coral Sea; Kermadec Islands; Fiji.

Benthesicymus armatus MacGilchrist, 1905. Arabian Sea.

Benthesicymus bartletti Smith, 1882. Western North Atlantic Ocean from Nova Scotia to Gulf

of Mexico and Caribbean; Belize; Honduras; Colombia; Venezuela; Surinam; French Guiana-

Azores; Canary Islands; Cape Verde Islands; Morocco; Mauritania; Gabon; Congo; Maidive

Islands; Bay of Bengal; Philippines; north-west Pacific Ocean.

Benthesicymus brasiliensis Bate, 1881. Gulf of Mexico; off Buenos Aires; north-east Atlantic

Ucean off Azores and Madeira; Atlantic Morocco; Tristan da Cunha.

Benthesicymus brevirostris Kikuchi and Nemoto, 1991. North-west Pacific Ocean.

Benthesicymus carinatus Smith, 1884. Caribbean; east coast of South Africa; Reunion-

Arabian Sea; north-west Pacific Ocean.

Benthesicymus cereus Burkenroad, 1936b. Bahamas; New Zealand.

Benthesicymus crenatus Bate, 1881. Central Pacific Ocean; North-west Pacific Ocean.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Fig. 27. — Bentheogennema intermedia : A, 3 16 mm cl, Northwest Atlantic Ocean. Petasma, dorsal view.

— B, $ 14 mm cl, Northwest Atlantic Ocean. Thelycum. Scales = 1 mm.

Benthesicymus hjorti Sund, 1920. Canary Islands.

Benthesicymus investigatoris Alcock and Anderson, 1899. East coast of Africa; Gulf of Aden,

Madagascar; Reunion; Saya de Malha Bank; Sri Lanka; Andaman Sea; Indonesia; Philippines,

Japan; New South Wales, Australia; Kermadec Island; Wallis and Futuna Islands; Fiji; Hawaii;

Sala-y-Gomez. , .

Benthesicymus iridescens Bate, 1881. Gulf of Mexico; Bahamas; north-east Atlantic Ocean

off Azores and Madeira; Tristan da Cunha; central Indian Ocean; Solomon Islands; New Zealand;

Fiji; North-west Pacific Ocean. _

Benthesicymus laciniatus Rathbun, 1906. Southern California; Azores; Canary Islands,

Madagascar; Reunion; Saya de Malha Bank; Japan; Hawaii.

Benthesicymus seymouri Tirmizi, 1960. Zanzibar; Madagascar.

Benthesicymus strabus Burkenroad, 1936b. South Pacific Ocean; north-west Pacific Ocean.

Benthesicymus tanneri Faxon, 1893. Off south-western U.S.A.; Pacific Mexico; Gulf of

California; Pacific Panama; Colombia; Galapagos Islands; south-west Ilo, Peru.

Benthesicymus tirmiziae Crosnier, 1978. Madagascar; Maidive Islands; Saya de Malha Bank;

Philippines. XT .

Benthesicymus urinator Burkenroad, 1936b. Comoro Islands; Reunion; Torres Straits, North¬

west Pacific Ocean; Hawaii.

Genus Benthonectes Smith, 1885

Figures 28-29

Benthonectes Smith, 1885a, Proc. U. S. natn. Mus., 7: 509. - Anderson and Lindner, 1943, Tran^Am. Fish.

Soc., 73: 290. — Tirmizi, 1960, Scient. Rep. John Murray Exped., 10(7): 333. — Crosnier, 1978, Faune

Madagascar, 46: 24.

ISABEL P£REZ FARFANTE & BRIAN KENSLEY

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1809, Bull. zool. Norn 25(4/5)- 138

La p NatUr ? , Jo a f iS ooo 3( 2, ): 229 ‘ Type species: By monotypy Hapalopoda investigator

Filhol, 1885a, La Nature, Paris, 13(2): 229, fig. 2. Type locality: 33°9'N, 11°58'W (of Paris = 9°38'W of

Greenwich) [see Holthuis, 1980a, Zool. Meded. Leiden, 55(15): 183-187],

Type Species: By monotypy, Benthonectes filipes Smith, 1885a, Proc. U. S. natn. Mus„

Type Locality: Off New Jersey, 1268-1908 m.

Gender: Masculine.

Fig. 28. — Benthonectes filipes : A, <? 19.8 mm cl, North Atlantic Ocean

Scale - 10 mm. — B, Dactylus of pereopod 5 enlarged. Scale = 5 mm.

off Virginia, U.S.A. Lateral

view.

cii«S!h G !!, 0SIS ' -, Inte g ument thin, fairly soft. Rostrum short, triangular, barely reaching to

slightly overreaching cornea of eye, bearing 2 dorsal teeth. Carapace with strong hepatic and

marked First^^founh ^' nes; . ce [ v,cal - postcervical, hepatic, and branchiocardfac sulci well

Srinate * JZ ? T S”?* 8 d ,° rSalIy r0unded ’ fifth and sixth somites dorsally

c annate. Telson with 4 pairs of movable lateral spines; apex acute. y

s m airtilhP^° mea h 0 Iil Siderab , ,y wider than eyestaIk ’ moderately well pigmented; eyestalk with

SSL?S f mCSial Ex °P° d of first maxilliped narrowing rapidly to fonn

slender multiarticulate section. Dactyli of fourth and fifth pereopods elongate multiarticulate

Podobmnch 13 present on second maxilliped to third pereopod, and one VeTS

arthrobranchia on somite VII, and two from somite VIII to XIII. y ^

PetaMM cons l } sti I n g of single narrow, apically subacute lobe. Thelycum open lacking seminal

receptacles; w.th plate of stemite XDI raised into narrowly eloag£S^S^.S^

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

rounded to subacute and free. Sternite XIV rectangular, overlapped laterally by basal lobe of fifth

pereopods.

Distribution of branchiae, epipods, and exopods in the genus Benthonectes.

Somite

(Mxpl)

Vffl

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 29. — Benthonectes filipes: A, <J 19.8 mm cl, North Atlantic Ocean off Virginia, U.S.A. Petasma, dorsal

view. — B, $ 21.5 mm cl, Hudson Canyon, northwest Atlantic Ocean. Thelycum. Scales = 1 mm.

Species. — Benthonectes filipes Smith, 1885a. Off North Carolina; off Morocco;

Madagascar; Wallis and Futuna Islands; Hawaii.

Genus Gennadas Bate, 1881

Figures 30-31

Gennadas Bate, 1881, Ann. Mag. nat. Hist., (5)8: 171, 191; 1888, Rep. scient. Results Voy. Challenger, 24: 339.

— Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 45. — De Man, 1911, Siboga Exped., 39a: 15. —

Kemp, 1913, Trans. Linn. Soc. Lond., Zool., (2)16: 60. — Calman, 1925, Rep. Fish. mar. biol. Surv. Un. S.

Afr., 4(3): 3. — Balss, 1927, Wiss. Ergebn. dt. Tiefsee-Exped. Valdivia, 23(6): 248. — Burkenroad, 1936b,

Source

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Bull. Bingham oceanogr. Coll., 5(2): 59. — Anderson and Lindner, 1943, Trans. Am. Fish. Soc., 73: 291.

— Barnard, 1950, Ann. S. Afr. Mus., 38: 627. — Tirmizi, 1960, Scient. Rep. John Murray Exped., 10(7):

340. — Kensley, 1971b, Ann. S. Afr. Mus., 57(12): 272. — Crosnier, 1978, Faune Madagascar, 46: 33. —

Burkenroad, 1983, Crust. Issues, 1: 282. — Squires, 1990, Can. Bull. Fish. Aq. Sci., 221: 26. — Hayashi,

1992, Dendr. Crust. Jap. Waters,: 44.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1811, Bull. zool. Norn., 25(4/5): 139.

Amalopenaeus Smith, 1882, Bull. Mus. comp. Zool. Harv., 10(1): 86. Type species: By monotypy,

Amalopenaeus elegans Smith, 1882, Bull. Mus. comp. Zool. Harv., 10(1): 87. Type locality: East coast of

United States, South Carolina to New Jersey, 680-2986 m.

Pasiphodes Filhol, 1885b, Vie Fond Mer, pi. 3. Type species: By monotypy, Pasiphodes purpureus Filhol, 1885,

Vie Fond Mer, pi. 3. Type locality: off Cap Ghir, Morocco, 2210 m. [Dr. L. Holthuis (in litt. 1996) refers to

A. Milne Edwards' original color sketch of Pasiphodes purpureus in the Paris Museum, probably from the

"Talisman" station 38, and suggests that it is Gennadas valens (Smith, 1884)).

Type Species: By monotypy, Gennadas parvus Bate, 1881, Ann. Mag. nat. Hist., (5)8: 192.

Type Locality: Off Japan, 26°29'N, 137°57'E, 4437 m.

Gender: Masculine.

Fig. 30. — Gennadas elegans , 2 11 mm cl, North Atlantic. Lateral view. Scale = 5 mm.

Diagnosis. — Integument thin, soft. Rostrum short, with single dorsal tooth, not reaching as

far as cornea of eye. Carapace with distinct cervical and postcervical sulci reaching dorsal

midline; antennal angle narrowly rounded; branchiostegal angle square; weak hepatic and

branchiocardiac carinae present. First to fifth abdominal somites dorsally rounded, sixth somite

dorsally carinate. Telson apically truncate, with single pair of movable posterolateral spines.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Cornea of eye moderately well pigmented; eyestalk with strong conical mesiodorsal tubercle.

Antennular peduncle segments 2 and 3 somewhat expanded. Exopod of first mmcilliped lacking

distal segmentation. Dactyli of fourth and fifth pereopods slender, u^^culate. Eodobrancba o

second maxilliped, lacking on third maxilliped and all pereopods. Arthrobranchia on somite

rudimentary, two arthrobranchiae from somite VIII to XIII. _ , ,

Petasma with distal margin divided into external, median, and internal lobes; accessory

always present. Appendix masculina bilamellate. Stemites XU-XIV variously modified in fern

to fomf thelycum, with small shallow seminal receptacles at base of third pereopods either

opening separately, or opening within a common median groove or depression.

Distribution of branchiae, epipods, and exopods in the genus Gennadas.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

xrv

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

—:—

Fig. 31. - Gennadas panms : A, <J 7 mm cl. South Atlantic. Petasma, dorsal view. - B, 2 5 mm cl, Indian

Ocean. Thelycum. Scales = 1 mm.

Species. — Gennadas barbari Vereshchaka, 1990. Naska and Sala-y-G6mez ridges,

southeastern Pacific Ocean. ^ , _ ...

Gennadas bouvieri Kemp, 1909. OffBaja California; Gulf of Me:

Rermuda- south-east Atlantic Ocean; south-west Indian Ocean, Zanzibar, Cult ot Aden,

Madagascar; Maidive Islands; Gulf of Arabia; Bay of Bengal; Indonesia; Philippines; Japan, east

coast of Australia; north-west Pacific Ocean.

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

Gennadas brevirostris Bouvier, 1905b. Caribbean; southern Spain; Eastern Atlantic Ocean

from Sierra Leone to South Africa.

Gennadas capensis Caiman, 1925. Off Baja California; Gulf of Mexico; Bahamas; Caribbean

Sea; Bermuda; Canary Islands; off Mauritania; south-east Atlantic Ocean; south-west Indian

Ocean; Madagascar; east coast of Australia; Wallis and Futuna Islands; north-west Pacific Ocean.

Gennadas clavicarpus De Man, 1907. South-east Atlantic Ocean off Cape of Good Hope,

South Africa; south-west Indian Ocean; Zanzibar; Gulf of Aden; Reunion; central Indian Ocean;

Indonesia.

Gennadas crassus Tirmizi, 1960. Zanzibar.

Gennadas elegans (Smith, 1882). Western north Atlantic Ocean from New York to Florida;

Bahamas; Bermuda; Cape Verde Islands; Mediterranean; North Atlantic Ocean off Morocco; off

Gabon; south-east Atlantic Ocean.

Gennadas gilchristi Caiman, 1925. South-east Atlantic Ocean; south-west Indian Ocean; east

coast of Australia.

Gennadas incertus (Balss, 1927). North-east Pacific Ocean off Oregon and Baja California;

South-east Atlantic Ocean; south-west Indian Ocean; east African coast off Kenya and Tanzania;

Zanzibar; Madagascar; Seychelles; Gulf of Arabia; east coast of Australia; Japan; north-west

Pacific Ocean.

Gennadas kempi Stebbing, 1914. South-east Atlantic Ocean; south-west Indian Ocean; east

coast of Australia.

Gennadas parvus Bate, 1881. North-east Pacific Ocean; south-east Atlantic Ocean off Cape of

Good Hope, South Africa; south-west Indian Ocean; Zanzibar; Gulf of Aden; Madagascar;

Arabian Sea; Japan; Hawaii; north-west Pacific Ocean.

Gennadas pectinatus Schmitt, 1921. Off Santa Catalina Island, California.

Gennadas propinquus Rathbun, 1906. North-east Pacific Ocean off Oregon and Baja

California; South-east Atlantic Ocean; south-west Indian Ocean; Gulf of Aden; Madagascar;

Arabian Sea, Philippines; east coast of Australia; Wallis and Futuna Islands; north-west Pacific

Ocean.

Gennadas scutatus Bouvier, 1906a. Off Baja California; Isla Clarion, off Mexico, and Isla

Lobos de Terra, northern Peru; Gulf of Mexico; Bahamas; Caribbean Sea; north-east Atlantic

Ocean off Azores; Cape Verde Islands; Gabon; Congo; Angola; south-east Atlantic Ocean off

Cape of Good Hope; south-west Indian Ocean off Natal; Gulf of Aden; Madagascar; Seychelles;

Indonesia; east coast of Australia; Indo-Pacific Ocean from east coast of Africa to west coast of

America.

Gennadas sordidus Kemp, 1910a. Off Baja California; Gulf of California and southern

Mexico; Gulf of Aden; Gulf of Oman; Arabian Sea.

Gennadas talismani Bouvier, 1906a. Gulf of Mexico; eastern Atlantic Ocean from Cape Verde

Islands to Cape of Good Hope.

Gennadas tinayrei Bouvier, 1906a. Off Baja California; north-west Atlantic Ocean; Bahamas-

Bermuda; Azores; Gibraltar; south-east Atlantic Ocean off St. Helena Island and Cape of Good

Hope; south-west Indian Ocean; Zanzibar; Madagascar; Seychelles; Maidive Islands; Japan- east

coast of Australia; north-west Pacific Ocean.

T Ge ™ adas A valens (Smith, 1884). Gulf of Mexico; Caribbean; Bahamas; Bermuda; Canary

Islands; Mediterranean; eastern Atlantic Ocean from Ireland to Cape of Good Hope, South

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Family PENAEIDAE Rafinesque-Schmaltz, 1815

Penedia Rafinesque-Schmaltz, 1815, Anal. Nat. Tabl. Univers,: 98 [subfamily of the Plyonuria].

Penaeidae - Dana, 1852a, Proc. Acad. nat. Sci. Philad., 6: 18; 1852b, Am. J. Sci. Arts, (2)14: 123; 1853, Class.

Geog. Distrib. Crust., : 1434. — Boas, 1880, Vidensk. Selsk. Skr., (6)1: 155. — Bate, 1881, Ann. Mag.

nat. Hist., (5)8: 171, 173; 1888, Rep. scient. Results Voy. Challenger, 24: 220. — Ortmann, 1898, Bronn's

Kl. Ordn. Tierreichs, (5)2: 1118. — Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 11. — Bouvier,

1908a, Result. Camp, scient. Prince Albert I, 33: 9. — De Man, 1911, Siboga Exped., 39a: 1. — Barnard,

1950, Ann. S. Afr. Mus., 38: 580. — Balss, 1957, Bronn's Kl. Ordn. Tierreichs, 5(1)7(12): 1516. — PErez

Farfante, 1978, FAO Sp. Idem. Sh., 6: 1. — Williams, 1984, Shr. Lob. Crabs Atl. Coast U.S., : 22. — De

Freitas, 1987, Investl. Rep. oceanogr. Res. Inst., Durban, 58: 1. — PErez Farfante, 1988, NOAA Tech.

Rep. NMFS, 64: iii, 8. — Dall et al., 1990, Adv. Mar. Biol., 27: 59, 60.

Placed on the Official List of Family Group Names in Zoology, International Commission on Zoological

Nomenclature, 1955, Direction 15, Op. Decl. Int. Comm. zool. Nom., 1C(C5): 70.

Penaeinae - Ortmann, 1898, Bronn's Kl. Ordn. Tierreichs, 5(2): 1120. — Bouvier, 1908a, Result. Camp, scient.

Prince Albert I, 33: 6. — Burkenroad, 1934b, Bull. Am. Mus. Nat. Hist., 68(2): 72. — Balss, 1957,

Bronn's Kl. Ordn. Tierreichs, 5(1)7(12): 1517.

Parapenaeinae Ortmann, 1898, Bronn's Kl. Ordn. Tierreichs, (5)2: 1120.

Placed on the Official List of Family-Group Names in Zoology, International Commission on Zoological

Nomenclature, 1961, Opinion 611, Bull. zool. Nom., 18(5): 307.

Funchaliae Bouvier, 1908a, Result. Camp, scient. Prince Albert I, 33: 6. — Balss, 1957, Bronn's Kl. Ordn.

Tierreichs, 5(1)7(12): 1518.

Haliporae Bouvier, 1908a, Result. Camp, scient. Prince Albert I, 33: 6. — Balss, 1957, Bronn's Kl. Ordn.

Tierreichs, 5(1)7(12): 1518.

Funchaliinae - Burukovsky and Romensky, 1991, Biull. Mosk. O-va Ispyt Prirod. Otd. Biol., 96(6): 61.

Diagnosis. — Body compressed, comparatively slender. Rostrum well developed, extending

to or beyond distal margin of eye; armed with dorsal and sometimes also with ventral teeth.

Carapace without postorbital spine; antennal and hepatic spines usually present; cervical sulcus

ending well ventral to dorsal midline. Posterior abdominal somites carinate. Telson sharply

pointed, with or without lateral spines.

Eye with optic calathus almost always lacking mesial tubercle; basal article of eyestalk

produced into moderately to slightly developed, never freely projecting, distomesial scale; ocular

plate lacking styliform projection. Antennule with foliaceous prosartema, flagella of about same

length, borne on apex of third segment. Exopod present on second maxilliped (except in Arte-

mesia, Macropetasma, and Protrachypene) and third maxilliped (absent only in Macropetasma),

and first four pereopods. Third through fifth pleopods biramous. Pleurobranchia on somites DC

through XII, and sometimes on XIII and XIV; rudimentary arthrobranchia usually present on

somite VII, two arthrobranchiae on VIII through XII, and posterodorsal one on XIII (sometimes

rudimentary anteroventral one also present on XIII); podobranchia on second maxilliped only.

Epipods borne on first maxilliped and usually on second, lacking on fourth and fifth pereopods.

Petasma semi-open or semi-closed. Second pleopod of males bearing appendix masculina

only, lacking appendix interna and distolateral projection. Thelycum open or closed.

Key to the Genera of the Family PENAEIDAE

1. Rostrum armed with dorsal and usually also ventral teeth (absent only in

Funchalia). Pleurobranchia on somite XIV (last thoracic somite) . 2

— Rostrum usually armed with dorsal teeth only. No pleurobranchia on

somite XTV .10

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

2. Integument glabrous and polished. Three short, well defined cicatrices on

sixth abdominal somite . 3

— Integument setose. Single, long (sometimes interrupted) cicatrix on sixth

abdominal somite or none . 8

3. Adrostral sulcus and carina short, falling distinctly short or extending to

about level of epigastric tooth. Gastrofrontal carina absent. 4

— Adrostral sulcus and carina long, reaching much beyond epigastric tooth,

usually almost to posterior margin of carapace. Gastrofrontal carina present

. 6

4. Hepatic carina absent or, if present, moderately to ill defined.

. Fenneropenaeus

— Hepatic carina prominent . 5

5. Thelycum open. Petasma with ventral costa short, not reaching distal

margin of lateral lobe. Litopenaeus

— Thelycum closed. Petasma with ventral costa long, reaching distal margin

of lateral lobe. Penaeus s.s.

6. Gastrofrontal carina not turning anterodorsally upon itself at posterior end.

Sixth abdominal somite with well defined dorsolateral sulcus. Telson

unarmed . Farfantepenaeus

— Gastrofrontal carina turning anterodorsally upon itself at posterior end.

Sixth abdominal somite lacking dorsolateral sulcus. Telson usually armed

with three pairs of movable lateral spines (absent only in Melicertus

camliculatus ). 7

7. Gastrofrontal sulcus not markedly bifid posteriorly. Thelycum with pair of

lateral plates on stemite XIV shielding saclike seminal receptacle opening

along midline. Melicertus

— Gastrofrontal sulcus markedly bifid posteriorly. Thelycum with single plate

on sternite XIV infolded laterally, forming pouch opening anteriorly,

functioning as seminal receptacle . Marsupenaeus

8. Integument with numerous sulci overlapped by rows of densely set stiff

setae projecting from one margin. Telson with four pairs of movable lateral

spines. Antennular flagella short, less than half length of carapace .

•. Heteropenaeus

— Integument with few sulci not overlapped by rows of stiff setae. Telson

with three pair of fixed lateral spines. Antennular flagella long, longer than

carapace. 9

9. Rostrum armed with dorsal teeth only. Carapace lacking dorsolateral

carina. Incisor processes of mandible extremely elongate, scythelike and

crossing posterior to labrum. Petasma asymmetrical with either right or left

half longer than opposite . Funchalia

— Rostrum armed with dorsal and ventral teeth. Carapace with dorsolateral

carina. Incisor processes of mandible short, not crossing. Petasma

symmetrical, both halves of same length . Pelagopenueus

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

10. Telson with pair of well developed fixed subapical spines (preceded by

lateral movable spines). First segment of antennular peduncle usually

bearing ventromesial (parapenaeid) spine . 11

— Telson without pair of subapical fixed spines (only occassionally present as

in Parapenaeopsis stylifera and Rimapenaeus fuscina), usually with

movable lateral spines. First segment of antennular peduncle lacking

parapenaeid spine. 14

11. Carapace with longitudinal suture (extending at least 0.8 of its length) and

transverse suture. Not more than one pair of minute lateral spines anterior

to subapical spines . Parapenaeus

— Carapace without longitudinal suture. Two or more pairs of conspicuous

spines anterior to subapical spines . 12

12. Pterygostomian spine absent (anteroventral margin of carapace rounded).

First pereopod without basial spine. Exopods lacking on second

maxillipeds and all pereopods. Artemesia

— Pterygostomian spine present (anteroventral margin of carapace toothlike).

First pereopod with basial spine. Exopods on all maxillipeds and

pereopods .

13. Third maxilliped and second pereopod with basial spine. Petasma asym-

metrical . Mctapcnavopsis

— Third maxilliped and second pereopod without basial spine. Petasma

symmetrical . Penaeopsis

14. Pleurobranchia on somite XIII (penultimate thoracic somite). Exopods on

maxillipeds and four anterior pairs of pereopods, lacking on fifth pereopod

. Metapenaeus

— Pleurobranchia absent on somite XEE. Exopods present on all pereopods or

absent from four posterior pairs . 15

15. Exopods on first maxilliped and first pereopod. Petasma with lateral lobes

distally produced into extremely long, filamentous processes. Thelycum

with lateral plates curved posteriorly and expanded into bulbous swelling

anteriorly . Macropetasma

— Exopods at least on third maxilliped and all pereopods. Petasma with lateral

lobes not distally produced into extremely long filamentous processes.

Thelycum with lateral plates neither strongly curved posteriorly nor

expanded into bulbous swelling anteriorly .16

16. Carapace lacking longitudinal and transverse sutures. Telson with subapical

pair of lateral movable spines mounted on elongate shoulder. Epipods not

furcate. Petasma with ventrolateral lobule produced into two or three distal

flaps . Trachypenaeopsis

— Carapace with either longitudinal or transverse sutures or both, absent only

in Miyadiella. Telson lacking spines or with movable ones not mounted on

shoulders or mounted on slight ones. Petasma with ventrolateral lobule not

produced into distal flaps . 17

17. Second maxilliped without exopod. First three pereopods with elongate

chela, propod lengthened and dactyl very short (much less than half length

of propod) . Protrachypene

ISABEL PSREZ FARFANTE & BRIAN KENSLEY

— Second maxilliped with well developed exopod. First three pereopods with

not elongate chela, propod not lengthened and dactyl not less than half

length of propod .18

18. Fourth and fifth pereopods very elongate, subflagelliform, much longer

than first three.19

— Fourth and fifth pereopods not elongate, only slightly longer than first

three, or only fifth elongate. 20

19. Integument pubescent. Telson armed with four pairs of movable spines.

Fourth and fifth pereopods with entire dactyl . Tanypenaeus

— Integument glabrous. Telson unarmed. Fourth and fifth pereopods with

multiarticulate dactyl. Xiphopenaeus

20. Carapace lacking longitudinal sutures. Second pereopod armed with ischial

spine . 21

— Carapace with longitudinal sutures. Second pereopod lacking ischial spine.

.22

21. Eyestalk not surpassing first antennular segment. Basal rostral teeth and

postrostral carina lacking spinules. Thelycum closed . Atypopenaeus

— Eyestalk conspicuously surpassing first antennular segment, sometimes

reaching or overreaching antennular peduncle. Basal rostral teeth and

postrostral carina studded with spinules. Thelycum open . Miyadiella

22. Body slender, integument thin. Third pereopod lacking epipod .

. Parapenaeopsis

— Body thickset, integument thick. Third pereopod armed with epipod .... 23

23. Carapace with longitudinal suture long, conspicuously overreaching hepatic

spine. Third maxilliped armed with basial spine. Rimapenaeus

— Carapace with longitudinal suture short, ending well anterior to hepatic

spine. Third maxilliped lacking basial spine . 24

24. Thelycum with plate on stemite XIV very short medially, deeply excavate,

embracing extremely long caudal extension of median protuberance.

Petasma with distolateral projections either moderately broad to rather

narrow basally and extending laterally to mesially or forward directed

hooklike tip or extremely broad basally but narrowing rapidly, ending

inforward directed tip . Megokris

— Thelycum with plate on stemite XIV relatively long, not excavate, median

protuberance lacking long caudal extension. Petasma with distolateral

projections either relatively narrow and directed laterally almost straight or

curving backwards, or extremely broad and ending in long, twisted process

.25

25. Thelycum with plate on stemite XIV not emarginate anteriorly, instead

continuous with median protuberance, lacking anterior transverse groove.

Petasma with distolateral projections extremely broad, extending laterally

and forming apically long, flat twisted process . Trachypenaeus

— Thelycum with plate on stemite XIV shallowly emarginate or occasionally

produced in small median prominence, not continuous with median

protuberance. Petasma with distolateral projections tapering gently from

relatively narrow base, extending almost straight laterally or curving

slightly backwards . Trachysalambria

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Genus Artemesia Bate, 1888

Figures 32-34

Artemesia Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 280. — Bouvier, 1908b, Bull. Inst. Oceanogr.,

119: 4. — Burkenroad, 1934a, Bull. Bingham oceanogr. Coll., 4(7): 11; 1934b, Bull. Am. Mus. nat. Hist.!

68(2): 73; 1983, Crust. Issues, 1: 282. — Dall et al., 1990, Adv. Mar. Biol., 27: 63.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1806, Bull. zool. Norn., 25(4/5): 138.

Artemisia - Mistakidis & Neiva, 1964, Nature, Lond., 202: 471. — Starobogatov, 1972, Akad. Nauk., USSR.,

Zool. Inst. Issled. Fauny Morei, 10(18): 385.

Type Species: By monotypy, Artemesia longinaris Bate, 1888, Rep. scient. Results Voy.

Challenger, 24: 281.

Type Locality: Off Montevideo, Uruguay, 35°02'S, 55°15'W, 24 m.

Gender: Feminine.

Fig. 32. — Artemesia longinaris, 2 28 mm cl, Rawson, Chubut, Argentina. Lateral view. Scale = 10 mm.

Diagnosis. — Integument glabrous. Rostrum bearing dorsal teeth only, limited to base, long,

slender, considerably overreaching antennular peduncle; epigastric tooth usually distinctly

separated from first rostral tooth, occasionally close to latter. Carapace with antennal and hepatic

spines, lacking orbital, pterygostomian (although angle produced) and branchiostegal spines;

postrostral carina extending to about two-thirds length of carapace; postocular sulcus present;

cervical sulcus short; hepatic sulcus well marked, accompanied by sharp carina anteriorly;

branchiocardiac carina clearly distinct, but not prominent; longitudinal and transverse sutures

absent. Sixth abdominal somite lacking cicatrices. Telson armed with pair of fixed posterolateral

spines and two or three pairs of more anterior movable spines.

Source: MNHN. Paris

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Fig. 33. —Artemesia longinaris, 6 25 mm cl,

Rawson, Chubut, Argentina. Petasma, dorsal

view. Scale = 1 mm.

cavity at base of fourth pereopod. Apparently,

exits through the slits.

Antennule bearing parapenaeid spine;

antennular flagella subfiliform, longer than

carapace. Palp of first maxilla short, entire,

gently tapering to rounded apex and bearing large

proximomesial lobe and lacking mesial spine.

Basial and ischial spines lacking on first and

second pereopods; fourth and fifth pereopods

subfiliform. Exopods lacking on pereopods;

rudimentary anteroventral filamentous

arthrobranchia on somite XIII.

Petasma symmetrical, semiclosed, with distal

part of dorsolateral lobules produced in strong

spoutlike shallow channelled distal projections

and bearing long, slender distodorsal horns

mesially. Appendix masculina greatly expanded

from narrow base. Thelycum closed, with single

heavily sculptured plate on sternite XIV and

strongly developed median protuberance on

sternite XIII. Seminal receptacles consisting of

paired pyriform sacs bearing distally two minute

lobules and connected distomesially by narrow

transverse troughlike duct situated on distal

margin of sternite XIV; opening of duct

continuous with slits extending along distal

extremity of sacs; slits, in turn, opening into

sperm enters into the duct, flows to the sacs, and

Fig. 34. — Artemesia longinaris , 2 28 mm cl, Rawson, Chubut, Argentina: A, Thelycum. Scale = 1 mm. —

B, 2 18 mm cl, off mouth of Rio de la Plata, Argentina. Seminal receptacles, dorsal view. Scale = 1 mm.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Distribution of branchiae, epipods, and exopods in the genus Artemesia

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(PS)

Pleurobranchiae

Arthrobranchiae

r+1

Podobranchiae

Epipods

Exopods

Species. —Artemesia longinaris Bate, 1888. Rio de Janeiro, Brazil to Chubut, Argentina.

Genus Atypopenaeus Alcock, 1905

Figures 35-38

Penaeus - Stimpson, 1860, Proc. Acad. nat. Sci. Philad., 12: 43. — Henderson, 1893, Trans. Linn. Soc. Lond.,

(2), Zool., 5: 450.

Atypopeneus Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 524; 1906, Cat. Indian Dec. Crust., 3(1): 45. —

Burkenroad, 1934a, Bull. Bingham oceanogr. Coll., 4(7): 41; 1934b, Bull. Am. Mus. Nat. Hist., 68(2): 95;

1983, Crust. Issues, 1: 283, 287.

Parapenaeopsis - Kubo, 1936, J. imp. Fish. Inst., Tokyo, 31(2): 55.

Fig. 35.—Atypopenaeus bicomis, 2 20 mm cl, Van Diemen Gulf, Australia. Lateral view. Scale = 10 mm

Source: MNHN. Paris

ISABEL P£REZ FARFANTE & BRIAN KENSLEY

Atypopenaeus [emendment of Atypopeneus Alcock, 1905, under the plenary powers by the International

Commission on Zoological Nomenclature, 1969]. Placed on the Official List of Generic Names in Zoology,

International Commission on Zoological Nomenclature 1969, Opinion 864, Name No. 1807, Bull. zool.

Nom., 25(4/5): 138. — De Man, 1911, Siboga Exped., 39a: 83. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1):

365. — Dall, 1957, Aust. J. mar. Freshwat. Res., 8(2): 198. — Racek and Dall, 1965, Verh. K. Ned. Akad.

Wet. Nat., 56(3): 84. — Starobogatov, 1972, Akad. Nauk SSSR Zool. Inst. Isled, Fauney Moreii, 10(18):

369, 386. — Holthuis, 1980b, FAO Fish. Synop., 125, 1: 13. — Dall et al., 1990, Adv. Mar. Biol., 27: 64.

— Hayashi, 1992, Dendr. Crust. Jap. Waters, : 70.

Type Species: By original designation, Penaeus compressipes Henderson, 1893, Trans. Linn.

Soc. Lond., (2), Zool. 5: 450.

Type Locality: Gulf of Martaban, India.

Gender: Masculine.

Fig. 36. — Atypopenaeus stenodactylus , 2 14 mm cl, southwest of Hainan Island, China. Lateral view.

Scale = 5 mm.

Diagnosis. — Integument glabrous, sparsely and microscopically setose, or sometimes partly

granulate. Rostrum armed with dorsal teeth only, variable in length; epigastric tooth usually

widely separated from first rostral tooth. Carapace with minute, occasionally absent orbital and

small non-buttressed antennal spines; hepatic spine present or absent; lacking pterygostomian

spine; postocular sulcus well defined; antennal carina and orbito-antennal sulcus absent; cervical

sulcus usually short; hepatic sulcus feeble or absent, hepatic carina marked anterior to hepatic

spine; branchiocardiac sulcus feeble; longitudinal suture lacking, transverse suture marked (in

A. compressipes, A. dearmatus and A. stenodactylus) or indistinct. Sixth abdominal somite

lacking cicatrices. Telson unarmed.

Antennule lacking parapenaeid spine; antennular flagella as long as or longer than antennular

peduncle, sometimes longer than carapace. Palp of first maxilla short, biarticulate, distal article

minute, much larger basal article bearing weak proximolateral lobule and conspicuous

proximomesial and smaller distomesial lobules, latter armed with spine; also distolateral row of

spinules. Fifth pereopod slender and very elongate, much longer than preceding ones, extending

at least by dactyl beyond antennal scale and as far as tip of antennular flagella. Basial spine

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS 75

lacking on first, present on second and third pereopods, ischial spine on first and second. Minute

ridge present in place of anteroventral arthrobranchia on somite XITT

Fig. 37 A. — Atypopenaeus bicomis, 2 20 mm cl, Van Diemen Gulf, Australia. Thelycum. Scale = 1 mm.

Fig. 37 B. — Atypopenaeus stenodactylus, 2 15 mm cl, southwest of Hainan Island, China. Thelycum.

Scale = 10 mm.

Fig. 37 C-D. — Atypopenaeus formosus, 2 22 mm cl, Van Diemen Gulf, Australia. C, Thelycum and outline of

ventral view of seminal receptacles. D, Seminal receptacles, dorsal view. Scales = 1 mm.

Source: MNHN, Paris

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Fig. 38 A. — Atypopenaeus bicornis, 6 14 mm cl, Van Diemen Gulf, Australia. Petasma, dorsal view.

Scale = 1 mm.

Fig. 38 B. —Atypopenaeus stenodactylus , c? 9 mm cl, southwest of Hainan Island, China. Petasma, dorsal view.

Scale = 0.5 mm.

Petasma symmetrical, semiclosed, subcylindrical, with lateral lobes variable in shape,

produced distally into elongate forwardly directed, acute or blunt projections, or ovoid, or with

minute laterally directed hornlike projections. Appendix masculina small, jointed to strong

dorsomesial rib of endopod. Thelycum closed, with paired, distinctly separated, long, flaplike

lateral plates often flanking posteriorly deep concavity on stemite XIV; median protuberance on

stemite XIII elongate (but falling conspicuously short of posterior thoracic ridge of stemite XIV),

roughly hourglass in shape, or acute anteriorly. Seminal receptacles consisting of paired

proximally rigid, distally membranous curved sacs continuous basally through sternal

invagination, between stemites XIV and XIII, forming transverse groove opening by horizontal

slit; sacs extending anteriorly into stemite XIII and opening through paired slits lying dorsal to

anterolateral part of median protuberance, above base of fourth pereopods. These receptacles, as

those in members of Metapenaeus, are unusual in that they are proximally rigid, instead of

entirely membranous, and extend anteriorly rather than posteriorly, from stemite XIV.

Distribution of branchiae, epipods, and exopods in the genus Atypopenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

Xffl

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Species. — Atypopenaeus bicornis Racek and Dali, 1965. New Guinea; Northern Territory,

Australia.

Atypopenaeus compressipes (Henderson, 1893). India to Malaysia; Indonesia; Hong Kong;

Philippines; Japan; New Guinea; Northern Territory, Australia.

Atypopenaeus dearmatus De Man, 1907. Indonesia; Philippines; Van Diemen Gulf, Australia.

Atypopenaeus formosus Dali, 1957. New Guinea; Northern Territory, Queensland, northern

New South Wales, Australia.

Atypopenaeus stenodactylus (Stimpson, 1860). India; Sri Lanka; Malaysia; Indonesia; Gulf of

Tonkin; South China Sea; Philippines; Hong Kong; Taiwan; Japan; New Guinea; Northern

Territory, Australia.

Genus Farfantepenaeus Burakovsky, 1997

Figures 39-40

Penaeus - Smith, 1885b, Proc. U. S. natn. Mus., 8(11/12): 170 [pari]; 1886. Rep. U. S. Commnr Fish., for

1885: 684 [part], — De Man, 1911, Siboga Exped., 39a: 95 [part]. — Burkenroad, 1934b, Bull. Am. Mus.

nat. Hist., 68(2): 74 [part]. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 268 [part], — Dall, 1957, Aust. J.

mar. Freshwat. Res., 8(2): 141 [part].

Paneus - Collins and Smith, 1892, Bull. U. S. Fish. Commn., 11: 102.

Peneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 513 [part]; 1906, Cat. Indian Dec. Crust., 3(1): 7 [part],

Penaeus (Melicertus) - P£rez Farfante, 1969, Fish. Bull. U.S., 67(3): 466 [part],

Penaeus (Farfantepenaeus) Burukovsky, 1972, Trudy AtlantNIRO, 42: 10, 13, 17. — P£rez Farfante, 1988,

NOAA Tech. Rep., NMFS, 64: 9. — Burukovsky, 1997, Proc. biol. Soc. Wash., 110: 154.

Fig. 39. — Farfantepenaeus aztecus, 2 51.5 mm cl, off Galveston, Texas. Lateral view. Scale = 10 mm

Source:

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

TyPE Species: By present designation, Penaeus brasiliensis var. aztecus Ives, 1891, Proc.

Acad. nat. Sci. Philad., 190; lectotype for Penaeus brasiliensis var. aztecus Ives, designated by

PfiREZ Farfante, 1969, Fish. Bull., U. S„ 67(3): 528.

Type Locality: Veracruz, Mexico.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum with dorsal and usually 2 (occasionally 1 or 3)

ventral teeth, moderately long, in young slightly surpassing antennular peduncle, in adult

sometimes reaching midlength of second antennular segment. Carapace with antennal and hepatic

spines pronounced, lacking orbital and pterygostomian spines; postocular sulcus absent;

postrostral carina long, adrostral carina and sulcus usually long, extending almost to posterior

margin of carapace, but sometimes only to posterior one-sixth of carapace; gastrofrontal carina

not turning anterodorsally upon itself at posterior end; gastro-orbital carina usually long,

extending more than three-quarters of distance between hepatic spine and orbital margin; orbito-

antennal sulcus well marked; cervical and hepatic carina sharp, accompanying sulci deep;

branchiocardiac carina lacking; longitudinal and transverse sutures absent. Sixth abdominal

somite bearing three cicatrices, dorsolateral sulcus narrow to quite broad. Telson unarmed.

Antennule lacking parapenaeid spine; antennular flagella much shorter than carapace. Palp of

first maxilla elongate, consisting of 3 or 4 articles, distal ones forming flagellum; large basal

article produced into setose proximal lobule on lateral and mesial margins and bearing 1,

occasionally 2, long subdistomesial spines and distolateral row of spinules. Basial spine on first

and second pereopods.

Petasma symmetrical, semiclosed, partly to almost completely hooded by well developed

distomedian projections; ventral costae long, strongly curved to almost straight distally, nearly

meeting midventrally. Appendix masculina subtriangular to suboval, armed with numerous

marginal spines. Thelycum closed, with pair of lateral plates on sternite XIV meeting along

midline, shielding simple, median, saclike seminal receptacle; sternite XIII with well developed

median protuberance.

Andrews (1911) gave an illustrated account of the seminal receptacle of P. brasiliensis (=

F. duorarum), and more recently Eldred (1958) figured the seminal receptacle of F. duorarum

and P£rez Farfante (1969) that of F. brasiliensis. The formation and structure of the

spermatophores of F. aztecus and F. duorarum were studied by Bauer and Cash (1991). They

consist of a main body containing sperm and a tail-like appendage which when exposed to sea

water during insemination unfolds, forming a membrane that fills the slit between the lateral

plates and extends along their mesial margins, or spreads over most of the lateral plates (in

F. brevirostris, pers. obs.), plugging the opening of the seminal receptacle. These

spermatophores exhibit a much simpler structure than those of genera with an open thelycum,

lacking attachment extensions, wing, flap, flange, and adhesive materials.

Distribution of branchiae, epipods, and exopods in the genus Farfantepenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

XUI

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Fig. 40. — Farfantepenaeus aztecus: A, 6 10 mm cl, off Galveston, Texas. Petasma, lateral view. Scale = 1 mm.

— B, $ 51.5 mm cl, off Galveston, Texas. Thelycum. Scale = 5 mm.

Species. — Farfantepenaeus aztecus (Ives, 1891). Massachusetts to Florida; Gulf of Mexico

to Yucatan.

Farfantepenaeus brasiliensis (Latreille, 1817). North Carolina to Florida Keys, southern Gulf

of Mexico to Yucatan; West Indies; Caribbean coast of Central and South America to Rio Grande

do Sul, Brazil; Bermuda.

Farfantepenaeus brevirostris (Kingsley, 1878). Western Baja California Sur and Gulf of

California to northern Peru; Galapagos Islands.

Farfantepenaeus califomiensis (Holmes, 1900). San Francisco Bay, California; Western Baja

California Norte; Gulf of California to Callao, Peru; Galapagos Islands.

Farfantepenaeus duorarum (Burkenroad, 1939). Chesapeake Bay to Florida; Gulf of Mexico;

Dry Tortugas to Cape Catoche, south to Islas Mujeres; Bermuda.

Farfantepenaeus notialis (Perez Farfante, 1967). Western Atlantic Ocean; from Cuba to Virgin

Islands; Quintana Roo, Mexico, Caribbean coast of Central and South America to Brazil; eastern

Atlantic Ocean from Mauritania to Angola.

Farfantepenaeus paulensis (Perez Farfante, 1967). Cabo Frio, Brazil, to Buenos Aires

Province, Argentina.

Farfantepenaeus subtilis (Perez Farfante, 1967). Cuba; Antilles; Caribbean coast of Central

and South America from Honduras to Rio de Janeiro, Brazil.

Genus Fenneropenaeus Perez Farfante, 1969

Figures 41-42

Cancer - Osbeck, 1765, Reise Ostind. China,: 151 [part].

Peneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 513; 1906, Cat. Indian Dec. Crust., 3(1): 7 [part].

Source:

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Penaeus - De Man, 1911, Siboga Exped., 39a: 95 [part], — Burkenroad, 1934b, Bull. Am. Mus. nat. Hist.,

68(2): 74 [pan]. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 268 [part]. — Dall, 1957, Aust. J. mar.

Freshwat. Res., 8(2): 141 [part].

Penaeus (Fenneropenaeus) P<5rez Farfante, 1969, Fish. Bull., U. S., 67(3): 466. — Liu and Zhong, 1988, Penaeoid

Shr. S. China Sea, : 105. — Hayashi, 1992, Dendr. Crust. Jap. Waters, : 122.

Penaeus (Penaeus) - Burukovsky, 1972, Trudy AtlantNIRO, 42: 5, 8, 13, 16.

Fig. 41. — Fenneropenaeus indicus, $ 46 mm cl, Joseph Bonaparte Gulf, Northern Territory, Australia. Lateral

view. Scale = 1 mm.

Type Species: By original designation, Penaeus indicus H. Milne Edwards, 1837, Hist. Nat.

Crust., 2: 415.

Type Locality: Coast of Coromandel, India.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum armed with dorsal and 2-5 ventral teeth, long in

young, conspicuously surpassing antennular flagella, rather short in adult, reaching between

midlength and about distal end of second antennular segment. Carapace with antennal and hepatic

spines pronounced, lacking orbital and pterygostomian spines; postocular sulcus absent;

postrostral carina short to rather long, reaching as far as posterior one-eighth of carapace;

adrostral carina and sulcus short, falling distinctly short or extending to about level of epigastric

tooth; gastrofrontal carina absent; gastro-orbital carina usually absent or feeble, but occasionally

rather well defined, if present occupying middle one-third or not more than posterior two-thirds

of distance between hepatic spine and orbital margin; orbito-antennal sulcus often shallow or

poorly marked; cervical carina well defined, accompanying sulcus well defined or feeble; hepatic

carina more often absent, if present, moderately or ill-defined; branchiocardiac carina lacking;

longitudinal and transverse sutures absent. Sixth abdominal somite bearing three cicatrices!

lacking dorsolateral sulcus. Telson unarmed.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Antennule lacking parapenaeid spine; antennular flagella much shorter than carapace. Palp of

first maxilla elongate, slender, either biarticulate, or consisting of 3 or 4 articles, large basal

article produced in setose proximal lobules on lateral and mesial margins, bearing 1 long

mesiodistal spine, and elongate distolateral patch of spinules. Basial and ischial spines on first

pereopod, only basial spine on second.

Petasma symmetrical, semiclosed, with distomedian projections usually quite short, ventral

costae long, strongly curved and gaping distally. Appendix masculina subelliptical or,

occasionally, broadly subtriangular and armed with closely set marginal spines and sometimes

with submarginal patches of spinules. Thelycum closed, with pair of lateral plates on stemite

XIV forming thick mesial lips meeting along midline, shielding simple, median, saclike seminal

receptacle; stemite Xm with well developed median protuberance.

Distribution of branchiae, epipods, and exopods in the genus Fenneropenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 42. — Fenneropenaeus indicus : A, 3 32 mm cl, off Yemen, Red Sea. Petasma, ventral view. Scale = 1 mm.

— B, 9 46 mm cl, Joseph Bonaparte Gulf, Northern Territory, Australia. Thelycum. Scale = 5 mm.

The seminal receptacles and spermatophores of Fenneropenaeus merguiensis were described

in detail by Tuma (1967). The formation and structure of the spermatophores of Fenneropenaeus

Source:

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

indicus were described by Champion (1987). They are similar to those of the species of genera

having a closed thelycum previously included in Penaeus s.l., with the exception of

Marsupenaeus.

Species. — Fenneropenaeus chinensis (Osbeck, 1765). Hong Kong; China; Korea; Japan.

Fenneropenaeus indicus (H. Milne Edwards, 1837). South-east and east Africa to Red Sea;

Madagascar; Yemen to southern China; Philippines; Japan; New Guinea; northern Australia.

Fenneropenaeus merguiensis (De Man, 1888). Persian Gulf; Pakistan; Western and southern

India; Sri Lanka; Malaysia; Singapore; Thailand; Indonesia; Gulf of Tonkin; South China Sea;

Philippines; Hong Kong; New Guinea; northern Australia; Fiji; New Caledonia.

Fenneropenaeus penicillatus (Alcock, 1905). Pakistan; India; Indonesia; China; Taiwan.

Fenneropenaeus silasi (Muthu and Motoh, 1979a). Singapore; Borneo; Sarawak.

Genus Funchalia Johnson, 1867

Figures 43-45

Funchalia Johnson, 1867, Proc. zool. Soc. Lond., 1867: 895. — Bouvier, 1908b, Bull. Inst. oceanogr. Monaco,

119: 3; 1908a, Result. Camp, scient. Prince Albert I, 33: 91. — Lenz and Strunck, 1914, Dt. Siidpol.-

Exped., 15, Zool. 7: 303. — Balss, 1925, Wiss. Ergebn. Dt.-Tiefsee Exped. Valdivia, 20(5): 227. — Calman,

1925, Rep. Fish. mar. biol. Surv. Un. S. Afr., 4, spec. rep. 3: 10. — Burkenroad, 1934b, Bull. Am. Mus.

nat. Hist., 68(2): 73-77; 1936b, Bull. Bingham oceanogr. Coll., 5(2): 126. — Roger, 1938, Bull. Soc. Zool.

Fr., 63: 23. — Barnard, 1950, Ann. S. Afr. Mus., 38: 608. — Gordon and Ingle, 1956, J. mar. biol. Ass.

U. K„ 35: 475 [part]. — Dall, 1957, Aust. J. mar. Freshwat. Res., 8(2): 163 [part], — Crosnier and Forest,

1973, Faune Trop., 19: 296 [part]. — Grippa, 1976, Atti Soc. ital. Sci. nat., 117(3/4): 124. — Burkenroad,

1983, Crust. Issues, 1: 282. — De Freitas, 1987, Investl. Rep. Oceanogr. Res. Inst., Durban, 58: 3. —

Fig. 43. — Funchalia woodwardi, 9 35.5 mm cl, southeast Atlantic Ocean off South Africa. Lateral view

Scale = 10 mm.

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Grippa, 1987, Investnes Pesq., 51 (suppl. 1): 73. — Wasmer, 1989, J. nat. Hist., 23: 476. — Dall et al.,

1990, Adv. Mar. Biol., 27: 66. — Burukovsky and Romensky, 1991, Biul. Moskovsk. Obsh. Ispyt. Prir.: 60.

Placed on the Official List of Generic Names in Zoology, International Commission of Zoological

Nomenclature, 1969, Opinion 864, Name No. 1810, Bull. zool. Norn., 25(4/5): 139.

Sicyonia - G£ny, 1873, Ann. Soc. Lettres Sci. Arts Alpes-Maritimes, 2: 162 [part],

Aristeus - Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 309 [part], — Senna, 1903, Boll. Soc. ent

ital., 34(4): 261 [part].

Penaeus - Alcock, 1905, Ann. Mag. nat. Hist., (7)36(95): 513 [part].

Grimaldiella Bouvier, 1905a, C. r. hebd. Seanc. Acad. Sci., Paris, 140: 981, 982. Type species: By monotypy,

Grimaldiella richardi Bouvier, 1905a, C. r. hebd. S6anc. Acad. Sci., Paris, 140: 982. (= Funchalia woodwardi

Johnson, 1867). Type locality: Off lie Saint-Michel, Azores.

Hemipenaeopsis Bouvier, 1905a, C. r. hebd. Seanc. Acad. Sci., Paris, 140: 981. Type species: By monotypy,

Hemipenaeopsis villosus Bouvier, 1905a, C. r. hebd. Seanc. Acad. Sci., Paris, 140: 981. Type locality: North¬

east Atlantic Ocean, between Canary Islands and Azores. Gender: Feminine.

Funchalia (Funchalia) - Burkenroad, 1934b, Bull. Am. Mus. nat. Hist., 68(2): 73, 76; 1936, Bull. Bingham

oceanogr. Coll., 5(2): 126.

Funchalis - Burkenroad, 1936b, Bull. Bingham oceanogr. Coll., 5(2): 136.

Type Species: By monotypy, Funchalia woodwardi Johnson, 1867, Proc. zool. Soc. Lond.,

1867: 895.

Type Locality: Off Madeira, north-east Atlantic Ocean.

Gender: Feminine.

Diagnosis. — Integument pubescent. Rostrum short, compressed, reaching as far

as midlength of second antennular segment, armed with dorsal teeth only. Carapace lacking

orbital spine (angle only occasionally slightly marked), with antennal and pterygostomian spines,

hepatic spine present or absent; postocular sulcus lacking;

gastro-orbital carina absent; orbito-antennal and cervical sulci

Fig. 44. — Funchalia woodwardi ,

9 35.5 mm cl, southeast Atlan¬

tic Ocean off South Africa.

A, Mandible, ventral view of left

side. Scale = 1 mm. B, Cutting

edge of mandible enlarged. Scale

= 0.5 mm.

weak or absent; postrostral carina pronounced or feeble;

antennal, pterygostomian, and branchiocardiac carinae well

marked; dorsolateral carina lacking; hepatic carina present or

lacking; without longitudinal and transverse sutures. Fourth,

fifth, and sixth abdominal somites bearing continuous,

prominent cicatrix. Telson with three pairs of fixed subapical

spines, preceded by much smaller ones.

Antennule lacking parapenaeid spine; antennular flagella

subfiliform, long, dorsal one as long as or longer than

carapace and slightly longer than ventral. Mandibles with

incisor processes extremely elongate, scythelike and crossing

posterior to labrum, ventral cutting edge minutely denticulate,

submarginally striate; distal article of mandibular palp large,

asymmetrical, truncate. Palp of first maxilla relatively long,

slender, triarticulate, basal article armed with minute

proximomesial lobe, distolateral row of spinules, and long

subdistal spine on mesial margin; distal article bearing long

apical setae. Dactyl of third maxilliped sexually dimorphic;

broadened and roughly spoon-shaped in males, narrow and

subcylindrical in females. Ridge of lateral ramus of uropod

produced into conspicuous spine. Basial and ischial spines on

first and second pereopods. Epipods strongly forked.

ISABEL pgREZ FARFANTE & BRIAN KENSLEY

Petasma semi-open, asymmetrical with either right or left half longer than opposite. Appendix

masculina subrectangular. Thelycum open, with anterior depression on stemite XIV flanked by

pair of large broad flaps or narrow ones bearing strong ridges; median protuberance on stemite

XIII deeply excavate, delimiting depression on stemite XIV.

Distribution of branchiae, epipods, and exopods in the genus Funchalia.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(Pi)

(P2)

(P3)

(P4)

XIV

(PS)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

“

■

Epipods

Exopods

Fig. 45. — Funchalia woodwardi: A, 6 37.5 mm cl, southeast Atlantic Ocean off South Africa. Petasma, dorsal

view. — B, $ 35.5 mm cl, southeast Atlantic Ocean off South Africa. Thelycum. Scales = 1 mm.

Remarks. — Funchalia sagamiensis was described from a single juvenile specimen (7 mm

carapace length and of undetermined sex), which possesses three characters that differ from those

typical of the members of Funchalia : it lacks exopods on all pereopods, basial and ischial spines

on the first and second pereopods, and the telson bears two pairs of movable lateral spines

anterior to the fixed subapical pair; this juvenile has not been associated with an adult. Hayashi

(1992) found a small female (8 mm carapace length) in Japanese waters which shares many

characters with F. sagamiensis, but it bears small prominences on the pereopods which resemble

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

buds of exopods. Although Hayashi tentatively assigned it to F. sagamiensis he suggested that

the specimen might possibly belong to F. villosa or F. taaningi. Because the members of

Funchalia undergo considerable changes in their development, the identities of these two juvenile

shrimps must await the study of additional, more nearly adult material.

Species. — Funchalia danae Burkenroad, 1940. Azores; Off Canary Islands; Saint Helena

Island; Congo; Arabian Sea; south-central Indian Ocean; north-west Pacific Ocean.

IFunchalia sagamiensis Fujino, 1975. Japan.

Funchalia taaningi Burkenroad, 1940. Madagascar; western and eastern Indian Ocean;

Indonesia; Japan; central and north-west Pacific Ocean.

Funchalia villosa (Bouvier, 1905a). Caribbean; Canary Islands; Mediterranean; southern

Atlantic Ocean off Tristan da Cunha; Antarctic; south-west Indian Ocean from Agulhas Bank to

Natal, South Africa; eastern Indian Ocean; Lord Howe Island and off New South Wales,

Australia.

Funchalia woodwardi Johnson, 1867. Outer Hebrides, north-east Atlantic Ocean; Azores;

Madeira; Mediterranean; south-east Atlantic Ocean off Cape of Good Hope, South Africa;

Arabian Sea; New South Wales, Australia.

Genus Heteropenaeus De Man, 1896

Figures 46-47

Heteropenaeus De Man, 1896, Zool. Anz., 19(498): 111; 1898, Zool. Jahrb., 10: 684. — Nobili, 1903, Boll.

Musei Zool. Anat. comp. R. Univ. Torino, 18(455): 4. — Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 515;

1906, Cat. Indian Dec. Crust., 3(1): 16. — De Man, 1924, Arch. Naturgesch., 90 A (2): 30. — Balss, 1957,

Bronn's Kl. Ordn. Tierreichs, 5(1)7: 12: 1518. — Hall, 1961, Bull. Raffles Mus., 26: 83. — Dall et al.,

1990, Adv. Mar. Biol., 27: 68.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1815, Bull. zool. Nom., 25(4/5): 139.

Heteropeneus - Burkenroad, 1983, Crust. Issues, 1: 282.

Type Species: By monotypy, Heterooenaeus longimanus De Man, 1896, Zool. Anz., 19(498):

111 .

Type Locality: Java Sea.

Gender: Masculine.

Diagnosis. — Integument of carapace and abdomen bearing rows of densely-set short stiff

variously-directed setae, mostly projecting from one margin of numerous sulci of varying length.

Rostrum armed with dorsal teeth along entire length, ventral teeth on anterior half, moderately

long, reaching or slightly overreaching antennular peduncle. Carapace with acutely pointed

antennal and hepatic spines; orbital and pterygostomian spines lacking; unique subrostral sulcus 1

present; orbito-antennal sulcus well defined only posteriorly. Postrostral carina well defined to

near posterior margin of carapace; short gastro-orbital and antennal carina present; cervical carina

sharp, sulcus deep; hepatic carina and sulcus well marked, extending anteroventrally below

hepatic spine then curving ventrally, also extending posterodorsally to about midlength of

'While this sulcus is in the postocular position, its unusual length and peculiar armature suggest that it is not

homologous with the postocular sulci of the rest of the 'tribe'. The term 'subrostral sulcus' may be more

appropriate.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

cervical sulcus; branchiocardiac carina and sulcus clearly defined; longitudinal and transverse

sutures absent. Abdomen with numerous, variously directed small grooves; sixth abdominal

somite without cicatrices, bearing long subdorsolateral sulcus. Telson with four pairs of

relatively long movable lateral spines.

Fig. 46. — Heteropenaeus longimanus, 2 12 mm cl, Masinloc Bay, Luzon, Philippines. Lateral view.

Scale = 10 mm.

Eye with basal plate produced into sharp spine. Antennule lacking parapenaeid spine;

antennular flagella short, less than half length of carapace. Mandible with incisor process short,

molar process broad, thick. Palp of first maxilla long, triarticulate, basal article with broad

proximolateral and smaller proximomesial setose lobules, bearing spine just distal to

proximomesial lobe and comb of subdistolateral spinules; distal article elongate, slender, bearing

five long slender spines. First pereopod in adult males sometimes stouter and considerably longer

than second and third; fingers of chela half length of propodal palm in male, subequal in length to

propodal palm in female. Basial and ischial spines on first pereopod, only basial spine on

second.

Petasma symmetrical, semi-open, not hooded and with long ventral costae widely gaping,

dorsolateral lobule extending apically into acute slender projection. Appendix masculina ovate,

with submesial setose rib where turning mesially at right-angle. Thelycum open, with paired,

strongly elevated (ventrally), lateral plates flanking deep depression on sternite XIV; median

protuberance on sternite XIII strongly excavate, forming roughly inverted V-shaped ridge,

crowning depression of XIV.

Remarks. — The extraordinarily elongate first pereopod was described by De Man (1896) for

the syntypic male from the Java Sea. This feature has since not been commented on nor

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

illustrated, neither was it seen in the seven males we have examined. However, Alain Crosnier

(in litt., 1996) assures us that Heteropenaeus longimanus can be bought by the kilogram in fish

markets in the Philippines, and that only very large males, which are not all that common,

possesses the very long first pereopods. He quotes a specimen with a carapace length of

17.5 mm as having a first pereopod of 66 mm, with an extremely elongate propod.

Distribution of branchiae, epipods, and exopods in the genus Heteropenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(PS)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 47. — Heteropenaeus longimanus : A, 2 12 mm cl, Tumindao Island, Sulu Archipelago, Philippines.

Petasma, dorsal view. — B, $ 12 mm cl, Masinloc Bay, Luzon, Philippines. Thelycum. Scales = 1 mm.

Species. —Heteropenaeus longimanus De Man, 1896. Singapore; Java, Temate, Indonesia;

Philippines; Japan; New Guinea; Australia; Palau; Caroline Islands.

Genus Litopenaeus Perez Farfante, 1969

Figures 48-49

Cancer - Linnaeus, 1767, Syst. Nat. (12th ed.), 1(2): 1054 [part],

Penaeus - Smith, 1885b, Proc. U. S. natn. Mus., 8(11/12): 170 [part]; 1886, Rep. U. S. Commnr Fish., for

1885: 684 [part]. — De Man, 1911, Siboga Exped., 39a: 95 [part]. — Burkenroad, 1934b, Bull. Am. Mus.

Source:

ISABEL PFRF7. FARFANTE & BRIAN KENSLEY

nat. Hist., 68(2): 74 [part]. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 268 [pan]. — Dall, 1957, Austr. J.

mar. Freshwat. Res., 8(2): 14 [part].

Panaeus - Thallwitz, 1892, Abh. Ber. zool. antrop.-ethn. Mus. Dresden, 1890-91, 3(3): 3. [Panaeus gracilirosiris

= p. setiferus according to Burkenroad, 1939, Bull. Bingham oceanogr. Coll., 6(6): 17].

Peneus - Alcock, 1905, Ann. Mag. nat. Hist., 7(16): 513 [part]; 1906, Cat. Indian Dec. Crust., 3(1): 7 [part],

Penaeus (Litopenaeus) P6rez Farfante, 1969, Fish. Bull., U. S., 67(3): 466. — Burukovsky, 1972, Trudy

AtlantNIRO, 42: 5, 8, 13, 15. — PfiREZ Farfante, 1975, Fish. Bull., U. S., 73(3): 463; 1988, NOAA Tech.

Rep., NMFS, 64: 9.

Fig. 48. — Litopenaeus setiferus, 2 38 mm cl, off Long Beach, North Carolina. Lateral view. Scale = 10 mm.

Type Species: By original designation, Penaeus vannamei Boone, 1931, Bull. Am. Mus. nat.

Hist., 63(2): 173.

Type Locality: Panama [Golfo de Panama].

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum armed with dorsal and usually, 2-4

(occasionally 1, 5-8) ventral teeth, moderately long, in young distinctly surpassing antennular

peduncle, shorter in adult, sometimes reaching only midlength of second antennular segment.

Carapace with antennal and hepatic spines pronounced, lacking orbital and pterygostomian

spines; postocular sulcus absent; postrostral carina variable in length, sometimes almost reaching

posterior margin of carapace; adrostral carina and sulcus short, extending to, or only slightly

beyond epigastric tooth; gastrofrontal carina absent; gastro-orbital carina relatively short, usually

extending (at most) anteriorly about two-thirds of distance between hepatic spine and orbital

margin; orbito-antennal sulcus well marked; cervical and hepatic carinae sharp, accompanying

sulci deep; branchiocardiac carina lacking; longitudinal and transverse sutures absent. Sixth

abdominal somite bearing three cicatrices, dorsolateral sulcus extremely narrow or absent. Telson

unarmed.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Antennule lacking parapenaeid spine; antennular flagella much shorter than carapace. Palp of

first maxilla elongate, consisting of 3 or 4 articles, distal ones together flagelliform; basal article

produced in setose proximal lobes on lateral and mesial margins, bearing 1 or 2 long distomesial

spines, and distolateral row of spinules. Basial and ischial spines on first pereopod, basial on

second.

Petasma symmetrical, semi-open, not hooded, lacking distomedian projections; ventral costae

short, not nearly reaching distal margin, and distinctly gaping. Thelycum open, stemite XIV

bearing ridges, prominences, depressions, or grooves. Spermatophores extremely complex,

consisting of a sperm mass encapsulated by a sheath and bearing various attachment structures

(anterior wing, lateral flap, caudal flange, dorsal plate), as well as adhesive and glutinous

materials.

Detailed descriptions of the spermatophores of all Litopenaeus species were given by P£rez

Farfante (1975), and that of Litopenaeus stylirostris by Cardenas Figueroa (1952). Studies of

the formation and structure of the spermatophore of L. setiferus were made by Bauer (1991),

and Chow et al. (1990); the latter also discussed the spermatophore of L. vannamei.

Distribution of branchiae, epipods, and exopods in the genus Litopenaeus.

Somite

(Mxpl)

Vffl

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 49. — Litopenaeus setiferus : A, 6 40 mm cl, off Galveston, Texas. Petasma, dorsal view. Scale = 1 mm. —

B, $ 47 mm cl, off Galveston, Texas. Thelycum. Scale = 10 mm.

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Species. —Litopenaeus occidentalis (Streets, 1871). Chiapas, Gulf of Tehuantepec, Mexico;

El Salvador; Gulf of Panama; Colombia to northern Peru; Galapagos Islands.

Litopenaeus schmitti (Burkenroad, 1936a). Cuba to Trinidad; Belize; Caribbean coast of

Central and South America to southern Brazil.

Litopenaeus setiferus (Linnaeus, 1767). Fire Island and Nyack, (Hudson River), New York

to Florida; Gulf of Mexico to Yucatan.

Litopenaeus stylirostris (Stimpson, 1874). Western Baja California Norte; Gulf of California

to northern Peru.

Litopenaeus vannamei (Boone, 1931). Gulf of California to northern Peru.

Genus Macropetasma Stebbing, 1914

Figures 50-51

Parapeneus - Balss, 1913b, in Schultze, Forschungsreise Sildafr., 5 (2), Jena Denkschr., 17: 105.

Macropetasma Stebbing, 1914, Ann. S. Afr. Mus., 15(1): 22. — Burkenroad, 1934a, Bull. Bingham oceanogr.

Coll., 4(7): 8; 1934b, Bull. Am. Mus. nat. Hist., 68(2): 73, 117. — Barnard, 1947, Ann. Mag. nat. Hist.,

(11)13: 382; 1950, Ann. S. Afr. Mus., 38: 605. — Holthuis, 1962, Bull. zool. Norn., 19(2): 108. —

Burkenroad, 1983, Crust. Issues, 1: 282. — Dall et al ., 1990, Adv. Mar. Biol., 27: 70.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1818, Bull. zool. Nom., 25(4/5): 139.

Fig. 50. —Macropetasma africana, $ 10.5 mm cl, Durban, South Africa. Lateral view. Scale = 10 mm.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Type Species: By monotypy, Parapeneus africanus Balss, 1913b, in Schultze, Forschungs-

reise Stidafr., 5(2), Jena Denkschr., 17: 105.

Type Locality: Swakopmund, Namibia.

Gender: Neuter.

Diagnosis. — Integument minutely and sparsely setose.

Opaque-spot photophores present on abdomen. Rostrum armed

with dorsal teeth only, in male short, reaching at most, to distal

margin of first antennular segment, in female extending to mid¬

length of third segment; epigastric tooth distinctly separated from

first rostral tooth. Carapace lacking orbital spine, with small an¬

tennal spine (subjacent margin usually produced into small pro¬

jection), and well developed hepatic spine; pterygostomian and

branchiostegal spines absent; postrostral carina lacking; postocular

sulcus well defined, antennal carina, gastrofrontal and orbito-

antennal sulci lacking, cervical sulcus well marked but short,

hepatic sulcus horizontal, shallow, not marked anterior to hepatic

spine; hepatic carina lacking; branchiocardiac carina feeble,

sometimes indistinct; longitudinal and transverse sutures absent.

Third abdominal somite expanded in caridean humpback fashion;

sixth somite lacking cicatrices; photophores on abdominal somites,

telson, uropods, and pleopod segments, their presence unique

within the family. Telson armed with four pairs of movable lateral

spines.

Fig. 51. — Macropetasma africana: A, 6 9.5 mm cl, off Great Fish Point, South Africa. Petasma, ventral

view. —- B, 9 10.5 mm cl, off Great Fish Point, South Africa. Thelycum. Scales = 1 mm.

Antennule lacking parapenaeid spine; antennular flagella subequal in length, long, longer than

carapace, sexually dimorphic: in males, mesial flagellum depressed and bearing broad tooth

ISABEL P£REZ FARFANTE & BRIAN KENSLEY

proximally on mesial margin. Palp of first maxilla entire, short, without proximolateral lobule,

with broad setose proximomesial and minute distomesial lobules. In females, first and second

pereopods lacking coxal, basial, and ischial spines; in males, second pereopod with coxal ridge

on mesial margin, and very small, proximally situated tooth on ischium, third pereopod with

coxa flattened, bearing blunt tooth on mesial margin, and small backwardly-directed proximal

tubercle on ischium. ,

Petasma (articulated close to proximal extremity of basis of endopod instead of more distally

as in other members of family) symmetrical, semiclosed, with lateral lobes uniquely

produced into extremely long, basally expanded then filamentous processes; dorsomedian

lobules terminating in well developed but relatively short distomedian projections. Appendix

masculina roughly bell-shaped; endopod much reduced. Male genital openings subcoxal.

Thelycum open, with paired lateral plates on sternite XIV sinuous, expanded into fleshy,

bulbous swelling anteriorly and forming narrow arcs flanking deep bowl-like depression

posteriorly; median protuberance on sternite XIII small, suboval to inverted heart-

shaped, slightly concave, with narrow posteromedian groove continuous with depression of

sternite XIV.

Distribution of branchiae, epipods, and exopods in the genus Macropetasma.

Somite

(Mxpl)

vrn

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Species. — Macropetasma africana (Balss, 1913b). Kunene River mouth, northern

Namibia, to Zululand, South Africa.

Genus Marsupenaeus Tirmizi, 1971

Figures 52-54

Peneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 513 [part]; 1906, Cat. Indian Dec. Crust., 3(1): 7 [part].

Penaeus - De Man, 1911, Siboga Exped., 39a: 95 [part]. — Burkenroad, 1934b, Bull. Am. Mus. nat. Hist.,

68(2): 74 [part], — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 268, 270A [part], — Dall, 1957, Aust. J. mar.

Freshwat. Res., 8(2): 141 [part].

Penaeus <Melicertus ) - P£rez Farfante, 1969. Fish. Bull., U. S., 67(3): 466 [part]. — Burukovsky, 1972, Trudy

AllantNIRO, 42: 5, 8, 17.

Penaeus (Marsupenaeus) Tirmizi, 1971, Pakist. J. Zool., 2(2): 193. — Liu and Zhong, 1988, Penaeoid Shr. S.

China Sea,: 105. — Hayashi, 1992, Dendr. Crust. Jap. Waters, : 298.

Type Species: By monotypy, Penaeus canaliculatus var. japonicus Bate, 1888, Rep. scient.

Results Voy. Challenger, 24: 245.

Type Locality: Japan.

Gender: Masculine.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Fig. 52. — Marsupenaeus japonicus, 2 56.5 mm cl, Tokyo, Japan. Lateral view. Scale = 10 mm.

Diagnosis. — Integument glabrous. Rostrum armed with dorsal and 1 (occasionally 2) ventral

teeth, long in young, conspicuously surpassing antennular peduncle, decreasing in length with

size, in adult reaching as little as distal end of first antennular segment. Carapace with spiniform

orbital angle; antennal and hepatic spines very pronounced, lacking pterygostomian spine;

postocular sulcus absent; postrostral carina and adrostral sulcus and carina long, almost reaching

posterior margin of carapace; gastrofrontal carina turning anterodorsally upon itself at posterior

end, short ramus projecting from base conspicuously separated from carina, gastrofrontal sulcus

markedly bifid posteriorly; gastro-orbital carina

long, almost reaching anterior margin of carapace;

orbito-antennal sulcus well defined; cervical carina

sharp, accompanying sulcus well marked; hepatic

carina and sulcus well marked; branchiocardiac

carina lacking; longitudinal and transverse sutures

absent. Sixth abdominal somite bearing three

cicatrices, lacking dorsolateral sulcus. Telson armed

with three pairs of movable lateral spines.

Antennule lacking parapenaeid spine; antennular

flagella much shorter than carapace. Palp of first

maxilla elongate, usually triarticulate, sometimes

consisting of 4 or 5 articles, basal article produced

into setose proximal lobule on lateral and mesial

margins, bearing one mesial spine and distolateral

row of spinules. Basial spines on first and second

pereopods.

Petasma symmetrical, semiclosed, with dis-

tomedian projections greatly developed and curved, forming hood. Appendix masculina subel¬

liptical, armed with strong marginal spines and broad patch of short strong ones. Thelycum

closed, with single plate on stemite XIV infolded laterally forming anteriorly open pocket, latter

Fig. 53. — Marsupenaeus japonicus, 6 45 mm

cl, Indian Ocean. Petasma. A, Distal part of

right side, dorsolateral view. B, Distal part of

right side, ventrolateral view. Scale = 1 mm.

Source:

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

functioning as seminal receptacle; stemite XHI bearing elongate platelike median protuberance

extending well into stemite XIV.

Hudinaga (1942) and Tirmizi (1958) gave a detailed account of the structure of the thelycum

and seminal receptacle of the monotypic M. japonicus, and included a study of the

spermatophore, the latter consisting of sperm sac and, when deposited on female, an unique large

subtriangular process or "wing", with anteriorly disposed broad base which, together with long

posteriorly curved dorsal rod, anchors spermatophore to pocket. Medina et al. (1994) reported

on the spermatozoal ultrastructure in M. japonicus.

Distribution of branchiae, epipods, and exopods in the genus Marsupenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

Xffl

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 54. — Marsupenaeus japonicus : A, 3 45 mm cl, Indian Ocean. Petasma, ventral view. — B, $ 42 mm cl,

Tokyo, Japan. Thelycum. Scales = 1 mm.

Species. —Marsupenaeus japonicus (Bate, 1888). Eastern Mediterranean; East coast of South

Africa; East Africa; Madagascar; Mauritius; Reunion; Red Sea; Persian Gulf; India; Malaysia;

Singapore; Thailand; Indonesia; South China Sea; Philippines; Taiwan; Korea; Japan; New

Guinea; Northern Territory, Queensland, Australia; Fiji.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Genus Megokris new genus

Figures 55-58

Penaeus - Haswell, 1879, Proc. Linn. Soc. N.S.W., 4: 38 [part, description of Penaeus granulosus only].

Peneus (Trachypeneus) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 15.

Trachypeneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 522 [part]. — Schmitt, 1926, Zool. (biol.) Results

Fish. Exp. 'Endeavour', 5(6): 348 [description of P. granulosus only]. — Burkenroad, 1983, Crust. Issues, 1:

283 [part].

Trachypenaeus - De Man, 1911, Siboga Exped., 39a: 87 [part]. — Kubo, 1949, J. Tokyo Fish., 36(1): 391 [part].

— Dall, 1957, Aust. J. mar. Freshwat. Res., 8(2): 202 [part]. — Racek and Dall, 1965, Verh. K. Akad.

Wet. Nat., 56: 87 [part]. — Racek and Yaldwyn, 1971, Proc. Linn. Soc. N.S.W., 95(3): 213. — Motoh and

Buri, 1984, Res. Crust., (13-14): 80. — Liu and Zhong, 1988, Penaeoid Shr. S. Ch. Sea, : 184 [part]. —

Dall etaU 1990, Adv. Mar. Biol., 27: 102 [part].

Trachypeneus (Trachypeneus) - Burkenroad, 1934a [September], Bull. Bingham oceanogr. Coll., 4(7): 49

[Division II (part)].

Trachypeneus - Burkenroad, 1934b [December], Bull. Am. Mus. nat. Hist., 68(2): 94 [Division 2 (part)].

Fig. 55. — Megokris granulosus , $ 26 mm cl, north-northeast of Bowen, Queensland, Australia. Lateral view.

Scale = 10 mm.

Type Species: By present designation, Penaeus granulosus Haswell, 1879, Proc. Linn. Soc.

N.S.W., 4: 41.

Type Locality: Australia, Damey Island, Torres Strait.

Gender: Masculine.

Source

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Etymology. — From the Greek 'mega' - large, and 'okris' - a projection, alluding to the

extraordinary posterior extension of the median protuberance of stemite XIII.

Diagnosis. — Body thickset. Integument thick, densely pubescent. Rostrum short, reaching

between distal margin of first antennular segment and about end of peduncle; armed with dorsal

teeth only; epigastric tooth distinctly separated from first rostral. Carapace with orbital angle

sharp or armed with small spine; antennal and hepatic spines present; pterygostomian angle blunt

or acute- postocular sulcus shallow or obsolete; gastro-orbital carina lacking; orbito-antennal

sulcus feeble or lacking; cervical sulcus short; hepatic sulcus extending just ventral to hepatic

region or farther anteroventrally; branchiocardiac carina usually absent, occasionally clear y

distinct; longitudinal suture short, ending well anterior to hepatic spine; transverse suture usual y

clearly distinct. Abdomen with sixth somite lacking cicatrix. Telson with 3 pairs of movable

lateral spines. „ , , „ i c r .

Antennule lacking parapenaeid spine; antennular flagella shorter than carapace. Palp of first

maxilla entire, distally tapering to acute apex, produced into small setose proximolateral, broad,

setose proximomesial and small acute distomesial lobules, latter bearing 1-3 sharp spines and

armed with distoventral row of spinules. Fourth and fifth pereopods with dactyl neither elongate

nor subdivided. Basial spine lacking on third maxilliped, present on first and second pereopods;

small ischial spine on first pereopod, lacking on second. Epipod on first to third pereopods 2 , or

on third only.

Fig. 56. — Megokris granulosus , 8 11 mm cl,

Manifa, Saudi Arabia. Petasma, dorsal view.

Scale = 1 mm.

Petasma symmetrical, semiclosed, variable in

shape, with lateral lobes distolaterally produced into

either moderately broad to narrow, roughly hornlike

projections extending laterally and with hook-like

tips curving distally or distomesially, or extremely

broad basally but narrowing rapidly, ending in

distally directed tips. Appendix masculina

subquadrangular with rounded corners. Thelycum

closed, with plate of sternite XIV deeply excavate

anteriorly, extremely short posteromedially,

expanding laterally, flanking strikingly large,

sometimes tongue-like, caudal extension of median

protuberance; mesial margins of plate in contact

with, or narrowly separated from protuberance, or

forming narrow flaps; anterior sternal invagination

(other than seminal receptacles) lacking, thus neither

transverse groove nor median pocket present;

median protuberance uniquely extending caudally

for almost entire length of stemite XIV.

Seminal receptacles consisting of paired bilobed

sacs, large posterior lobe and small anterolateral

one, with strongly curved slitlike opening lying on

anteromesial side of large lobe, joining cavity dorsal

to and at about midlength of median protuberance,

near base of fourth pereopod.

2 A few specimens of M. gonospinifer were found to lack an epipod on the third pereopod (Racek and Dall,

1965; Racek and Yaldwyn, 1971); further studies of the species are needed to determine the significance of these

observations.

Source . MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

Distribution of branchiae, epipods, and exopods in the genus Megokris.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

1 .

Arthrobranchiae

Podobranchiae

Epipods

1 (-)

Exopods

Fig. 57. — Megokris granulosus, 2 25 mm cl, north-northeast of Bowen, Queensland, Australia: A, Thelycum.

B, Seminal receptacles, dorsal view. Scales = 1 mm.

Remarks. — Burkenroad (1934a) divided Trachypenaeus in two subgenera:

Trachysalambria, "Division I", proposed by him, possessing an epipod on the first and second

pereopods and a pair of seminal receptacles opening mesially in an unpaired pocket into which

sperm-free male secretion is received; and Trachypenaeus s.s., "Division II", lacking an epipod

on the first and second pereopods and, apparently, a median pocket. Furthermore, Burkenroad

(1934a, b) subdivided Division I, Trachysalambria, into two sections: Section 1, possessing a

longitudinal suture extending behind the hepatic spine, basial spine on the third maxilliped,

lacking an ischial spine on the first pereopod, and having the posterior lip of the transverse

groove anterodorsal to the plate of sternite XIV, from which the median pocket is invaginated,

produced in a pair of free flaps; and Section 2, exhibiting a longitudinal suture not extending

behind the hepatic spine, lacking basial spine on the third maxilliped, possessing an ischial spine

on the first pereopod and the posterior lip of the transverse groove not produced as paired flaps.

The characters of these Divisions and Sections, cited by Burkenroad in 1934a, appeared in

another paper of his (1934b) only three months after the former was published.

Later, it was noticed by various authors (Kubo, 1949; Dall, 1957; Burkenroad, 1959) that

epipods on the first and second pereopods were also absent in some members of

Trachysalambria, Section 2, and, consequently, this taxon was not accepted. Kubo (1949)

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

rejected those subgenera but, as Burkenroad (1959) noted, for erroneous reasons: he mistook

Section 2 of Division I for Division II. Burkenroad (1934a) selected T. (Trachysalambria)

curvirostris as the type of this subgenus, including it in his Section 2 of Division I; in 1934b, he

further characterized Section 2.

Fig. 58 A-B. — Megokris pescadoreensis , 2 23.5 mm cl, 11 km north of Corregidor Island, Manila Bay,

Philippines: A, Thelycum. B, Seminal receptacles, dorsal view. Scales = 1 mm.

Fig. 58 C-D. — Megokris sedili, 2 23 mm cl, Chinde, Zambezia, Mozambique: C, Thelycum. D, Seminal

receptacles, dorsal view. Scales = 1 mm.

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

As a result of a detailed study of Trachypenaeus s.l., we are of the opinion that in this

extremely heterogeneous group, four different genera should be recognized. They can be

separated mainly by the structures of the genitalia, particularly those of the thelycum. One of

them ( Rimapenaeus , proposed herein) can be distinguished also by the length of the longitudinal

suture of the carapace which is long rather than short, by the presence of a basial spine on the

third maxilliped, and by possessing a trilobed, instead of a bilobed, seminal receptacle. The

occurrence of an epipod on the first and second pereopods varies in Megokris and

Trachysalambria but seems to be constant in Rimapenaeus and is lacking in the monotypic

Trachypenaeus, s.s. Finally, in two genera, Megokris and Trachypenaeus, the seminal

receptacles open independently, with no connecting sternal groove or median pocket. In

Trachysalambria, however, they open into a common transverse groove (from which they

actually originate) and in Rimapenaeus they open mesially into a common median pocket. The

four genera are discussed below; their diagnoses may be found under the relevant generic

headings.

Megokris comprises one of the species groups previously included in Trachypenaeus, and

constitutes part of Burkenroad's (1934a) Division II. Although it most closely resembles the

latter genus, as restricted herein, it differs in that the thelycal plate of stemite XIV is deeply

excavate, its posteromedian part is extremely short, and extends anterolaterally embracing the

typical caudal projection of the median protuberance. In contrast, in the species of Trachypenaeus

s.s., the plate of stemite XIV is moderately long medially and the median protuberance is short,

not invading the plate. Furthermore, in Megokris, the distolateral projections of the petasma are

either moderately broad to narrow and extend laterally or they are extremely broad basally,

narrowing rapidly, all ending in small hooklike tips; the distolateral projections in Trachypenaeus

are extremely broad throughout their length and end in an elongate flattened and twisted narrow

process directed almost straight forward. In both these genera the only sternal invaginations are

those giving rise to the seminal receptacles, thus these are neither connected by a transverse

groove, as they do in Trachysalambria, nor by a median pocket as they are in Rimapenaeus.

Species. — Megokris gonospinifer (Racek and Dali, 1965). Arafura Sea; west Irian Jaya,

Indonesia; Papua New Guinea; Northern Territory, Australia.

Megokris granulosus (Haswell, 1879). Saudi Arabia; Persian Gulf; Pakistan; South India; Sri

Lanka; Malaysia; Indonesia; Philippines; Taiwan; New Guinea; Queensland, Australia.

Megokris pescadoreensis (Schmitt, 1931). Persian Gulf; Malaysia; Singapore; China;

Philippines; Hong Kong; Taiwan.

Megokris sedili (Hall, 1961). Mozambique; India; Sri Lanka; Myanmar; Singapore; Straits of

Malacca; Gulf of Thailand; Gulf of Tonkin; South China Sea.

Genus Melicertus Rafinesque-Schmaltz, 1814

Figures 59-60

Cancer - ForskAl, 1775, Descr. Anim.,: 95 [part].

Melicertus Rafinesque-Schmaltz, 1814, Prec. Decouv. Trav. Somiol., :22.

Penaeus - H. Milne Edwards, 1837, Hist. Nat. Crust., 2: 414. — De Man, 1911, Siboga Exped., 39a: 95 [part],

— Burkenroad, 1934b, Bull. Amer. Mus. nat. Hist., 68(2): 74 [part]. — Kubo, 1949, J. Tokyo Coll. Fish.,

36(1): 268 [part], — Dall, 1957, Aust. J. mar. Freshwat. Res., 8(2): 141 [part].

Peneus - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 13 [part]; 1905, Ann. Mag. nat. Hist., (7)16: 513

[part]; 1906, Cat. Indian Dec. Crust., 3(1): 7 [part].

Salambria "Burkenroad" Balss, 1957, Bronn's Kl. Ordn. Tierreichs, 7(1)12: 1518 [nomen nudum].

100

ISABEL PFRFZ FARFANTE & BRIAN KENSLEY

Penaeus (Melicertus) - PErez Farfante, 1969, Fish, Bull., U. S„ 67(3): 466 [pan], — Burukovsky, 1972, Trudy

AtlantNIRO, 42: 6, 10, 14, 17. — Liu and Zhong, 1988, Penaeoid Shr. S. China Sea, : 105. — Hayashi,

1992, Dendr. Crust. Japan. Waters,: 122.

Fig. 59. — Melicertus kerathurus, 2 47 mm cl, El Maghreb, Tunisia. Lateral view. Scale = 10 mm.

Type Species: By monotypy, Melicertus tigrinus Rafinesque-Schmaltz, 1814: 22 (= Cancer

kerathurus Forskal, 1775).

Type Locality: Sicily.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum with dorsal and usually 1 ventral tooth (bearing

2 or 3 only in M. marginatus). Carapace with well marked orbital angle; antennal and hepatic

spines strong, lacking pterygostomian spine; postocular sulcus absent; postrostral carina and

adrostral carina and sulcus long, extending considerably beyond epigastric tooth, almost always

nearly to posterior margin of carapace; gastrofrontal carina turning anterodorsally upon itself at

posterior end, and with short ramus projecting anteriorly from base ventrally close to carina;

gastro-frontal sulcus not distinctly bifid posteriorly; gastro-orbital carina long, at least three-

quarters of distance between hepatic spine and orbital margin; orbito-antennal sulcus well

marked; cervical carina sharp, accompanying sulcus deep; hepatic carina and sulcus well marked;

branchiocardiac carina lacking; longitudinal and transverse sutures absent. Sixth abdominal

somite bearing three cicatrices, lacking dorsolateral sulcus. Telson usually armed with two or

three pairs of lateral movable spines (absent only in M. canaliculatus).

Antennule lacking parapenaeid spine; antennular flagella much shorter than carapace. Palp of

first maxilla elongate, consisting of 3 or 4 articles, distal ones together flagelliform; large basal

article produced in setose lateral and mesial lobules proximally, bearing 1 long, subdistal spine

and distolateral row of spinules. Basial spine on first and second pereopods reduced, ischial

spine occasionally present on first.

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

101

Petasma symmetrical, semiclosed, with distomedian projections short to moderately long;

ventral costae long, gently to strongly curved distally. Appendix masculina subelliptical, armed

with numerous marginal setae and, usually, broad distal patch. Thelycum closed, with paired

lateral plates on sternite XIV shielding simple median saclike seminal receptacle; median

protuberance on sternite XIII well developed. Spermatophores quite similar to those of

Farfantepenaeus species.

Observations on the structure of the spermatophore of Melicertus kerathurus were made by

Heldt (1938a, b), under the name Penaeus trisulcatus, while Malek and Bawab (1974a, b),

under P. kerathurus, studied its formation in detail. The spermatozoal ultrastructure of

M. kerathurus was reported by Medina et al., 1994.

Distribution of branchiae, epipods, and exopods in the genus Melicertus.

Somite

(Mxpl)

Vffl

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 60. — Melicertus kerathurus: A, 6 44 mm cl, El Maghreb, Tunisia. Petasma, dorsal view. Scale = 1 mm.

— B, 2 47 mm cl, El Maghreb, Tunisia. Thelycum. Scale = 5 mm.

Species. — Melicertus canaliculatus (Olivier, 1811). East coast of South Africa to Red Sea;

Madagascar; Mauritius; Reunion; Gulf of Oman; India; Sri Lanka; Bangladesh; Malaysia;

Indonesia; Thailand; South China Sea; Philippines; Taiwan; Japan; New Guinea; Northern

Queensland, Australia; Loyalty Islands; Fiji.

102

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Melicertus hathor Burkenroad, 1959. South-east coast of Africa to Red Sea; Madagascar;

Mauritius; Reunion; Seychelles; Persian Gulf; Gulf of Oman; Pakistan; South India; Sri Lanka;

Myanmar.

Melicertus kerathurus (Forskal, 1775). Mediterranean; eastern Atlantic Ocean from southern

England to Angola.

Melicertus latisulcatus (Kishinouye, 1896). Singapore; Malaysia; Thailand; Indonesia; South

China Sea; Philippines; Japan; Hong Kong; Korea; New Guinea; Western Australia, Northern

Territory to New South Wales, Australia; Fiji.

Melicertus longistylus (Kubo, 1943). Malaysia; Thailand; Singapore; South China Sea;

Philippines; Taiwan; Japan; Western Australia, Northern Territory, Queensland, New South

Wales, Australia.

Melicertus marginatus (Randall, 1840). East Africa; Madagascar; Arabian Sea; Bay of Bengal;

Singapore; Indonesia; Taiwan; South China Sea; Japan; Queensland and Western Australia,

Australia; Hawaii.

Melicertus plebejus (Hess, 1865). Southern Queensland to Victoria, Australia.

Genus Metapenaeopsis Bouvier, 1905

Figures 61-62

Penoeus - De Haan, 1844, Fauna Jap. (Crust.), (6, 7): pi. 46, fig. 3-5 [part].

Penaeus - De Haan, 1849, Fauna Jap. (Crust.), (6): 190-193.

Parapenaeus - Smith, 1885b, Proc. U.S. natn. Mus., 8(11/12): 170 [part].

Metapenaeus - Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 271 [part].

Metapenaeopsis Bouvier, 1905a, C. r. hebd. Seanc. Acad. Sci., Paris, 140: 981. — Kubo, 1949, J. Tokyo Coll.

Fish., 36(1): 408. — Dall, 1957, Aust. J. mar. Freshwat. Res., 8(2): 166. — Rucek and Dall, 1965, Verh.

K. Akad. Wet., 56(3): 17. — P£rez Farfante, 1971a, Smithson. Contrib. Zool., 79: 2. — Crosnier, 1987,

Bull. Mus. Hist. nat. Paris, (4)9, sect. A (2): 409. — Dall et al., 1990, Adv. Mar. Biol., 27: 71. —

Crosnier, 1991, Mem. Mus. Hist. nat. Paris, (A)152: 155. — Hayashi, 1992, Dendr. Crust. Japan. Waters, :

79. — Crosnier, 1994a, Mem. Mus. Hist. nat. Paris, 161: 258.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1819, Bull. zool. Nom., 25(4/5): 139.

Archipenaeopsis Bouvier, 1905b, C. r. hebd. Seanc. Acad. Sci., Paris, 141: 747. Type species: By monotypy,

Archipenaeopsis vestitus Bouvier, 1905b, C. r. hebd. Seanc. Acad. Sci., Paris, 141: 747 [= Metapenaeopsis

goodei (Smith, 1885b)]. Type locality: "Mer des Antilles" [actually, NW of Dry Tortugas Islands, 24°43'N,

83°25'W], 68 m, Blake sta 11 [data from label accompanying holotype. Museum of Comparative Zoology,

Harvard University, 7204].

Parapenaeopsis - Bouvier, 1905b, C. r. hebd. Seanc. Acad. Sci., Paris, 141: 748.

Metapeneus - Alcock, 1906, Cat. Indian Dec. Crust., 3(1): 16 [part].

Penaeopsis - Bouvier, 1908b, Bull. Inst, oceanogr., Monaco, 119: 5 [part]. — Barnard, 1950, Ann. S. Afr.

Mus., 38: 592. — Holthuis, 1952a, Result, scient. Exped. oceanogr. beige (1948-1949), 3(2): 80.

Leptopenaeus Kishinouye, 1929, Proc. imp. Acad. Japan, 5(7): 282. Type species: By present designation,

Leptopenaeus philippinensis (= Penaeus philippii Bate, 1881). Type locality: Philippines, 180 m.

Ceratopenaeus Kishinouye, 1929, Proc. imp. Acad. Japan, 5(7): 282. Type species: By present designation,

Ceratopenaeus mogiensis (= Parapenaeus mogiensis Rathbun, 1902). Type locality: Japan.

Erythropenaeus Kishinouye, 1929, Proc. imp. Acad. Japan, 5(7): 283. Type species: By present designation,

Erythropenaeus akayebi (= Parapenaeus akayebi Rathbun, 1902). Type locality: Japan.

[Penaeopsis] (Metapenaeopsis) - Burkenroad, 1934a, Bull. Bingham oceanogr. Coll., 4(7): 8.

Metapeneopsis - Burkenroad, 1983, Crust. Issues, 1: 282.

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

103

Fig. 61. — Metapenaeopsis miersi, $ 19 mm cl, off Mauritania. Lateral view. Scale = 10 mm.

Type Species: By monotypy, Metapenaeopsis pubescens Bouvier, 1905a, C. r. hebd. Seanc.

Acad. Sci., Paris, 140: 981 [not Penaeus pubescens Stimpson, 1871] = Penaeopsis miersi

Holthuis, 1952a, Exped. Oceanogr. beige Eaux Cot. Afr. Atlant. S. (1948-49), 3(2): 80.

Type Locality: Cape Verde Islands [Canal de St Vincent a St. Antoine, lies du Cap Vert,

16°06'N, 24°00'W, 137-165 m, Talisman sta. 115].

Gender: Feminine.

Diagnosis. — Integument thick, pubescent, usually densely so. Rostrum armed with dorsal

teeth only; variable in length, usually not overreaching antennular peduncle; epigastric tooth

conspicuously separated from first rostral tooth. Carapace with small orbital, and well developed

antennal, pterygostomian, and hepatic, spines, lacking branchiostegal spine; postocular sulcus

absent; orbito-antennal and cervical sulci usually feeble; hepatic sulcus usually weak, sometimes

indistinct anteriorly, never reaching pterygostomian angle; hepatic carina indistinct; sutures

lacking. Stridulating ridges on posterior parts of carapace present in some species. Sixth

abdominal somite without cicatrices. Telson armed with pair of fixed subapical spines preceded

by three pairs of conspicuous movable spines.

Antennule with parapenaeid spine, sometimes vestigial; antennular flagella variable in length,

usually much shorter but sometimes longer than carapace; sexually dimorphic, longer in males

than in females. Palp of first maxilla entire, short, often narrowing to blunt apex, bearing small

proximolateral lobule, conspicuous proximomesial and smaller distomesial lobules, latter armed

with 1-3 slender spines, and distolateral row of spinules. Basial spine on third maxilliped, first

and second pereopods, always lacking on third. Rudimentary anterior arthrobranchia present on

somite XIII.

Petasma asymmetrical, divided at about midlength into proximal and distal parts, latter

complex, subdivided into several projections and components, one (in Atlantic, eastern Pacific

and single Indo-West Pacific species) or two (in other Indo-west Pacific species) of these

104

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

lamellate distoventral projections, more or less covering others. Appendix masculina trumpet¬

shaped (small abrupt expansion of dorsomesial rib of endopod). Thelycum closed, with

transverse plate on sternite XIV and strongly developed median protuberance on sternite XIII;

sometimes intermediate plate present between latter and transverse plate on XIV.

Seminal receptacle not saclike, usually consisting of bulbous capsule situated dorsal to

anterolateral part of sternite XIV (dorsal to thelycal hood), continuous with horizontal or

obliquely directed narrow troughlike duct (fissure ventromesial), which sometimes turning

anteriorly in U and extending for variable length along sternite XIII. Apparently, sperm enters

through the duct fissure and flows into the capsule and exits through a slit (continuation of the

duct fissure) on the anterodorsal side of the capsule, at the base of the fourth pereopod. The

seminal receptacles of M. goodei and of various Western Atlantic species were described and

illustrated by Burkenroad (1934a) and P£rez Farfante (1971a), respectively.

Distribution of branchiae, epipods, and exopods in the genus Metapenaeopsis*

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

i+r

Podobranchiae

Epipods

Exopods

* Crosnier (1991) has pointed out that this formula is found in most species of this genus, but that there are

some exceptions in the number or development of the branchiae.

SPECIES. — Metapenaeopsis acclivis (Rathbun, 1902). Taiwan; Japan.

Metapenaeopsis aegyptia Galil and Golani, 1990. Eastern Mediterranean; Red Sea;

Madagascar; Seychelles; Arabian Sea; Maidive Islands; Sri Lanka; Japan; Chesterfield Islands;

New Caledonia.

Metapenaeopsis andamanensis (Wood-Mason, 1891). Southern India; Andaman Islands;

Straits of Malacca; Malaysia; South China Sea; Taiwan; Japan.

Metapenaeopsis angusta Crosnier, 1987. Singapore; Vietnam; Brunei; Philippines.

Metapenaeopsis assimilis (De Man, 1920). Indonesia; New Guinea.

Metapenaeopsis barbata (De Haan, 1844). Persian Gulf; Bay of Bengal; Malaysia; Indonesia;

Taiwan; Japan.

Metapenaeopsis beebei (Burkenroad, 1938). Western Baja California Sur, Gulf of California

to Michoacan, Mexico; Costa Rica; Gulf of Panama; Galapagos Islands.

Metapenaeopsis ceylonica Starobogatov, 1972. Natal, South Africa; Mozambique;

Madagascar; India; Maidive Islands; Sri Lanka; Indonesia; Philippines; Taiwan; New Guinea;

New Caledonia.

Metapenaeopsis commensalis Borradaile, 1898. Maidive Islands; Laccadive Islands;

Indonesia; Philippines; Taiwan; northwestern Australia; Torres Strait; New Caledonia; Ellice

Islands.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

105

Fig. 62. — Metapenaeopsis miersi : A-B, 6 12.5 mm cl, off Mauritania. A, Petasma, ventral view. B, Petasma,

dorsal view. — C, $ 19 mm cl, off Mauritania. Thelycum. — D, $ 17.5 mm cl, Ivory Coast. Seminal

receptacles, dorsal view. Scales = 1 mm.

Source:

106

ISABEL P£REZ FARFANTE & BRIAN KENSLEY

Metapenaeopsis coniger (Wood-Mason, 1891). Gulf of Aden; Gulf of Oman; India; Bay of

Bengal; Japan.

Metapenaeopsis costata C rosnier, 1991. Philippines.

Metapenaeopsis crassissima Racek and Dali, 1965. Western Australia, Northern Territory,

South Australia, Australia.

Metapenaeopsis dalei (Rathbun, 1902). Gulf of Tonkin; Taiwan; China; Japan; Korea.

Metapenaeopsis difficilis Crosnier, 1991. Indonesia; Philippines; Coral Sea; New Caledonia;

Marquesas Islands; Wallis and Futuna Islands.

Metapenaeopsis distincta (De Man, 1907). Indonesia; New Britain; New Caledonia; Palau;

Bikini Atoll, Marshall Islands; Wallis and Futuna Islands.

Metapenaeopsis dura Kubo, 1949. Taiwan; Japan.

Metapenaeopsis erythraea Crosnier, 1987. Red Sea.

Metapenaeopsis everrnanni (Rathbun, 1906). Indonesia; Chesterfield Islands; New Caledonia;

Molokai Island, Hawaii.

Metapenaeopsis faouzii (Ramadan, 1938). Seychelles; Maidive Islands.

Metapenaeopsis fusca R.J.G. Manning, 1988. Western Australia; New Guinea.

Metapenaeopsis gaillardi Crosnier, 1991. Kenya; southern India; Sri Lanka; Indonesia;

Chesterfield Islands; New Caledonia; Molokai Island, Hawaii.

Metapenaeopsis gallensis (Pearson, 1905). Tanzania; Madagascar; India; Maidive Islands; Sri

Lanka; Malaysia; Indonesia; Philippines; northwestern Australia; New Caledonia.

Metapenaeopsis gerardoi Perez Farfante, 1971a. Florida Keys; West Indies; Caribbean coast

of Central and South America; St. Helena Island.

Metapenaeopsis goodei (Smith, 1885b). North Carolina to Florida, Gulf of Mexico to

Pensacola; Yucatan; Bahamas; Caribbean Islands and coast of Central and South America to

Bahia, Brazil; Bermuda.

Metapenaeopsis hilarula (De Man, 1911). Natal, South Africa; Mozambique; Madagascar;

Gulf of Aden; Seychelles; Reunion; Maidive Islands; Sri Lanka; Singapore; Malaysia; Indonesia;

Philippines; New Guinea; Queensland, Australia; New Caledonia; Chesterfield Islands; Tahiti.

Metapenaeopsis hobbsi Perez Farfante, 1971a. West Indies; Caribbean coast of Central and

South America to Espfrito Santo, Brazil.

Metapenaeopsis incisa Crosnier, 1991. lies Glorieuses; Maidive Islands.

Metapenaeopsis ivanovi Crosnier, 1994b. Off Kenya.

Metapenaeopsis kishinouyei (Rathbun, 1902). Western Baja California Norte to tip of Baja

California Sur, Gulf of California to Jalisco, Mexico; Rocas Alijos; Islas Revillagigedo;

Clipperton Island; Panama; Galapagos Islands.

Metapenaeopsis kyushuensis (Yokoya, 1933). Japan.

Metapenaeopsis lamellata (De Haan, 1844). Malaysia; Indonesia; Thailand; Philippines;

Taiwan; Japan; China; Western Australia, Queensland, Northern Territory, Australia; New

Caledonia.

Metapenaeopsis lata Kubo, 1949. South China Sea; Japan.

Metapenaeopsis laubieri Crosnier, 1991. Malaysia; Indonesia; Philippines.

Metapenaeopsis lindae R.J.G. Manning, 1988. Western Australia.

Metapenaeopsis liui Crosnier, 1987. Indonesia; Philippines; north-western and northern

Australia.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

107

Metapenaeopsis mannarensis De Bruin, 1965. Sri Lanka; Indonesia; northwestern Australia;

Chesterfield Island; New Caledonia.

Metapenaeopsis manningi Crosnier, 1994b. Off Somalia.

Metapenaeopsis marquesas Crosnier, 1991. New Caledonia; Marquesas Islands.

Metapenaeopsis martinella Perez Farfante, 1971a. Antilles, Western Caribbean to

Pernambuco, Brazil.

Metapenaeopsis menoui Crosnier, 1991. Seychelles; Indonesia; Philippines; New Guinea;

Northwestern Australia; New Caledonia.

Metapenaeopsis miersi Holthuis, 1952a. Mauritania to Angola.

Metapenaeopsis mineri Burkenroad, 1934a. West coast of Baja California Sur, and Gulf of

California, Mexico; Panama.

Metapenaeopsis mogiensis complanata Crosnier, 1991. Northern Australia; Chesterfield

Islands; Bellona Island; New Caledonia.

Metapenaeopsis mogiensis consobrina (Nobili, 1904). Natal, South Africa; Djibouti, East

Africa; Red Sea; Gulf of Aden; Madagascar; Seychelles; Persian Gulf; India; Sri Lanka; South

China Sea; Indonesia.

Metapenaeopsis mogiensis intermedia Crosnier, 1991. Indonesia; Thailand; South China Sea;

Philippines; Taiwan.

Metapenaeopsis mogiensis mogiensis (Rathbun, 1902). Japan.

Metapenaeopsis novaeguineae (Haswell, 1879). New Guinea; Western Australia, Northern

Territory and Queensland, Australia.

Metapenaeopsis palmensis (Haswell, 1879). Singapore; Malaysia; Thailand; Vietnam;

Indonesia; Philippines; Taiwan; Japan; New Guinea; Western Australia, Northern Territory,

Queensland, and New South Wales, Australia.

Metapenaeopsis parahilarula Crosnier, 1991. Philippines.

Metapenaeopsis parapalmensis Crosnier, 1994a. Indonesia; Philippines.

Metapenaeopsis persica Crosnier, 1991. Saudi Arabia, Persian Gulf.

Metapenaeopsis philippii (Bate, 1881). Indonesia; Philippines; Admiralty Islands.

Metapenaeopsis propinqua Crosnier, 1991. New Caledonia.

Metapenaeopsis provocatoria longirostris Crosnier, 1987. Taiwan; Japan; Korea; Western

Australia.

Metapenaeopsis provocatoria provocatoria Racek and Dali, 1965. Indonesia; Philippines;

Queensland, Australia; New Caledonia; Wallis and Futuna Islands.

Metapenaeopsis proximo C rosnier, 1991. Seychelles.

Metapenaeopsis quadrilobata Crosnier, 1991. Nosy Be, Northwestern Madagascar.

Metapenaeopsis quinquedentata (De Man, 1907). Tanzania; Indonesia; Philippines; New

Caledonia.

Metapenaeopsis richeri Crosnier, 1991. Indonesia; Landsdowne-Fairway Bank; Chesterfield

Islands.

Metapenaeopsis rosea Racek and Dali, 1965. New Guinea; Queensland and Northern

Territory, Australia.

Metapenaeopsis scotti Champion, 1973. Natal, South Africa; Off Tanzania; Madagascar;

Reunion.

Metapenaeopsis sibogae (De Man, 1907). Indonesia; Philippines.

108

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Metapenae.opsis sinica Liu and Zhong, 1988. Malaysia; Indonesia; South China Sea;

Philippines; Taiwan; Japan; Queensland, Australia; Chesterfield Islands; New Caledonia.

Metapenaeopsis sinuosa Dali, 1957. Northern Territory and Queensland, Australia.

Metapenaeopsis smithi (Schmitt, 1924). South-east Florida to Tortugas Islands; Arrecife

Alacran, Mexico; West Indies; Caribbean coast of Central and South America.

Metapenaeopsis spatulata Crosnier, 1991. Philippines.

Metapenaeopsis spiridonovi Crosnier, 1991. Seychelles.

Metapenaeopsis stokmani Burukovsky, 1990. South-east Pacific Ocean near Sala-y-Gomez.

Metapenaeopsis stridulans (Alcock, 1905). Gulf of Aden; Persian Gulf; Gulf of Oman;

Arabian Sea; India; Maidive Islands; Sri Lanka; Bay of Bengal; Malaysia; Singapore; Thailand;

Vietnam; South China Sea; Philippines; Indonesia; Chesterfield Islands; New Caledonia.

Metapenaeopsis tarawensis Racek and Dali, 1965. Maidive Islands; Cocos Keeling Island;

Northern Australia; New Caledonia; Caroline Islands; Gilbert Islands; Tuamoto, Polynesia.

Metapenaeopsis tchekunovae Starobogatov, 1972. Gulf of Tonkin.

Metapenaeopsis tenella Liu and Zhong, 1988. South China Sea.

Metapenaeopsis toloensis Hall, 1962. Arabian Sea; Maidive Islands; Sri Lanka; Andaman Sea;

Bay of Bengal; Malaysia; Thailand; Indonesia; Gulf of Tonkin; South China Sea; Philippines;

New Guinea; Australia; Chesterfield Islands.

Metapenaeopsis vaillanti (Nobili, 1904). Suez Canal; Red Sea; Gulf of Aden; Arabian Sea.

Metapenaeopsis velutina (Dana, 1852a). Seychelles; South China Sea; Philippines;

Queensland and northwestern Australia; New Caledonia; Chesterfield Islands; Hawaii; Pitcairn

Island.

Metapenaeopsis wellsi Racek, 1967. Western Australia and northwestern Australia.

Genus Metapenaeus Wood-Mason, 1891

Figures 63-65

Penaeus - Miers, 1878, Proc. zool. Soc. London,: 301 [part].

Mangalura Miers, 1878, Proc. zool. Soc. London, : 303. — Burkenroad, 1959, Miss. Dollfus Egyp., Rdsult.

Scient., 3: 86. — Holthuis, 1962, Bull. zool. Nom., 19(2): 106. — Burkenroad, 1963a, Bull. zool. Nom.,

20(3): 169; 1963b, Bull zool. Nom., 20(4): 247; 1983, Crust. Issues, 1: 280. — Holthuis, 1963, Bull. zool.

Nom., 20(4): 246. — Burkenroad, 1983, Crust. Issues, 1: 280, 282.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1830, Bull. zool. Nom., 25 (4/5): 140 "Ruled under the plenary

powers not to be given precedence over Metapenaeus Wood-Mason, 1891." Type species: By monotypy,

Mangalura dobsoni Miers, 1878, Proc. zool. Soc. Lond., 1878: 302, pi. 17, fig. 2. Type locality: Mangalur

(Mangalore), west of India. Gender: Feminine.

Metapenaeus Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 271: [part]. — Burkenroad, 1934a, Bull. Bingham

oceanogr. Coll., 4(7): 4, 7, 29. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 327. — Barnard, 1950, Ann. S.

Afr. Mus., 38: 596. — Dall, 1957, Aust. J. mar. Freshwat. Res., 8(2): 182. — Holthuis, 1962, Bull. zool.

Nom., 19(2): 106. — Burkenroad, 1963b, Bull. zool. Nom., 20(4): 169; 1963c, Bull. zool. Nom., 20(4):

247. — Holthuis, 1963, Bull. zool. Nom., 20(4): 245. — Racek and Dall, 1965, Verh. K. Akad. Wet.,

56(3): 54. — Miquel, 1982, Zool. Verh., Leiden, 195: 5. — Motoh and Buri, 1984, Res. Crust., (13/14): 51.

— Dall et al., 1990, Adv. Mar. Biol., 27: 79.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1829, Bull. zool. Nom., 25 (4/5): 140. "Ruled under the plenary

powers to be given precedence over Mangalura Miers, 1878."

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

109

Peneus (Metapeneus) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 14.

Metapeneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 516 [part]; 1906, Cat. Indian Dec. Crust., 3(1): 16

[part].

Penaeopsis. — A. Milne Edwards and Bouvier, 1909, Mem. Mus. comp. Zool. Harv., 37(3): 220 [part], —

De Man, 1911, Siboga Exped., 39a: 53 [part]. — Schmitt, 1926, Zool. (biol.) Results Fish. Exp.

'Endeavour', 5(6): 319, [part].

Fig. 63. — Metapenaeus affinis, 2 35.5 mm cl, Orissa coast, India. Lateral view. Scale = 10 mm.

Type Species: By original designation: Penaeus affinis H. Milne Edwards, 1837, Hist. Nat.

Crust., 2: 416.

Type Locality: Cote de Malabar (now Kerala), southwest of India.

Gender: Masculine.

Diagnosis. — Integument variably pubescent, sometimes almost entirely glabrous. Rostrum

armed with dorsal teeth only and varying considerably in length; epigastric tooth often

conspicuously separated from first rostral tooth. Carapace with orbital angle usually sharp, but

lacking spine, antennal and hepatic spines pronounced; pterygostomian margin rounded, lacking

spine; postocular sulcus deep; gastro-orbital carina absent; orbito-antennal and cervical sulci well

marked; hepatic sulcus anterior to hepatic spine well defined and accompanied by ventral carina

often descending almost vertically from spine then turning towards pterygostomian angle, sulcus

posterior to hepatic spine ill-defined or absent; branchiocardiac carina variably developed, from

strong, long or short to indistinct; longitudinal and transverse sutures absent. Sixth abdominal

somite bearing single continuous cicatrix, or latter interrupted into two or three parts. Telson

lacking fixed subapical spines but bearing movable, sometimes minute and very numerous,

posterolateral ones.

Source:

110

ISABEL PIjREZ FARFANTE & BRIAN KENSLEY

Fig. 64. — Metapenaeus

affinis, 6 30 mm cl, Orissa

coast, India. Petasma, ventral

view. Scale = 1 mm.

Antennule lacking parapenaeid spine; antennular flagella shorter

than carapace. Palp of first maxilla biarticulate, distal article minute,

large basal article bearing weak proximolateral lobule and large and

small proximomesial lobules, latter armed with long mesial spine.

Basial spine on first through third pereopods, in some species

ischial spine on first. Fifth pereopod modified in male: ischium

usually bearing lateral keel, and merus exhibiting proximal

notch followed by conspicuous knob or spiniform process.

Pleurobranchia present on somite XIII. Exopod lacking on fifth

pereopod.

Petasma symmetrical, semiclosed, depressed, with median lobes

usually produced into simply curved, hoodlike, or convoluted distal

projections; sclerotized lateral lobes produced distally in spoutlike

obliquely or fully lateral projections and with ventrolateral recurved,

flaplike to complex, mesial process. Appendix masculina narrow

basally, greatly expanded distally and strongly convex ventrally.

Thelycum closed, with paired lateral plates of sternite XIV often

continuous across sternite, usually more or less enveloping

posterior end of elongate median protuberance of sternite XIII.

Seminal receptacles, for example in M. affinis and M. ensis ,

consisting of paired posteriorly rigid, anteriorly membranous

pyriform sacs, continuous basally and opening through sternal

transverse slit between sternites XIV and XIII; sacs extending

anteriorly into sternite XIII and opening through paired slits lying

lateral to posterior part of median protuberance, dorsal to base of

fourth pereopods. A general account of the seminal receptacles was

given by Burkenroad (1934a) and more recently Hassan (1981)

described in detail the development and structure of the seminal

receptacles of M. affinis.

Distribution of branchiae, epipods, and exopods in the genus Metapenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

XII

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

l+v

Podobranchiae

Epipods

Exopods

Species. — Metapenaeus affinis (H. Milne Edwards, 1837). Persian Gulf; Arabian Sea, from

Gulf of Oman to South India; Sri Lanka; Andaman Islands; Malaysia; Singapore; Borneo;

Thailand; Gulf of Tonkin; South China Sea; Philippines; Hong Kong; Taiwan; New Guinea;

Hawaii.

Metapenaeus alcocki George and Rao, 1968. Gulf of Kutch, India.

Metapenaeus anchistus (De Man, 1920). Malaysia; Borneo; Thailand; Philippines; Fiji; Wallis

and Futuna Islands.

Metapenaeus bennettae Racek and Dali, 1965. Queensland, New South Wales, Australia.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

111

Fig. 65. — Metapenaeus affinis, 2 35.5 mm cl, Orissa coast, India. A, Thelycum. B, 2 39 mm cl, Gulf of

Kutch, India. Seminal receptacles, dorsal view. Scales = 1 mm.

Metapenaeus brevicomis (H. Milne Edwards, 1837). Pakistan to Mangalore, India; Bay of

Bengal; Malaysia; Singapore; Borneo; Thailand; Vietnam.

Metapenaeus conjunctus Racek and Dali, 1965. Singapore; Malaysia; Indonesia; North

Borneo; Philippines; Northern Territory, Australia.

Metapenaeus dalli Racek, 1957. Indonesia; Philippines; Western Australia, Northern

Territory, Australia.

Metapenaeus demani demani (Roux, 1921). New Guinea; Northern Territory, Australia.

Metapenaeus demani stephani Miquel, 1982. New Guinea; Queensland, Australia.

Metapenaeus dobsoni (Miers, 1878). Western India; Sri Lanka; Andaman Islands; Malaysia;

Gulf of Thailand; Indonesia; Philippines; New Guinea.

Metapenaeus eboracensis Dali, 1957. New Guinea; Northern Territory, Queensland,

Australia.

Metapenaeus elegans De Man, 1907. Sri Lanka; Andaman Islands; Malaysia; Borneo;

Thailand; Philippines; New Guinea; Fiji; Wallis and Futuna Islands.

Metapenaeus endeavouri (Schmitt, 1926). New Guinea; Western Australia, Northern

Territory, Queensland, New South Wales, Australia.

Metapenaeus ensis (De Haan, 1844). India; Bangladesh; Sri Lanka; Bay of Bengal; Malacca

Straits; Malaysia; Indonesia; Borneo; Gulf of Tonkin; South China Sea; Philippines; Hong Kong;

Taiwan; Japan; New Guinea; Western Australia, Northern Territory, Queensland, New South

Wales, Australia; Wallis and Futuna Islands.

Metapenaeus insolitus Racek and Dali, 1965. Northern Territory, Queensland, Australia.

Metapenaeus intermedius (Kishinouye, 1900). Andaman Islands; Singapore; Malaysia;

Borneo; Gulf of Tonkin; South China Sea; Hong Kong; Taiwan; Japan.

112

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Metapenaeus joyneri (Miers, 1880). Gulf of Tonkin; South and East China; Taiwan; Japan;

Korea.

Metapenaeus krishnatrii Silas and Muthu, 1976. Andaman Islands.

Metapenaeus kutchensis George, George and Rao, 1963. Gulf of Kutch, India.

Metapenaeus lysianassa (De Man, 1888). India; Sri Lanka; Bay of Bengal; Singapore; North

Borneo; Thailand; Vietnam.

Metapenaeus macleayi (Haswell, 1879). Queensland, New South Wales, Australia.

Metapenaeus monoceros (Fabricius, 1798). Eastern Mediterranean; Red Sea; East coast of

South Africa; Mozambique; Madagascar; Reunion; Mauritius; Saudi Arabia; Pakistan; India; Sri

Lanka; Burma; Malaysia.

Metapenaeus tnoyebi (Kishinouye, 1896). South India; Sri Lanka; Bay of Bengal; Malaysia;

Borneo; Thailand; China; Phillipines; Taiwan; Japan.

Metapenaeus papuensis Racek and Dali, 1965. Thailand; Philippines; New Guinea.

Metapenaeus stebbingi Nobili, 1904. Eastern Mediterranean; Southeast African coast;

Mozambique; Gulf of Suez; Red Sea; Madagascar; Yemen; Saudi Arabia; Persian Gulf; Pakistan;

Northern India.

Metapenaeus suluensis Racek and Dali, 1965. Gulf of Thailand; Philippines; New Guinea.

Metapenaeus tenuipes Kubo, 1949. Malaysia; Singapore; North Borneo; Thailand;

Philippines; Taiwan.

Genus Miyadiella Kubo, 1949

Figures 66-67

Penaeus - Stimpson, 1860, Proc. Acad. nat. Sci. Philad., 12: 43.

Miyadiella Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 105, 264. — Holthuis, 1955b, Vidensk. Medd. Dansk

Naturh. Foren., 117: 75. — Starobogatov, 1972, Akad. Nauk SSSR Zool. Inst. Isled. Fauney Moreii,

10(18): 369. — Holthuis, 1980b, FAO Fish. Synop., 125, 1: 13. — Burkenroad, 1983, Crust. Issues, 1:

287. — Liu and Zhong, 1988, Pen. Shr. S. China Sea, : 102. — Dall et al, 1990, Adv. Mar. Biol., : 27. —

Hayashi, 1992, Dendr. Crust. Jap. Waters,: 72.

Type Species: By original designation, Miyadiella pedunculata Kubo, 1949, J. Tokyo Coll.

Fish., 36(1): 264 [= Penaeus podophthalmus Stimpson, 1860, Proc. Acad. Nat. Sci. Philad.,

12: 43].

Type Locality: Osaka-wan, Japan.

Gender: Feminine.

Diagnosis. — Integument thin, studded with microscopic setae. Rostrum short, not reaching

distal margin of first antennular segment, with dorsal teeth only; epigastric tooth markedly

separated from first rostral tooth. Carapace with orbital angle rounded, lacking spine; antennal

spine present; hepatic spine present or absent; lacking pterygostomian spine; postrostral carina

extending to about midlength or three-fourths of carapace, bearing rows of minute spinules,

which also present on first few rostral teeth; postocular sulcus well marked; cervical sulcus short

or long, almost reaching middorsum of carapace, accompanying carina sharp; hepatic sulcus and

carina clearly distinct anteriorly, there extending anteroventrally, both weak or indistinct

posteriorly; branchiocardiac carina long but feeble; longitudinal and transverse sutures lacking.

Sixth abdominal somite without cicatrices. Telson lacking movable posterolateral spines, or

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

113

armed with three pairs of small marginal, well-spaced ones, with few simple submarginal setae,

and considerably elongate, plumose marginal setae subapically.

Eye extremely elongate, reaching to between midlength of second antennular segment and

about midlength of third (first and second antennular segments considerably elongate).

Parapenaeid spine absent; stylocerite produced into sharply pointed spine and proximomesially

bearing small sharply pointed scale. Antennular flagella subequal or dorsal flagellum slightly

Fig. 66 A. — Miyadiella podophthalmus, 2 6.5 mm cl, East China Sea. Lateral view. Scale = 1 mm.

Fig. 66 B. — Miyadiella ornata, 2 holotype, 38 mm tl, northeast of Java. Lateral view (carapace slightly

displaced dorsally). Scale = 5 mm.

114

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

longer than ventral, moderately long, half to two-thirds as long as carapace. Palp of first maxilla

entire, produced into minute setose proximolateral lobe and broad setose proximomesial lobe

followed by sharp spine. Basial spine present or absent, ischial present on first pereopod, basial

and ischial on second and third. Fifth pereopod reaching about midlength of scaphocerite.

Petasma semi-open, symmetrical, with minute cincinnuli along dorsomesial margin; median

and lateral lobes simple, not folded, median bearing moderately broad proximal projection and

lateral one, long rather flexible costa. Appendix masculina subelliptical, very slightly convex

dorsally and bearing row of small marginal setae. Thelycum open, possessing basally broad

anteromedian spine on stemite XIV, preceded by rounded scales or possessing elongate flaplike

lateral plates (free along most of their length); median protuberance of sternite XIII either short,

not reaching stemite XIV, or extending well into latter.

Distribution of branchiae, epipods, and exopods in the genus Miyadiella.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

Xffl

(P4)

xrv

(P5)

Pleuro branchiae

Arthrobranchiae

i (->

Podobranchiae

Epipods

Exopods

Species. — Miyadiella podophthalmus (Stimpson, 1860). Maharashtra, west coast of India;

South China Sea; Japan.

Miyadiella omata Holthuis, 1955b. Northeast Java and Arafura Sea, Indonesia.

Remarks. — The generic status of Miyadiella pedunculata Kubo, 1949, and Miyadiella omata

Holthuis, 1955b, has been the object of two interpretations: that they belong to Atypopenaeus

Alcock, 1905, or that they should be retained in Miyadiella Kubo, 1949.

Alcock (1905) was the first to suggest that Penaeus stenodactylus Stimpson, 1860, and

P. podophthalmus Stimpson, 1860, might belong to the genus Atypopenaeus. Starobogatov

(1972) having the same opinion, recognized Penaeus podophthalmus and Miyadiella pedunculata

Kubo, 1949, as synonyms of A. stenodactylus and assigned it and M. ornata Holthuis, 1955b

(which he considered to be the young A. dearmatus De Man, 1907) to the genus Atypopenaeus.

Burkenroad (1983) also suggested such a tentative synonymy for the genus, and Holthuis

(1980) recognized Stimpson's species as well as M. pedunculata as synonyms of Atypopenaeus

stenodactylus. Recently, Hayashi (1992) stated that the long eyestalks, considered a character

typical of Miyadiella, become very short in specimens of A. stenodactylus having a carapace

length greater than 8.5 mm, in this respect being similar to those of the adult of the latter.

We have examined a specimen of Atypopenaeus compressipes with 7.2 mm carapace length,

total length of about 27 mm, which has short eyestalks that do not overreach the first antennular

segment, a length at which, according to Hayashi (1992) juveniles of the genus possess long

eyestalks. We also have examined specimens of Miyadiella omata from Indonesian waters with a

carapace length of 10 mm, a total length of 37.5 mm from the Museum National d'Histoire

Naturelle, Paris (material kindly made available by Dr. Alain Crosnier), with extremely long

eyestalks, thus, longer than specimens of Atypopenaeus exhibiting short ones; this specimen also

possesses clearly developed spinules on the first rostral teeth and along the postrostral carina,

another feature typical of Miyadiella; moreover, it has relatively short fifth pereopods, reaching to

about the end of the basal third of the scaphocerite, instead of extending by at least the length of

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

115

Fig. 67 A-C. — Miyadiella podophthalmus , 8 7 mm cl, Jiazi Lufeng, Guangdong, South China Sea.

A, Petasma, dorsal view. B, Petasma, ventral view. — C, 9 6.5 mm cl, East China Sea. Thelycum. Scales =

5 mm.

Fig. 67 D-E. — Miyadiella ornata , holotype, 9 38 mm tl, northeast of Java. D, Thelycum. Scale = 1 mm.

E, Posterior extremity of telson. Scale = 0.5 mm.

116

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

the dactyl beyond the antennular peduncle and, in one species (A. bicomis), as far as the tips of

the antennular flagella. On the basis of these observations we are tentatively recognizing

Miyadiella and Atypopenaeus as separate genera, the validity of which must await the laboratory

rearing of juvenile members of these genera.

Fig. 68 A. — Parapenaeopsis stylifera coromandelica , 9 28.5 mm cl, Singapore. Lateral view. Scale = 10 mm.

Fig. 68 B. — Parapenaeopsis balli , 9 12.5 mm cl, west of Sacapulco, Chiapas, Mexico. Lateral view

Scale = 5 mm. (From P£rez Farfante and Boothe, 1986).

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

117

Genus Parapenaeopsis Alcock, 1901

Figures 68-71

Penaeus - H. Milne Edwards, 1837, Hist. Nat. Crust., 2: 411 [part],

Penaeopsis - Bate, 1881, Ann. Mag. nat. Hist., (5)8: 183 [part].

Peneus (Parapeneopsis) Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 14; 1906, Cat. Indian Dec. Crust.,

3(1): 34.

Metapeneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16(95): 516 [part].

Parapenaeopsis [emendment of Parapeneopsis Alcock, 1901, under the plenary powers by the International

Commission on Zoological Nomenclature, 1969]. Placed on the Official List of Generic Names in Zoology,

International Commission on Zoological Nomenclature, 1969, Opinion 864, Name No. 1820, Bull. zool.

Nom., 25(4/5): 139. — De Man, 1911, Siboga Exped., 39a: 92. — Balss, 1914a, Abh. bayer. Akad. Wiss.,

suppl. 2(10): 14; 1925, Wiss. Ergebn. Dt.-Tiefsee Exped. Valdivia, 20: 229. — Kubo, 1949, J. Tokyo Coll.

Fish., 36(1): 368. — Barnard, 1950, Ann. S. Afr. Mus., 38: 604. — Dall, 1957, Austr. J. mar. Freshwat.

Res., 8: 213. — Racek and Dall, 1965, Verh. K. Ned. Akad. Wet. Nat., 56(3): 96. — Liu and Wang, 1987,

Oceanographia Limn. Sin., 18(6): 523. — Liu and Zhong, 1988, Penaeoid Shr. S. China Sea, : 195.

Trachypenaeus - Balss, 1916, Beitr. Kennt. Meeres. Westafr.,: 17.

Parapeneopsis - Burkenroad, 1934a, Bull. Bingham oceanogr. Coll.,4(7): 58. — Burkenroad, 1983, Crust.

Issues, 1: 283. — Dall eta/., 1990, Adv. Mar. Biol., 27: 83.

Type Species: By original designation, Penaeus styliferus H. Milne Edwards, 1837, Hist.

Nat. Crust., 2: 418.

Type Locality: Environs of Bombay, India.

Gender: Feminine.

Diagnosis. — Body slender. Integument thin, minutely setose-punctate, never densely so.

Rostrum armed with dorsal teeth only, variable in length and shape; epigastric tooth either close

to or distinctly separated from first rostral tooth. Carapace with small orbital spine, sometimes re¬

duced to not much more than sharp angle; well developed antennal (often continuous with carina)

and hepatic spines; pterygostomian spine lacking, but angle sometimes sharply produced; pos¬

tocular sulcus well defined; gastro-orbital carina absent; orbito-antennal and cervical sulci weak;

hepatic sulcus usually clearly marked anteriorly, sometimes feebly posteriorly; hepatic carina

often present, occasionally feeble, usually sharp and descending anteroventrally from hepatic

spine then extending almost to pterygostomian angle, usually blunt posteriorly; branchiocardiac

carina and sulcus clearly distinct to obsolete. Longitudinal suture falling short of posterior margin

of carapace, transverse suture usually present. Sixth abdominal somite with single elongate

cicatrix, or latter divided into two or three cicatrices. Telson armed with 3 to 5 (usually 4) small

movable lateral spines, with one or two pairs of small subapical fixed spines or unarmed.

Antennule lacking parapenaeid spine; antennular flagella subequal, rather short to considerably

long. Palp of first maxilla variable: (a) usually biarticulate, basial article produced into

proximolateral lobule, broad proximomesial and much smaller distomesial lobules, latter often

armed with spine; (b) sometimes entire, short, produced into same but less pronounced three

lobules, distomesial one armed with one to five spines; subdistolateral row of spinules usually

present on both. Basial spine present or absent on first and second pereopods; ischial spine

present or absent on first. Epipod lacking on third pereopod.

Petasma symmetrical, semiclosed, variable in shape; median lobes only barely perceptible,

moderately developed, or produced into oblique or fully lateral conspicuous projections;

moderately well sclerotized lateral lobes distally produced into relatively short, often roughly

spoutlike channeled projections, or elongate, hornlike ones (rarely semirectangular or strongly

bifid), variably directed, channeled, or tubular, and opening by minute apical apertures; petasma

118

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

sometimes expanded into

large, auriculate proximo-

lateral projections. Ap¬

pendix masculina simple,

or sometimes consisting

of two segments, proxi¬

mal one considerably

larger than horse-shoe or

cup-shaped distal one.

Thelycum closed, with

plate of sternite XIV

broad, sometimes deeply

emarginate anteriorly; me¬

dian protuberance strong,

usually concave, often

shieldlike, as broad as or

only slightly narrower

than plate of XIV, occa¬

sionally extending as far

as posterior thoracic

ridge. Seminal receptacles

exhibit considerable

variation indicated from

observations in limited

number of species:

minute elongate sacs [P.

hardwickii, Burkenroad

(1934a)], or strongly

developed bilobed (P.

atlantica) or trilobed

(P. stylifera, P. balli ) sacs (Perez Farfante and Boothe, 1986); if bilobed, consisting of large

pyriform posterior lobe and small anterolateral one; if trilobed, bearing additional anteromesial;

receptacles opening ventrally through slits which in single sac, elongate, extending mesially

along most of their length, in bilobed receptacles situated at the confluence of the posterior and

lateral lobes, and in trilobed ones on anterior margin of mesial lobe, slits lying at posterodorsal

margin of median protuberance, near base of fourth pereopods. As noted by Burkenroad

(1934a) in minute receptacles two openings, one for entrance (variably situated) and another for

exit mesial to hoods of sternite XIV.

Distribution of branchiae, epipods, and exopods in the genus Parapenaeopsis.

Somite

(Mxpl)

vrn

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

Xffl

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchi ae

Podobranchiae

Epipods

i(-)

i <-)

Exopods

Fig. 69 A. — Parapenaeopsis stylifera coromandelica, 3 18 mm cl, Karachi

market, Pakistan. Petasma, dorsal view. Scale = 1 mm.

Fig. 69 B. — Parapenaeopsis balli, 10.5 mm 3 cl, west of Sacapulco,

Chiapas, Mexico. Petasma, dorsal view. Scale = 1 mm. (From PErez

Farfante and Boothe, 1986).

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

119

Fig. 70 A-B. — Parapenaeopsis stylifera coromandelica : A, 2 28.5 mm cl, Singapore. Thelycum. — B, 2

24.5 mm cl, Ganjam coast, India. Seminal receptacles, dorsal view. Scales = 1 mm.

Fig. 70 C-D. — Parapenaeopsis atlantica, 2 30 mm cl, off St. Paul river mouth, Liberia. C, Thelycum and

outline of ventral view of seminal receptacles. D, Seminal receptacles, dorsal view. Scales = 1 mm.

Source: MNHN. Paris

120

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Fig. 71. — Parapenaeopsis balli, 2 12.5 mm cl, west of Sacapulco, Chiapas, Mexico: A, Thelycum. B, Seminal

receptacles, dorsal view. Scales = 1 mm. (From P£rez FaRFANTE and Boothe, 1986).

Remarks. — It is probable that species currently assigned to this very heterogeneous genus

will eventually be placed in several genera. The discussion above on the variations of the seminal

receptacles clearly indicates the striking differences of these structures among the species.

Species. — Parapenaeopsis acclivirostris Alcock, 1905. East coast of South Africa;

Mozambique; Madagascar (Europa Island); Red Sea; Gulf of Aden; Persian Gulf; Pakistan;

Western India; Sri Lanka.

Parapenaeopsis arafurica Racek and Dali, 1965. New Guinea; Northern Territory, Australia.

Parapenaeopsis aroaensis Hall, 1962. Malaysia.

Parapenaeopsis atlantica Balss, 1914c. West African coast from Senegal to Angola;

Mozambique.

Parapenaeopsis balli Burkenroad, 1934a. Pacific Mexico; El Salvador.

Parapenaeopsis cornuta (Kishinouye, 1900). West and south coast of India; Sri Lanka;

Singapore; Malaysia; Indonesia; South China Sea; Philippines; Hong Kong; Taiwan; Japan; New

Guinea; Australia.

Parapenaeopsis cultirostris Alcock, 1906. India; Malaysia; Indonesia; South China Sea.

Parapenaeopsis gracillima Nobili, 1903. Malaysia; Borneo.

Parapenaeopsis hardwickii (Miers, 1878). Pakistan; West and East coasts of India;

Bangladesh; Malaysia; Singapore; Borneo; Gulf of Tonkin; South China Sea; Taiwan; Japan.

Parapenaeopsis hungerfordi Alcock, 1905. Malaysia; Singapore; Indonesia; Gulf of Tonkin;

South China Sea; Hong Kong.

Parapenaeopsis incisa Liu and Wang, 1987. South China; Taiwan.

Parapenaeopsis maxillipedo Alcock, 1906. India; Sri Lanka; Malaysia; Borneo; Philippines;

New Guinea; northern Australia.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

121

Parapenaeopsis nana Alcock, 1905. India; Sri Lanka.

Parapenaeopsis sculptilis (Heller, 1862a). Pakistan; west coast of India; Malaysia; Indonesia;

South China Sea; Hong Kong; Philippines; Taiwan; New Guinea; Northern Australia.

Parapenaeopsis sinica Liu and Wang, 1987. South China Sea.

Parapenaeopsis stylifera coromandelica Alcock, 1906. East coast of India; Bangladesh;

Malaysia; Gulf of Thailand; Indonesia.

Parapenaeopsis stylifera stylifera (H. Milne Edwards, 1837). Persian Gulf; Arabian Sea;

Pakistan; Western India; Sri Lanka.

Parapenaeopsis tenella (Bate, 1888). India; Sri Lanka; Bangladesh; Malaysia; Indonesia; Gulf

of Tonkin; China; Philippines; Hong Kong; Taiwan; Japan; Korea; New Guinea; Northern

Australia.

Parapenaeopsis uncta Alcock, 1905. India; Sri Lanka; Bangladesh; Malaysia; Indonesia.

Parapenaeopsis venusta De Man, 1907. Malaysia; Indonesia; Queensland, Australia.

Genus Parapenaeus Smith, 1885

Figures 72-74

Peneus - Lucas, 1846, Hist. Nat. Anim. Artie., pt. 1: 46 [part].

Penaeus - Bate, 1881, Ann. Mag. nat. Hist., (5)8: 173 [part],

Parapenaeus Smith, 1885b, Proc. U.S. natn. Mus., 8(11/12): 170 [part]; 1886, Rep. U. S. Commnr Fish, for

1885: 685. — De Man, 1911, Siboga Exped., 39a: 77 [part], — Burkenroad, 1934b, Bull. Am. Mus. nat.

Hist., 68(2): 107. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 398. — Barnard, 1950, Ann. S. Afr. Mus.,

38: 600. — Dall, 1957, Aust. J. mar. Freshwat. Res., 8(2): 178. — Crosnier, 1986a, Mem. Mus. Hist. nat.

Paris, (A), Zool., 133: 303. — Dall el al ., 1990, Adv. Mar. Biol., 27: 87.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1961, Opinion 611, Name No. 1443, Bull. zool. Norn., 18(5): 306.

Peneus (Parapeneus) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 14 [part].

Parapeneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16(95): 519 [part]; 1906, Cat. Indian Dec. Crust., (3),

Macr., : 30 [part]. — Burkenroad, 1934b, Bull. Am. Mus. nat. Hist., 68(2): 107; 1983, Crust. Issues, 1:

282.

Neopenaeopsis Bouvier, 1905b, C. r. hebd. Seanc. Acad. Sci., Paris, 141: 747. Type species: By original

designation and monotypy, Neopenaeopsis paradoxus Bouvier, 1905b, C. r. hebd. Seanc. Acad. Sci., Paris,

141: 747 (= Parapenaeus longirostris). Type locality: Caribbean Sea. Gender: Feminine.

Type Species: By original designation: Penaeus longirostris Lucas, 1846, Explor. Sci.

l'Algerie, Zool. 2. Hist. Nat. Anim. Artie., pt. 1: 46.

Type Locality: Off Cape Matifou, Algeria.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum armed with dorsal teeth (5-9) only, very

variable in length; epigastric tooth situated conspicuously posterior to first rostral tooth. Carapace

with orbital spine more or less pointed; with well developed antennal and hepatic spines;

pterygostomian or branchiostegal spine usually present (absent in P. longipes Alcock, 1905,

except in some small specimens), sometimes continuous with well marked carina; postocular and

cervical sulci lacking; hepatic sulcus indistinct or weak; hepatic carina usually well defined only

anteriorly; branchiocardiac carina absent; longitudinal suture long, extending almost to posterior

Source:

122

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Fig. 72. — Parapenaeus longirostris, 2 34 mm cl, off Sicily, Mediterranean. Lateral view. Scale = 10 mm.

margin of carapace, transverse suture at level of second pereopod. Sixth abdominal somite

without cicatrix. Telson armed with pair of long fixed lateral spines at about posterior third.

Antennule with small parapenaeid spine; antennular flagella sexually dimorphic: much longer

in males than in females and ventral flagellum always longer than dorsal in females, whereas in

males dorsal sometimes longer. Palp of first maxilla entire, broad, blunt and bearing small setose

proximolateral and conspicuous mesial lobules. Basial and ischial spines on first pereopod only.

Somite XIII with pleurobranchia and posterior arthrobranchia only.

Distribution of branchiae, epipods, and exopods in the genus Parapenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

i (-)

Exopods

Petasma symmetrical, semiclosed, with numerous processes and folds; dorsomedian lobule

bearing one subdistally; dorsolateral lobule with one distally and ventrolateral lobule with two

terminal ones; distal parts of three lobules firmly connected by convoluted membrane. Appendix

masculina subovoid. Thelycum with single plate of sternite XIV raised in two pairs of lateral

prominences disposed longitudinally (anterior pair columnar in shape and more prominent), both

pairs separated by median groove, depression, or tubercles; median protuberance of sternite XIII

strongly developed, more or less excavate posteromedially. Seminal receptacles consisting of pair

of longitudinally or obliquely disposed sacs, opening by slits in groove between sternites XIV

and XIII.

Heldt (1938a) and PFrez Farfante (1982) illustrated the seminal receptacles of

P. longirostris and P. politus respectively. Heldt also described the spermatophore of the former

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

123

as consisting of a large slightly dorsoventrally depressed sperm mass containing strongly bound

spermatozoids and enveloped by a thin membrane, and a non-cellular cushion-like flange

narrowly attached to the sperm sac which is also accompanied by a gelatinous material.

Fig. 73. — Parapenaeus longirostris : A, 6 33 mm cl, off Sicily, Mediterranean. Petasma. ventral view.

— B, 2 34 mm cl, off Sicily, Mediterranean. Thelycum. Scales = 1 mm.

Fig. 74. — Parapenaeus politus,

20 mm cl, Sabanalarga, Colom¬

bia. Seminal receptacles, dorsal

view. Scale = 2 mm. (From

P£rez Farfante, 1982).

Species. — Parapenaeus americanus Rathbun, 1901. Off

New England; Gulf of Mexico; Bahamas; Cuba; Puerto Rico;

St. Lucia; Belize; Brazil; Northern Uruguay.

Parapenaeus australiensis Dali, 1957. Indonesia; Philip¬

pines; Queensland, New South Wales, Australia; Fiji.

Parapenaeus australiensis forma nodosa Crosnier, 1986a.

Philippines.

Parapenaeus fissuroides erythraeus Crosnier, 1986a. Red

Sea.

Parapenaeus fissuroides fissuroides Crosnier, 1986a.

Indonesia; South China Sea; Philippines; Hong Kong;

Taiwan; Japan; Korea.

Parapenaeus fissuroides indicus Crosnier, 1986a. East

coast of South Africa; Mozambique; Kenya; Zanzibar;

Madagascar; Gulf of Oman.

Parapenaeus fissurus (Bate, 1881). East coast of South

Africa to Zanzibar; Red Sea; Arabian Sea; India; Andaman

Islands; Malaysia; Indonesia; Gulf of Tonkin; South China

Source:

124

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Sea; Philippines; Taiwan; Japan; New Guinea; Wallis and Futuna Islands.

Parapenaeus investigatoris Alcock and Anderson, 1899. Natal, South Africa; Gulf of Aden;

India; Andaman Islands; Malaysia; Indonesia; Philippines; Taiwan; Japan.

Parapenaeus lanceolatus Kubo, 1949. Indonesia; Vietnam; South China Sea; Philippines;

Hong Kong; Taiwan; Japan; Korea; Western Australia.

Parapenaeus longipes Alcock, 1905. East coast of Africa from Tanzania to Somalia;

Madagascar; Pakistan; West and South India; Sri Lanka; Indonesia; South China Sea;

Philippines; Hong Kong; Taiwan; Japan; New Guinea.

Parapenaeus longipes forma denticulata Crosnier, 1986a. Indonesia; Philippines.

Parapenaeus longirostris (Lucas, 1846). Mediterranean; eastern Atlantic from Portugal to

Namibia.

Parapenaeus murrayi Ramadan, 1938. East coast of South Africa; Mozambique; Zanzibar;

Madagascar; Reunion; Indonesia; Philippines; Japan.

Parapenaeus perezfarfantae Crosnier, 1986a. Philippines.

Parapenaeus politus (Smith, 1881). Off New Jersey (south of Martha's Vineyard), south and

along the Gulf of Mexico; Caribbean Sea to French Guiana.

Parapenaeus ruberoculatus Hall, 1962. Malaysia; Indonesia; Philippines.

Parapenaeus sextuberculatus Kubo, 1949. Madagascar; India; Indonesia; Gulf of Tonkin;

South China Sea; Philippines; Taiwan; Japan; New South Wales, Australia.

Genus Pelagopenaeus Perez Farfante & Kensley, new designation 3

Figures 75-76

Peneus - Faxon, 1893, Bull. Mus. comp. Zool. Harv., 24(7): 211; 1895, Mem. Mus. comp. Zool. Harv., 18(15):

181. — Alcock, 1906, Cat. Indian Dec. Crust., 3(1): 49 [Placed in "uncertain or doubtful species" category],

Penaeus - Lenz and Strunck, 1914, Dt. Sudpol.-Exped., 15 (zool. 7): 298.

Funchalia (Pelagopenaeus) Burkenroad, 1934b, Bull. Am. Mus. nat. Hist., 68(2): 73, 76; 1936b, Bull. Bingham

oceanogr. Coll., 5(2): 136. — Ramadan, 1938, Sci. Rep. John Murray Expcd., 5(3): 64.

Funchalia - Gordon and Ingle, 1956, J. mar. biol. Ass. U. K., 35: 476 [part], — Dall, 1957, Aust. J. mar.

Freshwat. Res., 8(2): 163 [part]. — Crosnier and Forest, 1973, Faune Trop., 29: 296 [part],

Pelagopenaeus - Grippa, 1976, Atti Soc. ital. Sci. nat., 117(3/4): 126; 1987, Investnes Pesq., 51 (Suppl. 1): 73.

— Dall etal., 1990, Adv. Mar. Biol., 27: 91.

Type Species: By present selection, Penaeus balboae Faxon, 1893, Bull. Mus. comp Zool

Harv., 24(7): 211.

Type Locality; Off Isla del Coco, Costa Rica, 5°26'20"N, 86°55’00"W, 1280 m.

Gender: Masculine.

As Burkenroad proposed the name Pelagopenaeus for two species in 1934, i.e. after 1930, and did not select

a type species, Pelagopenaeus is an unavailable name under Article 13b of the International Code of Zoological

Nomenclature. Dr. L. Holthuis (in lilt, 1996) notes further that the indication by W. L. Sclater [1935,

Zoological Record for 1934, vol. 71, part 9 (Crustacea), page 34]: "Pelagopenaeus subg. n. for Penaeus balboae

Faxon" is not a type selection either. Thus it is with the greatest reluctance, and in light of Dr. Burkenroad being

deceased, that we make the name available and assume in this work the authorship of Pelagopenaeus.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

125

Fig. 75 . _ Pelagopenaeus balboae , 3 28 mm cl, off Gulf of Panama. Lateral view. Scale = 10 mm.

Diagnosis. — Integument minutely pubescent. Rostrum short, reaching to about midlength of

second antennular segment; armed with dorsal and ventral teeth. Carapace lacking orbital spine

(angle only occasionally slightly marked) and antennal spine, with pterygostomian and small

hepatic spines, latter often indistinct in adults; postocular sulcus lacking; gastro-orbital carina,

orbito-antennal, and cervical sulci absent; postrostral carina long and well marked; antennal carina

clearly defined; elongate dorsolateral carina present; long hepatic carina joining long

branchiocardiac; branchiostegal carina also long; without longitudinal and transverse sutures.

Fourth through sixth abdominal somites each bearing single long, strong cicatrix. Telson with

three pairs of fixed lateral spines preceded by numerous pairs of much smaller ones; lateral ramus

of uropod armed with large lateral spine.

Antennule lacking parapenaeid spine; antennular flagella subfiliform, dorsal one longer than

carapace and much longer than ventral. Mandibles with incisor processes short, straight; distal

article of biarticulate palp small, symmetrical, triangular. Palp of first maxilla relatively short,

slender, biarticulate, basal article armed with small proximolateral comb of short setae,

subdistolateral row of spinules, and long subdistal spine on mesial margin; distal article bearing

long apical setae. Dactyl of third maxilliped elongate and flat, slightly longer in females than in

males. Ridge of lateral ramus of uropod produced into conspicuous spine. Basial and ischial

spines on first and second pereopods.

Distribution of branchiae, epipods, and exopods in the genus Pelagopenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XJY

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

126

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Petasma semi-open, symmetrical, with lateral lobe not produced in free, distally directed

projection. Appendix masculina suboval. Thelycum open, with deep anterior depression on

sternite XIV flanked by pair of moderately large, semicircular flaps; sternite XIII with deeply

excavate median protuberance delimiting depression on sternite XIV.

Spermatophores elongate-ovate in outline, and when attached to female are tightly held in

place and partly covered by the thelycal flaps.

Fig. 76. — Pelagopenaeus balboae: A, 6 27.5 mm cl, Line Islands, east of Christmas Island. Petasma, dorsal

view. — B, 2 31.5 mm cl. Line Islands, east of Christmas Island. Thelycum. Scales = 1 mm.

Species. — Pelagopenaeus balboae (Faxon, 1893). Eastern Pacific Ocean off Gulf of

Panama; Isla de Cocos; Caribbean; Bermuda; Equatorial mid-Atlantic; off Cape Verde Islands;

eastern Atlantic Ocean off Congo; South-eastern Atlantic; Arabian Sea; north-west Pacific Ocean.

Genus Penaeopsis Bate, 1881

Figures 77-79

Penaeus - Bate, 1881, Ann. Mag. nat. Hist., (5)8: 173 [part],

Penaeopsis Bate, 1881, Ann. Mag. nat. Hist., (5)8: 182; 1888, Rep. scient. Results Voy. Challenger, 24: 273. —

Bouvier, 1905a, C. r. hebd. Seanc. Acad. Sci., Paris, 140: 981; 1908b, Bull. Inst, oceanogr., Monaco, 119: 5.

Schmitt, 1926, Zool. (biol.) Results Fish. Exp. 'Endeavour', 5(6): 319 [part], — Kubo, 1949, J. Tokyo

Coll. Fish., 36(1): 320. — PErez Farfante, 1980a, Fish. Bull., U. S„ 77(4): 722. — Liu and Zhong 1988

Penaeoid Shr. S. China Sea, : 83. — Dall et al„ 1990, Adv. Mar. Biol., 27: 91.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1821, Bull. zool. Norn., 25(4/5): 139.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

127

Parapenaeus - Smith, 1885b, Proc. U.S. natn. Mus., 8(11/12): 172 [part].

Metaperiaeus - Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 271 [part].

Peneus (Parapeneus) - Alcock, 1901, Descr. Cat. Indian Deep-sea Crust.,: 14 [part].

Artemesia. - Bouvier, 1905a, C. r. hebd. Seanc. Acad. Sci., Paris, 140: 982 [part].

Parapeneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 519 [part]; 1906, Cat. Indian Dec. Crust., 3(1): 30

[part].

Penaeopsis (Penaeopsis) - Burkenroad, 1934a, Bull. Bingham oceanogr. Coll., 4(7): 4, 12.

Peneopsis - Burkenroad, 1983, Crust. Issues, 1: 282.

Fig. 77. — Penaeopsis serrata, $ 27 mm cl, west of Puerto Rico. Lateral view. Scale = 10 mm. (From

P£rez Farfante, 1979).

Type Species: By designation by Bouvier, 1905a, Penaeopsis serratus Bate, 1881, Ann. Mag.

nat. Hist., (5)8: 183, C. r. hebd. Seanc. Acad. Sci., Paris, 140: 981.

Type Locality: Off Barbados, "Gulf of Mexico", 12°58'33"N, 59°36'45"W, 399 m.

[Discussion of types in P£rez Farfante, 1980a: 760.]

Gender: Feminine.

Diagnosis. — Integument glabrous in appearance, various areas microscopically

setose. Rostrum armed with dorsal teeth only, extending along its entire length; epigastric tooth

Source

128

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

conspicuously separated from first rostral tooth.

Carapace with well developed antennal,

pterygostomian, and hepatic spines, lacking orbital

and branchiostegal spines; postocular sulcus lacking;

cervical sulcus and carina well marked; hepatic

sulcus with anterior part accompanied by sharp

carina and descending from hepatic spine then

turning toward and reaching pterygostomian spine,

carina absent from posterior part; branchiocardiac

carina variably developed, from long and strong to

short; longitudinal and transverse sutures absent.

Sixth abdominal somite bearing long, interrupted

cicatrix. Telson armed with pair of moderately long,

fixed, posterolateral spines preceded by two or three

pairs of small, movable spines.

Antennule with long parapenaeid spine;

antennular flagella long, 0.75 to almost twice length

of carapace; ventral flagellum sexually dimorphic: in

male shorter than dorsal and with proximal part

forming rigid, flattened, strongly arched loop; in

female straight and longer than dorsal. Palp of first

maxilla entire, short, blunt or truncate distally,

bearing weak (sometimes indistinct) proximolateral

lobe, conspicuous proximomesial lobe, and slender

Fig. 78. — Penaeopsis serraia, 3 22 mm cl, distomesial spine. Basial spine on first pereopod,

Gulf of Mosquito, Panama. Petasma, dorsal lacking on third maxilliped, second and third

view. Scale = 1 mm. (From P£rez pereopods. Anterior arthrobranchia lacking on

Farfante, 1979). somite XIII.

Petasma symmetrical, semiclosed, with lateral

lobes not produced distally into spoutlike projections. Appendix masculina jointed to dorsomesial

rib of endopod, heavily sclerotized, wider than long. Thelycum closed, with single plate of

sternite XIV occupying entire sternite and bearing variably developed median prominence;

median protuberance of sternite XIII well developed. Seminal receptacles consisting of paired

bilobed sacs (one large submedian, the other smaller extending laterally), each opening through

single long slit located at anterior extremity of sternite XIV, at base of fourth pereopod.

The compound spermatophore of Penaeopsis is not as complex as those of members of genera

with open thelyca (e.g. Litopenaeus, Pleoticus). Perez Farfante (1980a) presented an

illustration of that of P. rectacuta.

Distribution of branchiae, epipods, and exopods in the genus Penaeopsis.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

129

Fig. 79 A-B. — Penaeopsis serrata , $ 29.5 mm cl, Dry Tortugas Islands, Florida: A, Thelycum. Scale = 2 mm.

B, Seminal receptacles, dorsal view. Scale = 1 mm. (From P£rez Farfante, 1979).

Fig. 79 C-D. — Penaeopsis eduardoi , 9 32.5 mm cl, southwest of Taiwan: A, Thelycum. Scale = 1 mm.

B, Seminal receptacles, dorsal view. Scale = 1 mm.

Source: MNHN. Paris

130

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Species. — Penaeopsis balssi Ivanov and Hassan, 1976. East coast of South Africa;

Mozambique; Kenya; Tanzania; Somalia; Madagascar; Saya de Malha Bank.

Penaeopsis challengeri De Man, 1911. Fiji; Wallis and Futuna Islands.

Penaeopsis eduardoi Perez Farfante, 1977b. Mozambique; Madagascar; Reunion; Bay of

Bengal; Indonesia; South China Sea; Philippines; Taiwan; Japan; New South Wales, Australia;

Fiji.

Penaeopsis jerryi Perez Farfante, 1979. Off Mozambique; Yemen; Gulf of Aden; Southwest

and East India; off Andaman Islands.

Penaeopsis rectacuta (Bate, 1881). East coast of South Africa; Andaman Islands; Indonesia;

South China Sea; Taiwan; Philippines; Japan; Fiji.

Penaeopsis serrata Bate, 1881. Western North Atlantic Ocean from New Jersey to Gulf of

Mexico, Caribbean Sea to French Guiana; southern Brazil; eastern North Atlantic Ocean from

southern Portugal to Mauritania.

Genus Penaeus Fabricius, 1798

Figures 80-81

Peneus Weber, 1795, Nomencl. Entomol. Sec. Ent. Syst. Illust., : 94 [nomen nudum, International Commission

on Zoological Nomenclature, Opinion 104, Smithson, misc. Colins, 1928, 73(5): 25].

Penaeus Fabricius, 1798, Suppl. Entomol. Syst., : 408 [part], — Latreille, 1803, Hist. nat. Crust. Ins., 6: 246

[part]; 1806, Gen. Crust. Ins., 1: 53; 1810, Consid. Gener. Crust. Arach. Ins., : 102, 422. — Smith, 1885b,

Proc. U. S. natn. Mus., 8(11/12): 170 [part]; 1886, Rep. U. S. Commnr Fish., for 1885: 684 [part], —

De Man, 1911, Siboga Exped., 39a: 95 [part], — Burkenroad, 1934b, Bull. Am. Mus. nat. Hist., 68(2): 73

[part], — Kubo, 1949, J. Tokyo Coll. Fish., 6(1): 268 [part]. — Dall et al ., 1990, Adv. Mar. Biol., 27: 93

[part].

Placed on the Official List of Generic Names in Zoology, 1928, Smithson, misc. Colins, 73(5): 25.

Penoeus - Lamarck, 1818, Hist. Nat. Anim. s. Vert., 5: 205. — Lucas, 1840, Hist. nat. Crust. Arach. Myr., :

195.

Peneus - Alcock, 1901, Descr. Cat. Ind. Deep-Sea Crust., : 13 [part]; 1905, Ann. Mag. nat. Hist., (7)16: 513

[part]; 1906, Cat. Indian Dec. Crust., 3(1): 7 [part], — Burkenroad, 1983, Crust. Issues, 1: 282.

Penaeus (Penaeus) - PErez Farfante, 1969, Fish. Bull., U. S., 67(3): 461. — Burukovsky, 1972, Trudy

AtlantNIRO, 42: 3, 5, 8, 13, 16. — Motoh and Buri, 1984, Res. Crust., (13, 14): 24. — Liu and Zhong,

1988, Penaeoid Shr. S. China Sea, : 104. — Hayashi, 1992, Dendr. Crust. Japan. Waters, : 122.

Type Species: By subsequent designation of Latreille, 1810, Penaeus monodon Fabricius,

1798, Suppl. Entomol. Syst., : 408; neotype for Penaeus monodon, Rijksmus. Nat. Hist.,

Leiden, designated by Holthuis, 1949, Proc. K. ned. Akad. Wet., 52(9): 10.

Type Locality: Bay of Batavia (= Jakarta), Indonesia.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum with dorsal and 2-4 (occasionally 5) ventral

teeth, moderately long, in young slightly exceeding antennular peduncle, in adult reaching to

about midlength of second antennular segment. Carapace with antennal and hepatic spines

pronounced, lacking orbital and pterygostomian spines; postocular sulcus absent; cervical sulcus

shallow; postrostral carina long, reaching almost to posterior margin of carapace; adrostral carina

and sulcus short, extending only to about level of epigastric tooth; gastrofrontal carina absent;

gastro-orbital carina short, orbito-antennal sulcus well marked, extending anteriorly not more

than two-thirds of distance from hepatic spine to orbital margin; cervical and hepatic carinae well

defined, cervical sulcus shallow; branchiocardiac carina lacking; longitudinal and transverse

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

131

sutures absent. Sixth abdominal somite bearing three cicatrices, lacking dorsolateral sulcus.

Telson unarmed.

Fig. 80. — Penaeus monodon, 2 50 mm cl, Micaune, Mozambique. Lateral view. Scale = 10 mm.

Antennule lacking parapenaeid spine; antennular flagella shorter than carapace. Palp of first

maxilla elongate, consisting of 2-5 articles, distal ones slender; large basal article produced into

setose proximal lobule on lateral and mesial margins, bearing long subdistomesial spine and

distolateral comb of small spinules. Basial spine on first and second pereopods. Exopod on fifth

pereopod absent in P. monodon , unique in series.

Petasma symmetrical, semiclosed with distomedian projections quite short, ventral costae

narrowly gaping or contiguous, long and strongly curved distally. Appendix masculina

subelliptical, or suboval, armed with numerous marginal spines. Thelycum closed, with pair of

lateral plates of stemite XIV meeting along midline, shielding simple median saclike seminal

receptacle; stemite XIII with well developed median protuberance.

Spermatophores lacking attachment structures. Observations by Malek and Bawab (1974a,

b), Champion (1987), and Bauer and Cash (1991), on the formation and structure of the

spermatophore in other closed thelycum species previously included in Penaeus s.l.

( Faifantepenaeus, Fenneropenaeus, Melicertus) indicate that they may be similar in all of them.

Distribution of branchiae, epipods, and exopods in the genus Penaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

i (-)

132

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Fig. 81. — Penaeus monodon : A, 6 54 mm cl, Indian Ocean. Petasma, dorsal view. Scale = 1 mm. —

B, $ 50 mm cl, Micaune, Mozambique. Thelycum. Scale = 5 mm. — C-D, $ 35 mm cl, Parati Taoch,

Palau Islands. C, Seminal receptacle, ventral view (lateral plates of thelycum removed). D, Seminal receptacle,

dorsal view. Scales = 5 mm.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

133

Species. — Penaeus esculentus Haswell, 1879. Western Australia, Northern Territory,

Queensland to New South Wales, Australia.

Penaeus monodon Fabricius, 1798. East coast of Africa; Red Sea; Madagascar; Mauritius;

Reunion; Pakistan; India; Sri Lanka; Malaysia; Singapore; Indonesia; China; Philippines; Hong

Kong; Taiwan; Japan; Korea; New Guinea; Western Australia to Queensland, Australia; Fiji.

Penaeus semisulcatus De Haan, 1844. Eastern Mediterranean; east coast of Africa; Red Sea;

Madagascar; Persian Gulf; Pakistan; India; Sri Lanka; Malaysia; Indonesia; South China Sea;

Philippines; Hong Kong; Taiwan; Japan; Korea; New Guinea; Western Australia, Northern

Territory, Queensland, New South Wales, Australia; Fiji.

Genus Protrachypene Burkenroad, 1934

Figures 82-84

Protrachypene Burkenroad, 1934a, Bull. Bingham oceanogr. Coll., 4(7): 43; 1983, Crust. Issues, 1: 280. — Dall

et al., 1990, Adv. Mar. Biol., 27: 99.

Fig. 82. — Protrachypene precipua, $ 24.5 mm cl, Playas, Ecuador: A, Lateral view. Scale = 1 mm. B, Chela of

pereopod 1 enlarged.

Type Species: By monotypy, Protrachypene precipua Burkenroad, 1934a, Bull. Bingham

oceanogr. Coll., 4(7): 43.

Type Locality: Bella Vista Beach, Panama City, Panama.

Gender: Feminine.

Diagnosis. — Integument pubescent. Rostrum bearing dorsal teeth only, limited to base, very

long, considerably overreaching antennular peduncle; epigastric tooth distinctly separated from

first rostral tooth. Carapace with orbital angle well marked, well developed antennal and hepatic

134

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

spines; pterygostomian angle produced but without spine; postocular sulcus well marked; gastro-

frontal carina lacking; orbital-antennal sulcus deep; cervical sulcus short, shallow, without

accompanying carina; hepatic carina short, merging with extremely strong branchiocardiac carina;

longitudinal and transverse sutures present. Sixth abdominal somite bearing strong, long cicatrix.

Telson armed with eight or more minute, movable, lateral spines.

Antennule lacking parapenaeid spine; antennular flagella unequal

in length, dorsal flagellum considerably longer than both ventral

one and carapace. Palp of first maxilla biarticulate, basal article

tapering to minute distal one, and produced into weak

proximolateral, large proximomesial, and quite small mediodistal

lobules, latter bearing strong spine; palp also bearing row of spines

on midlateral margin. First three pereopods with weak extremely

elongate chela, lengthened propod and very short dactyl, less than

half propod (in juveniles chelae of normal form); fourth and fifth

pereopods very long and extremely slender, subflagelliform. Long

basial spine on first pereopod and spine or minute protuberance on

basis of second. Exopod absent on second maxilliped.

Petasma symmetrical, semiclosed, depressed, with lateral lobes

produced distolaterally in broad horns; ventrolateral lobule

terminating in subcircular flap overlapping or surpassing

dorsolateral lobule. Appendix masculina subrectangular, almost as

wide as long and strongly concave ventrally. In female, endopod of

first pleopod absent. Thelycum closed, with single plate of sternite

XIV conspicuously cleft anteriorly, produced into rounded

anterolateral lobules, their anteromesial margins bordering grooves

(sternal invaginations), leading to relatively shallow median pocket.

Median protuberance of sternite XIII roughly triangular, with

anterior margin broadly curved or trilobed, narrowing posteriorly

toward median pocket; latter supported laterally by paired heavily

sclerotized, convoluted lamellae. Seminal receptacles large,

bladderlike, opening on anterolateral extremities of grooves.

Distribution of branchiae, epipods, and exopods in the genus Protrachypene.

Somite

(Mxpl)

Vffl

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 83. — Protrachypene

precipua , S 24.5 mm cl,

Playas, Ecuador. Petasma,

ventral view. Scale =

1 mm.

Spermatophores apparently entering median pocket with sperm mass passing to seminal

receptacles, and trailing sperm-free material plugging pocket. As noted by Burkenroad (1934a)

the plug blocks the entrance and the sperm is forced to leave the receptacle through the laterally

located slits at the base of the fourth pereopods, close to the gonopores. In general, this seems to

be the method of impregnation and release of sperm in members of the genera of the

Trachypenaeus 'series'.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

135

Fig. 84. — Protrachypene precipua, 2 24.5 mm cl, Playas, Ecuador: A, Thelycum. B, Seminal receptacles, dorsal

view. Scales - 1 mm.

Species. — Protrachypene precipua Burkenroad, 1934a. El Salvador to Northern Peru.

Genus Rimapenaeus , new genus

Figures 85-87

Penaeus - Stimpson, 1874, Ann. Lyc. nat. Hist. N.Y., 10: 135 [description of Petiaeus constrictus only].

Parapenaeus - Smith, 1885b, Proc. U. S. natn. Mus., 8(11/12): 170 [part].

Peneus (Trachypeneus) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 15 [part].

Trachypeneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)26: 522 [part]. — Burkenroad, 1983, Crust. Issues, 1:

283 [part].

Trachypeneus (Trachysalambria) - Burkenroad, 1934a [September], Bull. Bingham oceanogr. Coll., 4(7): 49

[Division I, Section 1].

Trachypeneus - Burkenroad, 1934b [December], Bull. Am. Mus. nat. Hist., 68(2): 94 [Division 1, Section 1].

Trachypenaeus - Dall, 1957, Aust. J. mar. Freshwat. Res., 8(2): 202 [part]. — P£rez Farfante, 1971b, Fish.

Bull. U.S., 69(3): 365 [part]. — Dall et al 1990, Adv. Mar. Biol., 27: 102 [part].

Type Species: Trachypenaeus faoe Obarrio, 1954, in Segundo Centro Latinoamericano de

Capacitacion Pesquera (= CLACP, FAO), Tr. 41: 3[a]. Lectotype, here designated, USNM

256545.

136

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

Type Locality: Off Panama, 12 m.

Gender: Masculine.

Etymology. — From the Greek rima, cleft, in combination with the generic name Penaeus,

alluding to the conspicuously or more often, deeply cleft thelycal plate of stemite XIV.

Fig. 85. — Rimapenaeus faoe, 9 37 mm cl, off Juan Diaz, Gulf of Panama. Lateral view. Scale = 10 mm.

Diagnosis. — Body thickset. Integument thick, pubescent, usually densely so. Rostrum

reaching between distal margin of first antennular segment and about proximal fourth of dorsal

flagellum; armed with dorsal teeth only; epigastric tooth distinctly separated from first rostral

tooth. Carapace with orbital, antennal, and hepatic spines present; pterygostomian angle blunt,

lacking spine; postocular sulcus clearly distinct; short cervical and well marked hepatic sulcus;

orbito-antennal sulcus weak; branchiocardiac carina well defined, accompanying sulcus almost

indistinct; longitudinal suture long, ending well posterior to epigastric tooth; transverse suture

well marked. Abdomen with sixth somite bearing one continuous cicatrix or latter more often di¬

vided into two well defined parts. Telson with 3-4 pairs of movable lateral spines, or 3 movable

and subapical fixed, subapical pair projecting from slight shoulder, or unarmed (in Rimapenaeus

byrdi only).

Antennule lacking parapenaeid spine, antennular flagella shorter than carapace. Palp of first

maxilla entire, rapidly tapering to produced apex, bearing setose proximolateral, broad, setose

proximomesial, and small acute distomesial lobules, latter bearing 5 elongate spines; palp also

armed distally with ventrolateral rows of spinules. Fourth and fifth pereopods with dactyl neither

elongate nor subdivided. Basial spine on third maxilliped, first and second pereopods; ischial

spine lacking on first pereopod. Epipod on first through third pereopods.

Petasma symmetrical, semi-closed, with lateral lobes produced distolaterally into large,

occasionally small (in R. byrdi only) hornlike projections either extending straight laterally or

sinuously curving forward. Appendix masculina subrectangular with rounded comers. Thelycum

closed, with plate of stemite XIV entirely cleft medially, resulting in two truncate flaps or, more

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

137

often, deeply cleft anteriorly, and produced into

moderately to strongly elongate, tongue-like flaps;

sternal invagination forming pronounced median

pocket dorsally continuous with median pro¬

tuberance of stemite XIII; latter strongly developed,

but not produced caudally. Seminal receptacles

consisting of paired trilobed sacs, each with large

posterior lobe and two small lobes, one lateral,

another mesial, flanking median protuberance;

sinuous slitlike openings of receptacles lying on

anterior extremity of posterior lobe connecting

mesially with median pocket and extending to

anterolateral lobe (dorsal to posterolateral part of

median protuberance), where opening to exterior

near base of fourth pereopod. A general description

of the seminal receptacles of Trachypenaeus

[Rimapenaeus] similis was presented by

Burkenroad (1934b) and recently, Bauer and Lin

(1993) conducted a thorough investigation of these

structures, resulting in an excellent and finely

illustrated contribution. Years before, Perez

Farfante (1971b) described and depicted the

seminal receptacles of T. [R.] fuscina.

Material leaving the ejaculatory duct, i.e. the

spermatophore, consists of numerous minute

packets of sperm ' (spermatophores sensu

Burkenroad, 1934b, Bauer and Lin, 1993) in a

seminal fluid and a viscous adhesive material, the plug substance. Apparently, during copulation

the spermatophores are introduced into the seminal receptacles and the trailing plug substance fills

and seals the median pocket where it hardens considerably. This arrangement prevents sperm

from flowing back to the median pocket, forcing it to exit close to the gonopores. Earlier,

Burkenroad (1934a) had inferred the differing places of entrance and exit of the sperm in

members of Trachypenaeus s.l. and later (1934b) described the transit of the sperm mass in

Trachypenaeus [Rimapenaeus] similis, and the role of the trailing material. Bauer and Lin (1993:

184) also discuss the role of the plug substance as a mechanism to preclude subsequent

inseminations by other males.

Distribution of branchiae, epipods, and exopods in the genus Rimapenaeus.

Somite

(Mxpl)

Vffl

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

1 +v

Podobranchiae

Epipods

Exopods

Fig. 86 . — Rimapenaeus faoe, 3 20 mm cl,

Bellevista Beach, Panama. Petasma, dorsal

view. Scale = 1 mm.

Remarks. — Rimapenaeus , one of the genera that has been segregated from Trachypenaeus

s.l., includes some of the species (all from American waters) grouped by Burkenroad (1934a) in

138

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

the subgenus Trachysalambria (Section 1, Division I). This genus can be distinguished by the

thelycum which exhibits the plate of sternite XIV either divided in two by a longitudinal slit

separating truncate plates or, more often, produced in relatively short or elongate tonguelike

flaps, with sternite XIV typically invaginated anteriorly forming a well defined median pocket,

instead of a broad bracket-shaped transverse groove. Rimapenaeus can also be distinguished by

the petasma, the lateral lobes of which are distally produced into usually large, occasionally

small, projections, extending laterally, often in a sinuous course with apical parts curving

forward. The longitudinal suture is long, conspicuously overreaching the hepatic spine, rather

than short, not extending behind the hepatic spine, and a basial spine is present on the third

maxilliped. The seminal receptacles are trilobed, whereas in the other three genera they are

bilobed. Finally, in Rimapenaeus the openings of the receptacles are mesially connected to a

median pocket, and from there extend anterolaterally. Although in species of the other genera

( Megokris , Trachypenaeus, Trachysalambria ) formerly included in Trachypenaeus s.l. the

epipods may be present on, or absent from, the first and second pereopods (in all four genera an

epipod is always present on the third), all observations to the present indicate that in

Rimapenaeus they are always present.

Fig. 87. —Rimapenaeus faoe, $ 34 mm cl, Tortugas grounds, south of Buenaventura, Colombia: A, Thelycum.

B, Seminal receptacles, dorsal view. Scales = 1 mm.

Species. — Rimapenaeus byrdi (Burkenroad, 1934a). Southern Gulf of California off Isla

Altamura, Sinaloa, Mexico; Guatemala; Panama; Golfo de Guayaquil, Ecuador; Colombia; to

Paita, northern Peru.

Rimapenaeus constrictus (Stimpson, 1874). Nova Scotia; Chesapeake Bay to Florida Keys,

Gulf of Mexico; Caribbean Sea to Bahia de Zimbros, Brazil; Bermuda.

Rimapenaeus faoe (Obarrio, 1954). From southern Gulf of California to Golfo de

Guayaquil, Ecuador.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

139

Rimapenaeus fuscina (Perez Farfante, 1971b). From Golfo de Tehuantepec, Mexico, to

Paita, northern Peru.

Rimapenaeus pacificus (Burkenroad, 1934a). South of Punta San Felipe, northwest of Baja

California, and Gulf of California, Mexico, to Huacho, Peru.

Rimapenaeus similis (Smith, 1885b). Florida Keys, Gulf of Mexico; Caribbean Sea to Sao

Paulo, Brazil.

Genus Tanypenaeus Perez Farfante, 1972

Figures 88-90

Tanypenaeus Perez Farfante, 1972, Bull. mar. Sci. Gulf Caribb., 22: 186. — Dall et al., 1990, Adv. Mar. Biol.,

27: 99.

Type Species: By original designation, Tanypenaeus caribaeus Perez Farfante, 1972, Bull,

mar. Sci. Gulf Caribb., 22(1): 187.

Type Locality: Off Punta Faro, Colombia, 11°08'N, 74°47'W, 155-157 m.

Gender: Masculine.

Fig. 88 . — Tanypenaeus caribaeus, paratype, 2 18.5 mm cl, off Punta Faro, Colombia. Lateral view.

Scale = 10 mm.

Diagnosis. — Integument pubescent. Rostrum pronouncedly elongate, with dorsal teeth only,

extending along entire length; epigastric tooth distinctly separated from first rostral tooth.

Carapace with minute orbital, well developed antennal and hepatic spines; pterygostomian spine

140

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

lacking; postocular sulcus clearly distinct; gastro-orbital carina and orbito-antennal sulcus absent;

cervical sulcus relatively long but weak, without accompanying carina; hepatic carina and sulcus

present; branchiocardiac carina well developed; longitudinal suture well marked, transverse

suture lacking. Sixth abdominal somite without cicatrix. Telson armed with movable, lateral

spines.

Antennule lacking parapenaeid spine; antennular flagella filiform, markedly unequal in length,

both longer than carapace. Palp of first maxilla entire, broad, bearing small proximolateral lobule,

large on proximomesial margin, and acute spine accompanied by rigid setae on distomesial

margin. Fourth and fifth pereopods subflagelliform with entire dactyls (differing from those of

Xiphopenaeus in which they are multiarticulate). Basial spine on first and second pereopods and

on ischium of first.

Distribution of branchiae, epipods, and exopods in the genus Tanypenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

1 +r

Podobranchiae

Epipods

Exopods

Fig. 89. — Tanypenaeus caribaeus : A, Paratype, 6 18 mm cl, off Punta Faro, Colombia. Petasma, dorsal view.

— B, Holotype, 2 21 mm cl, off Punta Faro, Colombia. Thelycum. Scales = 1 mm.

Petasma symmetrical, semiclosed, produced distolaterally into pair of broad hornlike

projections. Appendix masculina thick, almost subglobular, about as wide as long. Thelycum

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

141

closed, with single plate of sternite XIV not produced into paired flaps, forming anteriorly

shallow pocket with relatively long and deeply excavate median protuberance of sternite XIII.

Seminal receptacles consisting of paired trilobed sacs: large posterior lobe and two anterior lobes,

one directed mesially, other laterally; sacs opening by slits on anterolateral extremities of sternite

XIV.

A detailed account and figures of the seminal receptacles were given by P£rez Farfante

(1972).

Species. — Tanypenaeus caribaeus Perez Farfante, 1972. Off Colombia, western Atlantic

Ocean.

Genus Trachypenaeopsis Burkenroad, 1934

Figures 90-92

Penaeus - Miers, 1884, Rep. Voy. Alert zool. Coll., : 564.

Penaeopsis - De Man, 1911, Siboga Exped., 39a: 53 [part].

Metapenaeus - Rathbun, 1906, Bull. U. S. Fish Comm., 23(3): 904 [part]; 1915, Proc. biol. Soc. Wash., 28:

117; 1920, Rapp. Betref. Vissch. Kol. Curasao, II, 2: 320 [part].

Trachypeneopsis Burkenroad, 1934a, Bull. Bingham Oceanogr. Coll., Yale Univ., 4(7): 40. — Kubo, 1949,

J. Tokyo Coll. Fish., 36(1): 361. — Burkenroad, 1983, Crust. Issues, 1: 282. — Dall et a/., 1990, Adv.

Mar. Biol., 27: 100.

Trachypenaeopsis [emendment of Trachypeneopsis Burkenroad, 1934a, under the plenary powers by the

International Commission of Zoological Nomenclature, 1969, Opinion 864, Bull. zool. Nom., 25(4/5): 139].

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1826, Bull. zool. Nom., 25(4/5): 139.

Fig. 90. — Trachypenaeopsis mobilispinis , 2 11.5 mm cl, Saba Bank, Lesser Antilles. Lateral view.

Scale = 5 mm.

Source:

142

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Type Species: By original designation, Metapenaeus mobilispinis Rathbun, 1915, Proc. biol.

Soc. Wash., 28: 117.

Type Locality: Cave Round Bay, Saba, Netherlands Antilles, 7 m.

Gender: Feminine.

Diagnosis. — Integument pubescent on anterior part of carapace. Rostrum deep, short, falling

distinctly short of distal margin of first antennular segment; armed with dorsal teeth only;

epigastric tooth well separated from rostral series. Carapace with small orbital, buttressed

antennal, and hepatic spines, lacking pterygostomian spine (pterygostomian margin rounded);

postocular sulcus deep; gastro-orbital carina lacking; orbito-antennal sulcus well defined; cervical

and hepatic sulci well marked, latter usually curving anteroventrally, accompanied by sharp

carina; branchiocardiac carina absent; longitudinal and transverse sutures lacking.

Sixth abdominal somite without cicatrix. Telson armed with three pairs of lateral spines, either

all movable and subapical pair projecting from strongly produced

shoulders, or anterior pairs movable and subapical pair fixed.

Antennule lacking parapenaeid spine; antennular flagella

short, unequal in length, dorsal ramus shorter than ventral, latter

about half to three-fifths as long as carapace. Palp of first maxilla

biarticulate, basal article relatively broad, narrowing to minute

distal article, with row of proximolateral setae, broadly rounded

or subtriangular, setose proximomesial lobule followed by long

spine. Basial spine and minute ischial spine on first pereopod

only. All epipods non-furcate (unique in family, as far as

known).

Petasma symmetrical, semiclosed, with ventrolateral lobule

produced into three flat distal processes, one terminal, remainder

lateral; dorsolateral lobule forming minute spout; exopod of first

pleopod in adult males uniquely bearing laterally two elongate

rigid projections, these contiguous along sinuous margins.

Appendix masculina roughly ovoid, projecting from thickened

dorsomedial margin of endopod. Thelycum closed, with paired

narrow lateral plates of stemite XIV, anteriorly flanking base of

median protuberance of sternite XIII; protuberance strong, with

broad base; posterior thoracic ridge bearing small, pointed,

median prominence. Seminal receptacles consisting of paired

membranous subspherical sacs opening by oblique slit on each

side of median protuberance immediately anterodorsal to anterior

margin of lateral plates, sacs uniquely attached posteriorly by

narrow septum.

Distribution of branchiae, epipods, and exopods in the genus Trachypenaeopsis.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

l+r

Podobranchiae

Epipods

Exopods

Fig. 91. — Trachypenaeopsis

mobilispinis , cj 8.9 mm cl,

Saba Bank, Lesser Antilles.

Petasma, dorsal view. Scale =

1 mm.

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

143

Fig. 92. — Trachypenaeopsis mobilispinis : A, <? 8.9 mm cl, Saba Bank, Lesser Antilles. Petasma and first

pleopods, ventral view. — B-C: 2 11.5 mm cl, Saba Bank, Lesser Antilles. B, Thelycum. C, Seminal

receptacles, dorsal view. Scales = 1 mm.

Species. — Trachypenaeopsis minicoyensis Thomas, 1972. Laccadive Sea.

Trachypenaeopsis mobilispinis (Rathbun, 1915). Bermuda; Gulf of Campeche; Turks and

Caicos Islands; Cuba; Jamaica; Saba, Netherlands West Indies; Brazil.

144

ISABEL P£REZ FARFANTE & BRIAN KENSLEY

Trachypenaeopsis richtersii (Miers, 1884). St. Helena Island; Madagascar; Mauritius;

Seychelles; Indonesia; Japan; Hawaii.

Genus Trachypenaeus Alcock, 1901

Figures 93-95

Penaeus - Bate, 1881. Ann. Mag. nat. Hist., (5)8: 173 [part, description of Penaeus anchoralis only].

Petieus (Trachypeneus) Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 15 [part].

Trachypeneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)16: 522 [part]. — Burkenroad, 1983, Crust. Issues, 1:

283 [part].

Trachypeneus (Trachypeneus) - Burkenroad, 1934a [September], Bull. Bingham oceanogr. Coll., 4(7): 49

[Division II, (part)].

Trachypeneus - Burkenroad, 1934b [December], Bull. Am. Mus. nat. Hist., 68(2): 94 [Division 2 (part)].

Trachypenaeus [emendment of Trachypeneus Alcock, 1901, under the plenary powers by the International

Commission on Zoological Nomenclature, 1969, Opinion 864, Bull. zool. Norn., 25(4/5): 139].

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1827, Bull. zool. Nom., 25(4/5): 139. — Kubo, 1949,

J. Tokyo Coll. Fish., 36(10: 391 [part]. — Dall, 1957, Austr. J. mar. Freshwat. Res., 8(2): 202 [part]. —

Racek and Dall, 1965, Verh. K. Akad. Wet., 56(3): 87 [part]. — Motoh and Buri, 1984, Res. Crust., (13-

14): 80 [part]. — Liu and Zhong, 1988, Penaeoid Shr. S. Ch. Sea, : 184 [part]. — Dall et al ., 1990, Adv.

Mar. Biol., 27: 102 [part].

Fig. 93. — Trachypenaeus anchoralis , 9 23 mm cl, off Northern Territory, Australia. Lateral view.

Scale = 10 mm.

Type Species: By original designation, Penaeus anchoralis Bate, 1881, Ann. Mag. nat. Hist.,

(5)8: 181.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOrD AND SERGESTOID SHRIMPS

145

Type Locality: South of New Guinea, [Torres Strait], 9°59'S, 139°42'E, 51 m, Challenger

sta. 188.

Gender: Masculine.

Diagnosis. — Body thickset. Integument thick, densely pubescent. Rostrum relatively short,

reaching distal end of second antennular segment; armed with dorsal teeth only; epigastric tooth

distinctly separated from first rostral tooth. Carapace with orbital, antennal, and hepatic spines

present; pterygostomian angle sharp but lacking spine; postocular sulcus shallow; short faint

cervical sulcus, clearly marked hepatic sulcus; orbito-antennal sulcus absent; branchiocardiac

carina absent; longitudinal suture weak, short, ending quite anterior to hepatic spine; transverse

suture absent. Abdomen with sixth somite lacking cicatrix. Telson with 3 pairs of movable lateral

spines, subapical pair projecting from slight shoulder.

Antennule lacking parapenaeid spine; antennular flagella

shorter than carapace. Palp of first maxilla entire, tapering to

blunt apex, produced into setose proximolateral lobule,

rounded setose proximomesial and small acute distomesial

lobules, latter bearing sharp spines; also armed distally with

ventrolateral row of sharp spinules. Fourth and fifth

pereopods with dactyl neither elongate nor subdivided. Basial

spine lacking on third maxilliped, present on first and second

pereopods; small ischial spine on first pereopod, lacking on

second. Epipod present on third pereopod only.

Petasma symmetrical, semiclosed, with lateral lobes

produced into very broad distolateral projections, extending

laterally, then turning forward and forming long, flat, twisted

process directed almost straight forward. Appendix masculina

subquadrangular with rounded corners. Thelycum closed,

with plate of sternite XIV continuous with median

protuberance, lacking anterior sternal invagination, thus

neither tranverse groove nor median pocket present; plate

markedly depressed anteromedially; median protuberance of

sternite XIII comparatively short, not produced caudally.

Seminal receptacles consisting of paired bilobed sacs, large

posterior lobe and small, depressed, anterolateral one;

strongly curved, slitlike openings lying on anteromesial side

of large lobe, joining cavity dorsal to posterolateral part of

median protuberance, near base of fourth pereopods.

Fig. 94. — Trachypenaeus

anchoralis, 6 13 mm cl, Van

Diemen Gulf, Northern Territory,

Australia. Petasma, dorsal view.

Scale = 1 mm.

Distribution of branchiae, epipods, and exopods in the genus

Trachypenaeus.

Somite

XIV

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

146

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Fig. 95. — Trachypenaeus anchoralis : A, 2 24 mm cl, Van Dienem Gulf, Northern Territory, Australia.

Thelycum. — B, 2 23 mm cl, off Northern Territory, Australia. Seminal receptacles, dorsal view.

Scales = 1 mm.

Remarks. — The characters that differentiate Trachypenaeus from Megokris are discussed

under "Remarks" in the treatment of the latter genus.

Species. — Trachypenaeus anchoralis (Bate, 1881). Taiwan; Torres Straits; Western

Australia, Northern Territory, Queensland, New South Wales, Australia.

Genus Trachysalambria Burkenroad, 1934

Figures 96-98

Penaeus - Stimpson, 1860, Proc. Acad. nat. Sci. Phiiad., 12: 44 [description of Penaeus curvirostris only], —

Kishinouye, 1900, J. Fish. Bur., Tokyo, 8(1): 6 [part, features of P. curvirostris only],

Peneus (Trachypeneus) - Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust.,: 15 [part],

Trachypeneus - Alcock, 1905, Ann. Mag. nat. Hist., (7)26: 522 [part], — Burkenroad, 1959, in Mission

R. P. Dollfus Egypte, Result, scient., 3(23-24): 285; 1983, Crust. Issues, 1: 281 [part].

Metapenaeus - Nobili, 1906, Ann. Sci. nat. Zool. Paris, (9)4: 20 [part].

Trachypeneus (Trachysalambria) Burkenroad, 1934a [September], Bull. Bingham oceanogr. Coll., 4(7): 49

[Division I, Section 2],

Trachypeneus - Burkenroad, 1934b [December], Bull. Am. Mus. nat. Hist., 68(2): 94 [Division I, Section 2],

Trachypenaeus - Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 391 [part], — Dall, 1957, Aust. J. mar. Freshwat.

Res., 8(2): 202 [part], — Motoh and Buri, 1984, Res. Crust., (13-14): 80. — Liu and Zhong, 1988, Penaeoid

Shr. S. Ch. Sea, : 184 [part], — Dallc? al, 1990, Adv. Mar. Biol., 27: 102 [part], — Hayashi, 1992, Dendr.

Crust. Japan Waters, : 137.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

147

Fig. 96. — Trachysalambria curvirostris, 2 24 mm cl, Mogi, Japan. Lateral view. Scale = 10 mm.

Type Species: By original designation, Trachypenaeus (Trachysalambria) curvirostris

(Stimpson)[= Penaeus curvirostris Stimpson], I860, Proc. Acad. nat. Sci. Philad., [12]: 44.

Type Locality: Port "Simoda" [Shimoda Ko], Japan.

Gender: Feminine.

Diagnosis. — Body thickset. Integument thick, densely pubescent, pubescence occasionally

in large patches. Rostrum relatively short, reaching approximately between base of second

antennular segment and distal margin of third; armed with dorsal teeth only; epigastric tooth

distinctly separated from first rostral tooth. Carapace with orbital, antennal, and hepatic spines

present; pterygostomian angle usually blunt, or rather sharp, always lacking spine; postocular

sulcus absent; orbito-antennal sulcus shallow; weak, short, moderately long, or absent cervical

sulcus, marked or indistinct hepatic sulcus; branchiocardiac carina extremely weak or lacking;

longitudinal suture short, faint, sometimes almost indistinct, ending anterior to hepatic spine;

transverse suture short, well marked or, usually, weak, sometimes extremely so. Abdomen with

sixth somite lacking cicatrix. Telson with 1-4, usually 3, pairs of movable lateral spines,

subapical pair projecting from slight shoulder.

Antennule lacking parapenaeid spine; antennular flagella shorter than carapace. Palp of first

maxilla entire, tapering to blunt apex, produced into setose proximolateral, rounded setose

proximomesial and small acute distomesial lobules, latter bearing sharp spines; also armed

distally with ventrolateral row of spinules. Fourth and fifth pereopods with dactyl neither

elongate nor subdivided. Basial spine lacking on third maxilliped, present on first pereopod and

usually on second; small ischial spine present or absent on first pereopod. Epipod present on first

through third pereopods, on second and third, or on third only.

148

ISABEL P&REZ FARFANTE & BRIAN KENSLEY

Fig. 97. — Trachysalambria curvirostris, 6 21 mm

cl, Hong Kong. Petasma, dorsal view. Scale =

1 mm.

Petasma symmetrical, semiclosed, with

lateral lobes produced distally into usually

large, hornlike or winglike projections,

extending either horizontally or curving

distinctly backwards; if lobes small, subovate

and horizontal. Appendix masculina

subglobular, or subquadrate with rounded

comers. Thelycum closed, with plate of stemite

XIV usually moderately long medially (except

in T. brevisuturae), its anterior margin

shallowly emarginate or produced in small

median prominence (in T. brevisuturae only);

anterior sternal invagination forming bracket¬

shaped groove extending anterolaterally dorsal

to plate, not extending posteriorly into median

pocket; median protuberance on sternite XIII

very strong, but not produced caudally.

Seminal receptacles consisting of paired bilobed

sacs, each with large posterior lobe and small

anterolateral one; their curved slitlike openings

on anteromesial side of large lobe continuing

into anterolateral part of bracket-shaped groove,

lying dorsal to posterolateral part of median

protuberance, near base of fourth pereopods.

Distribution of branchiae, epipods, and exopods in the genus Trachysalambria.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

1+r*

Podobranchiae

Epipods

i(-)

1 (-)

Exopods

*A rudimentary anteroventral arthrobranchia is found only in T. brevisuturae.

Remarks. — The members of Trachysalambria are rather closely allied to those of

Rimapenaeus ; however, they differ in that in the former the plate of stemite XIV is neither deeply

cleft anteriorly, at most slightly emarginate, nor produced into paired flaps; anterior sternal

invagination of XIV forming a typical bracket-shaped transverse groove and not a well defined

median pocket; and the distolateral projections of the petasma extend almost straight laterally or

with their tips curving slightly backwards. Also, in Trachysalambria, unlike that of

Rimapenaeus, the longitudinal suture of the carapace is short, not reaching the level of the

epigastric tooth; the basial spine is lacking on the third maxilliped, and the seminal receptacles are

bilobed, with their openings continuing into the common groove.

It should be pointed out that the validity of two of the species cited below, T. albicoma and

T. malaiana, has been questioned and needs to be confirmed.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

149

Fig. 98. — Trachysalambria curvirostris, $ 25 mm cl, off Natal South Africa: A, Thelycum. B, Seminal

receptacles, dorsal view. Scales = 1 mm.

Species. — Trachysalambria albicoma (Hayashi and Toriyama, 1980). Japan.

Trachysalambria aspera (Alcock, 1906). India; Andaman Islands; Indonesia; Philippines.

Trachysalambria brevisuturae (Burkenroad, 1934a). Eastern Pacific Ocean from Bahia

Almejas, southwest of Baja California, Gulf of California, Sinaloa, Gulf of Tehuantepec,

Mexico; El Salvador; northern Peru.

Trachysalambria curvirostris (Stimpson, 1860). Eastern Mediterranean; Natal, South Africa to

Tanzania; Red Sea; Madagascar; Yemen to South India including Persian Gulf; Sri Lanka;

Malaysia; Indonesia; Gulf of Tonkin; China; Hong Kong; Taiwan; Philippines; Japan; Korea;

New Guinea; Western Australia, Northern Territory, New South Wales, Australia.

Trachysalambria fulva (Dali, 1957). Malaysia; Indonesia; Philippines; New Guinea; Western

Australia, Queensland, Australia.

Trachysalambria longipes (Paulson, 1875). Red Sea; Gulf of Tonkin; South China Sea;

Japan.

Trachysalambria malaiana (Balss, 1933). Borneo; Gulf of Tonkin; South China Sea;

Philippines; New Guinea.

Trachysalambria villaluzi (Muthu and Motoh, 1979b). Philippines.

Genus Xiphopenaeus Smith, 1869

Figures 99-101

Penaeus - Heller, 1862b, Sber. Akad. Wiss. Wien, 45(1): 425.

Xiphopeneus Smith, 1869, Trans. Conn. Acad. Arts Sci., 2(1): 27; 1885b, Proc. U. S. natn. Mus., 8(11/12):

188. — Rathbun, 1901, Bull. U. S. Fish Commn., 20(2): 102. — Burkenroad, 1934b, Bull. Am. Mus. nat.

Hist., 63(2): 102; 1983, Crust. Issues, 1: 283. — Dallci al., 1990, Adv. Mar. Biol., 27: 106.

Source: MNHN, Paris

150

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Xiphopenaeus [emendment of Xiphopeneus Smith, 1869, under the plenary powers by the International

Commission on Zoological Nomenclature, 1969, Opinion 864, Bull. zool. Norn., 25(4/5): 139].

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1828, Bull. zool. Nom., 25(4/5): 139. — Holthuis, 1948,

Proc. K. ned. Akad. Wet., 51(9): 1105.

Fig. 99. — Xiphopenaeus kroyeri, 2 27 mm cl, northwest of Puerto Limon, Costa Rica. Lateral view.

Scale = 10 mm.

Type Species: By monotypy, Xiphopeneus hartii Smith, 1869, Trans. Conn. Acad. Arts Sci.,

2(1): 27 [= Xiphopenaeus kroyeri (Heller, 1862b)].

Type Locality: Caravelas, Estado da Bahia, Brazil.

Gender: Masculine.

Diagnosis. — Integument glabrous. Rostrum long, considerably overreaching antennular

peduncle, usually longer than carapace in adults, sinuous, styliform anteriorly; armed with dorsal

teeth only, situated basally; epigastric tooth distinctly separated from first rostral. Carapace with

orbital angle well marked, antennal, and hepatic spines present; pterygostomian angle produced

but lacking spine; postocular sulcus well marked; clearly distinct orbito-antennal sulcus; short,

almost indistinct cervical sulcus, clearly distinct orbito-antennal sulcus; hepatic sulcus and sharp

hepatic carina reaching only base of pterygostomian region and posteriorly merging with long

branchiocardiac sulcus and carina, respectively; longitudinal suture extending to about midlength

of carapace, transverse suture lacking (in adults). Abdomen with sixth somite bearing interrupted

cicatrix. Telson unarmed.

Antennule lacking parapenaeid spine; antennular flagella long, dorsal (twice or more as long

as carapace) longer than ventral. Palp of first maxilla entire, gradually tapering distally, produced

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

151

Fig. 100. — Xiphopenaeus kroyeri, <5

23 mm cl, Punta Araguapiche, Venezue¬

la. Petasma, dorsal view. Scale = 1 mm.

into small, triangular, setose proximolateral lobule,

broad, setose proximomesial and quite small, acute

setose distomesial lobules, latter armed with slender

spine; distolateral row of small spines present on ventral

surface. Fourth and fifth pereopods long, much longer

than third, subflagelliform, each with multiarticulate

dactyl. Basial and ischial spines on first pereopod only.

Petasma semiclosed, symmetrical, with lateral lobes

produced into large, hornlike, distolateral projections.

Appendix masculina subcircular. Male genital openings

subcoxal. In females, first pleopod lacking rudimentary

endopod. Thelycum closed, with single plate of sternite

XIV smooth, broad, not produced anteriorly into pair of

flaps, its anterolateral hoods much reduced; anterior

sternal invagination nearly as broad as sternite, forming

spacious pocket extending as far as posterior thoracic

ridge; median protuberance of sternite XIII also broad

but quite short. Paired seminal receptacles bilobed, each

consisting of large posterior lobe and small anterolateral

one; sinuous slitlike opening lying on anterior extremity

of posterior lobe, connecting with median pocket and

extending laterally dorsal to posterolateral extremity of

median protuberance.

Burkenroad's observations (1934b) indicate that the

spermatophore (sperm packets and plug substance) of

X. kroyeri is similar to that reported for Rimapenaeus (=

Trachypenaeus) similis by Bauer (1991).

Fig. 101. — Xiphopenaeus kroyeri , $ 27 mm cl. Northwest of Puerto Limon, Costa Rica: A, Thelycum.

B, Seminal receptacles, dorsal view. Scales = 1 mm.

152

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

The study of extensive material of specimens of this genus demonstrated that there are no

differences between those from the Atlantic and Pacific, that would justify their separation into

two species.

Distribution of branchiae, epipods, and exopods in the genus Xiphopenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Species. —Xiphopenaeus kroyeri (Heller, 1862a). Punta Piaxtla, Sinaloa, Mexico to Paita,

Peru; North Carolina to Gulf of Mexico, Caribbean Sea to Santa Catarina, Brazil.

Family SICYONIIDAE Ortmann, 1898

Sicyoninae Ortmann, 1898, Bronn’s Kl. Ordn. Tierreichs, (1st ed.) (5)2: 1121. — Balss, 1957, Bronn's KI. Ordn.

Tierreichs, (2nd ed.) 5(1)7(12): 1520.

Placed on the Official List of Family-Group Names in Zoology, International Commission on Zoological

Nomenclature, 1956b, Direction 54, Op. Decl. Int. Comm. zool. Norn., 12(26): 460.

Penaeidae - Barnard, 1950, Ann. Mus. Nat. Hist., 38: 580 [part].

Sicyonidae - Grey et al., 1983, Guide Austr. Prawns,: 13,31. — Dall etal., 1990, Adv. Mar. Biol., 27: 60.

Sicyoniidae - P£rez Farfante, 1978, FAO Ident. Sh., 6: SICYON. — Williams, 1984, Shr. Lob. Crabs All.

Coast U.S., : 42. — P£rez Farfante, 1988, NOAA Tech. Rep. NMFS, 64: iv, 22.

Diagnosis. — Body thickset. Integument rigid, pubescent. Rostrum armed with dorsal and

usually apical teeth, lacking ventral teeth, short, not overreaching antennular peduncle. Carapace

without postorbital and pterygostomian spines, bearing or lacking antennal spine; cervical sulcus

very weak or absent; hepatic carina weak, branchiocardiac carina strong to barely distinct.

Abdomen marked by transverse sulci, often tuberculate. Telson armed with pair of lateral fixed

subterminal spines.

Eye with optic calathus articulated directly to basal segment of eyestalk, intermediate segment

not apparent, without mesial tubercle; basal segment without ocular scale; ocular plate bearing

styliform mesial projection (stylet). Antennule with prosartema rudimentary, flagella short,

cylindrical. Third through fifth pleopods uniramous, lacking endopods (unique in Penaeoidea).

Exopod on first maxilliped, absent from second and third maxillipeds and all pereopods.

Pleurobranchia on somite IX only; rudimentary arthrobranchia on somite VII, two

arthrobranchiae on somites VIII through XIII, anteroventral one of XIII rudimentary;

podobranchia on second maxilliped; epipods on first and second maxillipeds and first three

pereopods. Petasma closed, its lateral lobes heavily sclerotized. Second pleopods in male bearing

appendix masculina only. Thelycum closed.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

153

Genus Sicyonia H. Milne Edwards, 1830

Figures 102-104

Cancer - BrOnnich, 1768, Ichthyol. Massil., : 102 [part].

Palaemon - Olivier, 1811, Encycl. method. Hist, nat., 8: 664 [part].

Hippolyte - De Haan, 1844, Fauna Jap. (Crust.), (6, 7): pi. 45, fig. 6, 9, 10 [part].

Sicyonia H. Milne Edwards, 1830, Annls Sci. nat., (1)19: 339. — Bate, 1888, Rep. scient. Results Voy.

Challenger, 24: 292. — Holthuis, 1952b, Bull. zool. Nom., 6: 339. — Burkenroad, 1983, Crust. Issues, 1:

282. — De Freitas, 1984, Invest. Rep. oceanogr. Res. Inst. Durban, 60: 1. — P£rez Farfante, 1985, Fish.

Bull., U. S., 83(1): 1.

Placed on Official List of Generic Names in Zoology, International Commission on Zoological Nomenclature,

1956a, Opinion 382, Name No. 922, Opin. Decl. Internat. Comm. Zool. Nom., 12(3): 45.

Ruvulus De Natale, 1850, Descriz. zool. Plojaria crost. Messina, : 20. (Published as synonym of Sicyonia

H. Milne Edwards, 1830). Type species: By monotypy: Sicyonia sculpta H. Milne Edwards, 1830, Annls. Sci.

nat., (1)19: 340 (= Cancer carinatus Briinnich, 1768, Ichthyol. Massil., : 102). Type locality: Bay of Naples.

Gender: Masculine.

Synhimantites Boeck, 1864, Forh. VidenskSelskr. Christ., (1863): 189. Type species: By monotypy,

Synhimantites typicus Boeck, 1864, Forh. VidenskSelsk. Christ., (1863): 189. Type locality: Norway.

Gender: masculine.

Eusicyonia Stebbing, 1914, Ann. S. Afr. Mus., 15: 25. [Substitute name for Sicyonia H. Milne Edwards, 1830,

Annls Sci. nat., (1)19: 339]. — Burkenroad, 1934a, Bull. Bingham oceanogr. Coll., Yale Univ., 4(7): 70;

1934b, Bull. Am. Mus. nat. Hist., 68(2): 116. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 437. —

Barnard, 1950, Ann. S. Afr. Mus., 38: 635. Type species: By monotypy for Sicyonia H. Milne Edwards,

1830: Sicyonia sculpta H. Milne Edwards, 1830, Annls Sci. nat., (1)19: 340. Type locality: Bay of Naples,

Mediterranean Sea. Gender: feminine.

Fig. 102. — Sicyonia carinata , 2 19.4 mm cl, Gibraltar, Mediterranean Sea. Lateral view. Scale = 10 mm.

Source

154

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Type Species: By subsequent designation, by Desmarest,

1858, Encycl. Hist. Nat. (Crust. Moll. Zooph.): 42, Sicyonia

sculpta H. Milne Edwards, 1830, Annls Sci. nat., (1)19: 340

(= Cancer carinatus Briinnich, 1768, Ichthyol. Massil., 1768:

102 ).

Type Locality: Bay of Naples, Mediterranean Sea.

Gender: Feminine.

Diagnosis. — Body robust, strongly sclerotized, of stony

appearance, variably pubescent. Rostrum armed with dorsal and

usually apical teeth, lacking ventral teeth, short, not

overreaching antennular peduncle, more often falling short of it.

Carapace without postorbital and pterygostomian spines,

bearing or lacking antennal spine; hepatic spine well developed;

cervical sulcus very weak or absent; hepatic carina weak;

branchiocardiac carina strong to virtually indistinct. Abdomen

marked by transverse sulci bordered by closely set setae, often

tuberculate; sixth somite bearing cicatrix. Telson armed with pair

of lateral fixed, subterminal spines.

Eye with optic calathus articulated directly to basial article of

eyestalk; basal article without scale; ocular plate armed with

styletlike projection. Antennule with prosartema rudimentary,

flagella short, subcylindrical. Palp of first maxilla entire, broad,

blunt to subacute apically, lacking or bearing weak

proximolateral lobule and produced mesially into broad proximal

lobule and much smaller distal lobule, latter armed with spine;

also bearing sublateral row of spinous setae. Both basial and

ischial spine present or absent on first pereopod. Third through

fifth pleopods lacking endopods.

Distribution of branchiae, epipods, and exopods in the genus Sicyonia.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

r+1

Podobranchiae

Epipods

Exopods

Fig. 103. — Sicyonia carinata, 6

19.4 mm cl, Gibraltar,

Mediterranean Sea. Petasma,

dorsal view. Scale = 1 mm.

Petasma closed, lateral lobes heavily sclerotized, depressed, its dorsolateral and ventrolateral

lobules each produced into distal projections. Appendix masculina small, roughly bellshaped but

with membranous terminal wall. Male genital openings sternal. Thelycum closed, with single

plate of sternite XIV flat or raised in paired, weak or prominent, lateral bulges; median plate of

sternite XIII roughly lanceolate, long, sometimes conspicuously overreaching sternite XI.

Seminal receptacles consisting of paired trilobed membranous sacs: large, longitudinally disposed

posterior lobe, and two small anterior lobes, one directed anteromesially, the other laterally;

openings lying between lateral hoods of sternite XIV and posterolateral margin of median plate of

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

155

stemite XIII. Seminal receptacles of several American species were studied by Perez Farfante

(1985) and Bauer (1986, 1991).

True spermatophores are not formed; only sperm in a fluid matrix is transferred to female

seminal receptacles, without any plug substance (Burkenroad, 1934b; Bauer, 1991).

Fig. 104 A. — Sicyonia carinata, 9 16 mm cl, Gibraltar, Mediterranean Sea. Thelycum. Scale = 1 mm.

Fig. 104 B. — Sicyonia disedwardsi, 9 30 mm cl, Golfo de Panama, Panama. Seminal receptacles, dorsal view.

Scale = 1 mm.

Species. — Sicyonia affinis Faxon, 1893. Costa Rica; Panama; Colombia; Northern Peru.

Sicyonia aliaffinis (Burkenroad, 1934a). Eastern Pacific Ocean from southwestern Baja

California Sur and Gulf of California to southern Mexico; Guatemala; Costa Rica; Panama;

Ecuador; Paita, northern Peru.

Sicyonia benthophila De Man, 1907. Indonesia.

Sicyonia bispinosa (De Haan, 1849). Indonesia; Japan.

Sicyonia brevirostris Stimpson, 1871. Eastern Pacific Ocean off southern Mexico; Western

Atlantic Ocean from Virginia to Florida, Gulf of Mexico; Yucatan; Bahamas; Cuba.

Sicyonia burkenroadi Cobb, 1971. North Carolina to Florida through Gulf of Mexico; West

Indies; Caribbean coast of Central and South America to Bahia, Brazil.

Sicyonia carinata (Briinnich, 1768). Banyuls, France; Mediterranean; east Atlantic Ocean;

Portugal; Mauritania; Guinea; Principe; Sao Tome; Congo; Angola.

Sicyonia cristata (De Haan, 1844). South China Sea; Hong Kong; Taiwan; Japan; northern

Australia.

Sicyonia curvirostris Balss, 1913a. Indonesia; South China Sea; Taiwan; Japan.

Sicyonia disdorsalis (Burkenroad, 1934a). Eastern Pacific, south-western Baja California

Sur, Gulf of California, Mexico to Pisco, Peru.

156

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Sicyonia disedwardsi (Burkenroad, 1934a). Eastern Pacific Ocean, southwestern Baja

California Sur and Gulf of California, to Colima, Mexico; Costa Rica; Panama.

Sicyonia disparri (Burkenroad, 1934a). Southwestern Baja California Sur and Gulf of

California to Nayarit, Mexico.

Sicyonia dorsalis Kingsley, 1878. North Carolina to Gulf of Mexico; Caribbean coast of

Central and South America to Central Brazil.

Sicyonia fallax De Man, 1907. Madagascar; Indonesia.

Sicyonia formosa Chan and Yu, 1985. Taiwan.

Sicyonia furcata Miers, 1878. Philippines.

Sicyonia galeata Holthuis, 1952a. Eastern Atlantic Ocean from Western Sahara to Angola.

Sicyonia inflexa (Kubo, 1949). Indonesia; Japan.

Sicyonia ingentis (Burkenroad, 1938). Monterey Bay, California to southern tip of Baja

California Sur; Gulf of California to Nayarit, Mexico.

Sicyonia japonica Balss, 1914a. Gulf of Aden; South China Sea; Hong Kong; Taiwan; Japan.

Sicyonia laevigata Stimpson, 1871. Eastern Pacific Ocean; southern Gulf of California; Costa

Rica; Panama; western Atlantic Ocean, North Carolina to southern Florida; Gulf of Mexico; West

Indies; Caribbean coast of Mexico, Central and South America to Santa Catarina, Brazil.

Sicyonia laevis Bate, 1881. Indonesia; New Guinea; Hawaii.

Sicyonia lancifer (Olivier, 1811). Mozambique; Red Sea; Gulf of Aden; Madagascar; Maidive

Islands; Sri Lanka; Malaysia; Indonesia; Gulf of Tonkin; South China Sea; Philippines; Japan.

Sicyonia longicauda Rathbun, 1906. East coast of South Africa; Mozambique; Taiwan; Japan;

Hawaii.

Sicyonia martini Perez Farfante and Boothe, 1981. Eastern Pacific Ocean, southwest Baja

California Sur, Gulf of California, southern Mexico to Panama.

Sicyonia mixta Burkenroad, 1946. Southwest Baja California Sur, and southern Gulf of

California; Panama; Puerto de Eten, northern Peru.

Sicyonia nasica Burukovsky, 1990. South-east Pacific Ocean near Sala-y-Gomez.

Sicyonia nebulosa (Kubo, 1949). Indonesia; Japan.

Sicyonia ocellata Stimpson, 1860. Sri Lanka; Singapore; Indonesia; Torres Straits; South

China Sea.

Sicyonia olgae Perez Farfante, 1980b. Dry Tortugas, Antilles to Surinam.

Sicyonia ommanneyi Hall, 1956. Singapore; Malaysia; South China Sea.

Sicyonia parafallax Crosnier, 1995. Red Sea.

Sicyonia parri (Burkenroad, 1934a). North Carolina through Gulf of Mexico; West Indies;

Colombia to Brazil.

Sicyonia parvula (De Haan, 1850). Indonesia; Japan.

Sicyonia penicillata Lockington, 1879. Baja California; Gulf of California; Costa Rica.

Sicyonia picta Faxon, 1893. Baja California Sur, Gulf of California to Islas Lobos de Afuera,

northern Peru.

Sicyonia rectirostris De Man, 1907. Indonesia.

Sicyonia stimpsoni Bouvier, 1905a. North Carolina to Florida through Gulf of Mexico; West

Indies; Caribbean coast of Mexico, Central and South America to Surinam.

Sicyonia trispinosa De Man, 1907. Indonesia.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

157

Sicyonia truncata (Kubo, 1949). Indonesia; Japan.

Sicyonia typica (Boeck, 1864). North Carolina through Gulf of Mexico; Antilles; Caribbean

coast of Central and South America to Santa Catarina, Brazil.

Sicyonia vitulans (Kubo, 1949). Japan.

Sicyonia wheeled Gurney, 1943. Bermuda.

Family SOLENOCERLDAE Wood-Mason, 1891

Solenocerina Wood-Mason, 1891, Ann. Mag. nat. Hist., (6)8: 275.

Parapetiaeinae - Ortmann, 1898, Bronn's Kl. Ordn. Tierreichs, (5)2: 1119, 1120 [part].

Peneinae - Alcock, 1901, Descr. Cat. Indian Deep-sea Crust.,: 13 [part].

Solenocerinae - Burkenroad, 1934b, Bull. Am. Mus. nat. Hist., 68: 63. — Ramadan, 1938, Scient. Rep. John

Murray Exped., 5(3): 56. — Crosnier, 1978, Faune Madagascar, 46: 14, 96. — Burkenroad, 1983, Crust.

Issues, 1: 281, 285. — Crosnier, 1985, Mar. Res. Indonesia, 24: 31; 1988, Bull. Mus. nat. Hist. Nat., Paris,

4(10)[A](3): 563; 1989, Sencken. Biol., 69: 379.

Placed on the Official List of Family-Group Names in Zoology, International Commission on Zoological

Nomenclature, 1961, Opinion 611, Bull. zool. Norn., 18(5): 307.

Penaeidae - Barnard, 1950, Ann. S. Afr. Mus., 38: 580 [part].

Solenoceridae - P£rez Farfante, 1978, FAO Sp. Indent. Sh., 6: 2. — Grey et al., 1983, Guide Austr. Pen.

Prawns, : 14. — Williams, 1984, Shr. Lob. Crabs Atl. Coast U.S., 17. — De Freitas, 1985, Investl. Rep.

oceanogr. Res. Inst., Durban, 57: 28. — P£rez Farfante, 1988, NOAA Tech. Rep. NMFS, 64: iv, 26. —

Dall et al., 1990, Adv. Mar. Biol., 27: 59.

Diagnosis. — Integument thin or firm. Rostrum laterally compressed, relatively short, seldom

reaching beyond antennular peduncle, armed with dorsal teeth, usually lacking ventral teeth.

Carapace with postorbital or postantennal and hepatic spines, antennal spine almost always pre¬

sent; orbital, branchiostegal, and pterygostomian spines present or absent. Cervical sulcus well

defined, reaching or almost reaching dorsal midline. Telson apically acute, usually armed with

subapical pair of fixed spines, occasionally with movable lateral spines, rarely lacking spines.

Eye with optic calathus bearing small mesial tubercle; basal segment of eyestalk produced into

strongly to barely developed ocular scale; ocular plate lacking styliform projection. Antennule

with prosartema variable in length, usually long and foliaceous, sometimes reduced to short rigid

projection; flagella usually very long, slender, subcylindrical or flattened. Exopods on all

maxillipeds and pereopods. Third through fifth pleopods biramous. Pleurobranchia on somites

IX through XIV; one or two rudimentary or small arthrobranchia on VII, two well developed

arthrobranchiae on VEU through XIII; podobranchia on second maxilliped, rarely on following

appendages, never on fourth and fifth pereopods.

Petasma open or semi-open. Second pleopod of male bearing appendix masculina, appendix

interna, and with basis produced into distolateral, ventrally inclined projection or spur. Thelycum

open.

Key to the Genera of the Family SOLENOCERIDAE

1. Movable lateral telsonic spines present . 2

— Movable lateral telsonic spines absent . 3

2. Post-cervical spine present, situated dorsal to hepatic spine .. Gordonella

— Post-cervical spine lacking . Ha lip or us

158

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

3. Orbital spine present . 4

— Orbital spine absent . 6

4. Both antennular flagella flattened . Solenocera

— Both antennular flagella subcylindrical or ventral flagellum flattened. 5

5. Both antennular flagella subcylindrical . Pleoticus

— Ventral antennular flagellum flattened. Mesopenaeus

6. Suprahepatic spine present . Haliporoides

— Suprahepatic spine absent . 7

7. Neither fourth nor fifth pereopods flagelliform . Cryptopenaeus

— Fifth or fourth and fifth pereopods flagelliform . 8

8. Fifth pereopod flagelliform, considerably longer than fourth .

. Hadropenaeus

— Fourth and fifth pereopods flagelliform . Hymenopenaeus

Genus Cryptopenaeus De Freitas, 1979

Figures 105-106

Cryptopenaeus De Freitas, 1979, Ann. S. Afr. Mus., 77: 123; 1985, Invest. Rep. oceanogr. Res. Inst. Durban,

57: 50.

Crassipenaeus Liu and Zhong, 1983, Chin. J. Oceanol. Limnol., 1: 171. Type species: By original designation,

Crassipenaeus sinensis Liu and Zhong, 1983, Chin. J. Oceanol. Limnol., 1(2): 171. Type locality:

Northeastern Guangdong near Shantou (Swatow), 261 m. Gender: Masculine.

Fig. 105. — Cryptopenaeus catherinae , 6 47.5 mm cl, off Cabo Santa Maria, Mozambique. Lateral view.

Scale — 1 mm.

Type Species: By original designation, Cryptopenaeus catherinae De Freitas, 1979 Ann

S. Afr. Mus., 77: 125, fig. 1 A-I.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

159

Type Locality: Off Cape Santa Maria, southern Mozambique, 26°06'S, 33°08'E, 350 m.

Gender: Masculine.

Diagnosis. — Body robust, firm. Integument glabrous or minutely setose punctate. Rostrum

short, not overreaching distal end of second antennular peduncle, armed with dorsal teeth

only. Carapace with small antennal spine and well developed postorbital, hepatic, and

pterygostomian spines; lacking orbital, suprahepatic, and branchiostegal spines; cervical carina

and sulcus well marked; branchiocardiac carina distinct to almost indistinguishable; submarginal

carina present. Telson armed with pair of short, fixed, subapical spines, movable marginal spines

lacking.

Antennular flagella similar, subcircular in cross section, as long as or slightly longer than

carapace. Mandibular palp biarticulate, articles subequal in length, distal one narrower than

proximal and tapering to rounded apex. Palp of first maxilla entire, short, gently tapering to blunt

apex, and bearing weak but densely setose, proximolateral lobule, and proximomesial

and distomesial lobules, latter with long spines. Basial spine on first and second pereopods,

ischial spine on first only. Lateral ramus of uropod bearing small or barely distinct distolateral

spine.

Petasma semi-open, symmetrical, with short dorsomedian lobule; ventromedian lobules

heavily sclerotized distally there ending in conspicuous process, and ventrolateral lobules

virtually occupied by ventral costae. Appendix masculina roughly trapezoidal; appendix interna

present; basis of pleopod produced distally into ventrolateral spur. Thelycum open, with

protuberances (2-3) or plates on anterior part of stemite XIV.

Fig. 106. — Cryptopenaeus catherinae , 6 47.5 mm cl, off Cabo Santa Maria, Mozambique: A, Petasma,

ventral view. Scale = 1 mm. — B, 9 63.2 mm cl, off Monte Bello, Mozambique. Thelycum.

Scale = 10 mm.

160

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Distribution of branchiae, epipods, and exopods in the genus Cryptopenaeus.

Somite

(Mxpl)

Vffl

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

xra

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Species. — Cryptopenaeus catherinae de Freitas, 1979. Natal, South Africa; southern

Mozambique; Kenya.

Cryptopenaeus clevai Crosnier, 1985. Indonesia; Wallis and Futuna Islands.

Cryptopenaeus crosnieri Perez Farfante and Kensley, 1985. New South Wales, Australia.

Cryptopenaeus sinensis (Liu and Zhong, 1983). Off Western Australia; South China Sea.

Genus Gordonella Tirmizi, 1960

Figures 107-109

Haliporus - Alcock and Anderson, 1894, J. Asiat. Soc. Beng., 63(2)3: 146 [part]. — Alcock, 1901, Descr. Cat.

Indian Deep-Sea Crust., : 22 [part]. — Bouvier, 1906b, Bull. Mus. oceanogr. Monaco, 81: 1 [part]; 1908a,

Result. Camp, scient. Prince Albert I, 33: 78 [part]. — Kensley, 1968, Ann. S. Afr. Mus., 50(12): 299. —

P£rez Farfante, 1977a, Fish. Bull., U. S., 75(2): 263 [part]. — Crosnier, 1978, Faune Madagascar, 46: 97

[part].

Gordonella Tirmizi, 1960, Scient. Rep. John Murray Exped. 1933-34, 10(7): 372. — Gordon, 1960 (in Tirmizi,

1960), Scient. Rep John Murray Exped 1933-34, 10(7): 379. — Crosnier, 1988, Bull. Mus. Hist. nat. Paris,

(4)10: 586.

Fig. 107. — Gordonella polyarthra (= G. villosa ), holotype, 2 58 mm cl, southern Arabian Sea. Lateral view.

Scale = 10 mm. (Adapted from Tirmizi, 1960, Fig. 86).

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

161

Fig. 108. — Gordonella kensleyi , holotype, 2 53.3 mm cl. New Caledonia: A, Lateral view. B. Telson, dorsal

view. Scale = 10 mm.

Type Species: By monotypy, Gordonella polyarthra Tirmizi, 1960, Scient. Rep. John

Murray Exped. 1933-34, 10(7): 373 [= Haliporus villosus Alcock and Anderson, 1894],

Type Locality: Southern Arabian Sea, 4°47'42"N, 72°35'36"E, 2727 m.

Gender: Feminine.

Diagnosis. — Integument glabrous or pubescent, soft, but carapace and abdomen reinforced

by numerous carinae. Rostrum short, not overreaching second antennular segment, with dorsal

teeth only; epigastric tooth separated from first rostral by more or less conspicuous interval.

Carapace high, distinctly convex posterior to cervical sulcus, with antennal, postantennal,

postcervical, hepatic, and pterygostomian spines present, postcervical spine and sometimes

pterygostomian followed by several denticles or spinules; well marked cervical sulcus crossing

postrostral carina where forming deep notch; hepatic, branchiocardiac, and branchiostegal sulci

clearly defined, postcervical sulcus absent; antennal, postantennal, cervical, hepatic,

branchiocardiac, sublateral, subhepatic-branchiostegal, and submarginal carinae present.

Abdomen slender, with dorsomedian carina along entire length, on each somite ending or not in

spine. Telson, tapering to point, armed with subapical pair of fixed spines preceded by three

pairs of widely spaced spinules.

Eye with well developed cornea, densely pigmented; bearing mesial tubercle; ocular scale

reduced. Prosartema absent or well developed but never foliaceous; stylocerite clearly defined;

antennular flagella very long, equal, filiform. Mandibular palp triarticulate, first article very short,

annular; second very elongate; third slender and much shorter than second. Palp of first maxilla

entire. Lateral ramus of uropod with distolateral spine falling conspicuously short of distal

margin of lamella.

Petasma open, symmetrical, dorsomedian lobule usually short, about 0.33 length of

petasma, ventromedian lobule markedly curved distally, lateral lobule without conspicuous distal

thickenings. Appendix masculina longer than appendix interna, latter somewhat flattened, basis

162

ISABEL P£REZ FARFANTE & BRIAN KENSLEY

of pleopod produced distolaterally into foliaceous projection or spur. Thelycum open, with

strong protuberance on stemite XIV and smaller one on stemite XIII.

Distribution of branchiae, epipods, and exopods in the genus Gordonella *

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(Pi)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

♦Above formula applies to G. kensleyi and G. paravillosa ; that of G. villosa unknown.

Fig. 109 A. — Gordonella paravillosa , 6 46 mm cl, Gulf of Boni, Celebes, Indonesia. Petasma, dorsolateral view

of right half. Scale = 1 mm.

Fig. 109 B. — Gordonella kensleyi , holotype, 2 53.5 mm cl, New Caledonia. Thelycum. Scale = 1 mm.

Species. — Gordonella kensleyi Crosnier, 1988. South-east Atlantic Ocean off Cape of Good

Hope; south of Mozambique; New Caledonia.

Gordonella paravillosa Crosnier, 1988. Celebes, Indonesia; east coast of Australia.

Gordonella villosa (Alcock and Anderson, 1894). North-west Indian Ocean in vicinity of

Laccadive Islands; north of Chagos Archipelago.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

163

Genus Hadropenaeus Perez Farfante, 1977

Figures 110-112

Solenocera - Bate, 1881, Ann. Mag. nat. Hist., (5)8: 185 [part].

Hymenopenaeus - Smith, 1885b, Proc. U. S. natn. Mus., 8(11/12): 179 [part]. — Burkenroad, 1936b, Bull.

Bingham oceanogr. Coll., 5(2): 102 [part]. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 212 [part].

Philonicus - Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 273 [part].

Pleoticus - Bate, 1888, Rep. scient. Results Voy. Challenger, 24: xii [part, excluding Philonicus miilleri Bate,

1888 = Pleoticus muelleri (Bate, 1888), and Philonicus pectinatus Bate, 1888 = Solenocera pectinata Bate,

1888].

Haliporus - Bouvier, 1906b, Bull. Mus. oceanogr. Monaco, 81:1 [part]. — A. Milne Edwards and Bouvier,

1909, Mem. Mus. comp. Zool. Harv., 27: 206 [part]. — De Man, 1911, Siboga Exped., 39a: 31.

Hadropenaeus Perez Farfante, 1977a, Fish. Bull., U. S., 75(2): 315. — Liu and Zhong, 1983, Chin. J. Oceanol.

Limnol., 1(2): 172. — Hayashi, 1992, Dendr. Crust. Jap. Waters, : 170.

Fig. 110. — Hadropenaeus affinis, $ 21.5 mm cl, Mona Lisa Passage, off Puerto Rico. Lateral view (third

pereopod slightly raised). Scale = 5 mm. (From Perez Farfante, 1977).

Type Species: By original designation, Hymenopenaeus modestus Smith, 1885b, Proc. U. S.

natn. Mus., 8(11/12): 183.

Type Locality: [Off Bethany Beach, Delaware], 38°31'N, 73°21'W, 285 m.

Gender: Masculine.

Diagnosis. — Body stout, integument firm. Rostrum short, not overreaching distal margin

of first antennular segment, deep, ventral margin pronouncedly convex; armed with dorsal teeth

Source

164

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

only; epigastric tooth and first rostral separated by

interval equal to, or only slightly greater than that

between first and second rostral teeth. Carapace

proportionately short; orbital and pterygostomian

spines absent; postorbital, antennal, hepatic, and

branchiostegal spines present. Cervical sulcus long,

almost reaching middorsum of carapace; hepatic

sulcus deep, subhorizontal posteriorly, inclined

anteroventrally from depressed area below hepatic

carina; branchiocardiac carina and sulcus absent;

posthepatic and submarginal carinae lacking. Telson

with pair of conspicuous fixed lateral spines.

Prosartema long, flexible; antennular flagella

longer than carapace, usually subcylindrical, ventral

flagellum occasionally depressed. Mandibular palp

biarticulate, articles moderately broad, distal one as

long as or slightly shorter than basal, and tapering to

blunt apex. Palp of first maxilla entire, gently

narrowing to rounded apex, lateral margin with

brush of short setae, mesial margin bearing row of

longer setae, and apex with considerably longer

ones. Fifth pereopod subflagelliform and

considerably longer than fourth. Basial, ischial, and

meral spines on first pereopod. Exopods well

developed on all maxillipeds and pereopods. Lateral

ramus of uropod armed with distolateral spine

reaching distal margin of lamella. Single

arthrobranch on somite VII small, bearing numerous branched filaments.

Petasma semi-open, symmetrical, with ventral costa free from heavily sclerotized, platelike

terminal part of ventrolateral lobule. Appendix masculina and appendix interna present; basis of

second pleopod produced distally into subelliptical or paddlelike ventrolateral spur. Thelycum

open.

Fig. 111. — Hadropenaeus affinis , 2 19 mm

cl, east of Cape Island, South Carolina. Thely¬

cum. Scale = 1 mm. (From PSrez Farfante,

1977).

Distribution of branchiae, epipods, and exopods in the genus Hadropenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

xrv

(P5)

Pleurobranchiae

Arthrobranchi ae

Podobranchiae

Epipods

Exopods

Species. —Hadropenaeus affinis (Bouvier, 1906b). Western Atlantic Ocean from North

Carolina to Florida; Gulf of Mexico; Caribbean; Cape Verde Islands.

Hadropenaeus lucasii (Bate, 1881). Madagascar; Maidive Islands; India; Indonesia; South

China Sea; Philippines; Japan; New Guinea; New South Wales, Australia; Hawaii; Wallis and

Futuna Islands.

Source . MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

165

Hadropenaeus modestus (Smith, 1885b). Western Atlantic Ocean from Delaware to Florida,

Gulf of Mexico, Bahamas, Caribbean, and northern Brazil.

Hadropenaeus spinicauda Liu and Zhong, 1983. South China Sea.

Fig. 112. — Hadropenaeus affinis , 3 16 mm cl, southeast of Cape Fear, North Carolina: A, Petasma, dorsal view

of right half. B, Petasma, ventral view. Scale = 1 mm. (From PErez Farfante, 1977).

Genus Haliporoides Stebbing, 1914

Figures 113-114

Penaeopsis - Faxon, 1893, Bull. Mus. comp. Zool. Harv., 24(1): 212.

Faxonia - Bouvier, 1905a, C. r. hebd. S6anc. Acad. Sci., Paris, 140: 981 [part, excluding type species, Penaeopsis

ocularis Faxon, 1895 = Pleoticus robustus (Smith, 1885b)].

Haliporus - Bouvier, 1906b, Bull. Mus. ocdanogr. Monaco, 81:1 [part]. — De Man, 1911, Siboga Exped., 39a:

31 [part],

Haliporoides Stebbing, 1914, Ann. S. Afr. Mus., 15: 20. — PErez Farfante, 1977a, Fish. Bull., U. S., 75(2):

289. — De Freitas, 1985, Invest. Rep. oceanogr. Res. Inst. Durban, 57: 37. — Hayashi, 1992, Dendr. Crust.

Jap. Waters, : 174.

Hymenopenaeus - Burkenroad, 1936b, Bull. Bingham oceanogr. Coll., 5(2): 102 [part], — Crosnier, 1978,

Faune Madagascar, 46: 108, 112 [part].

Parahaliporus Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 207. Type species: by original designation, Parahaliporus

sibogae De Man, 1907 [= Haliporus sibogae De Man, 1907], Notes Leyden Mus., 29(2): 138. Type locality:

Syntypes from four localities from East Flores Sea to Kai Islands (type locality not restricted). Gender:

Masculine.

Hymenopenaeus (Haliporoides) - Barnard, 1950, Ann. S. Afr. Mus., 38: 619.

Type Species: By monotypy, Haliporoides triarthrus Stebbing, 1914, Ann. S. Afr. Mus.,

Source:

166

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Type Locality: Off East London NW1/2N, 18 mi (929 km), Cape Province, South Africa,

457-549 m.

Gender: Masculine.

Fig. 113. — Haliporoides triarthrus vniroi, $ 43 mm cl, off llha do Bazaruto, Mozambique. Lateral view.

Scale = 10 mm.

Diagnosis. — Body moderately robust, integument firm. Rostrum relatively long, extending

at least to, often beyond, second antennular peduncle segment, ventral margin straight or

concave; armed with dorsal and, frequently, with ventral teeth; epigastric tooth separated from

rostral teeth by interval noticeably longer than spaces between latter. Carapace relatively elongate;

orbital and branchiostegal spines absent; postorbital, antennal, pterygostomian, hepatic, and

suprahepatic spines present. Cervical sulcus deep, long, extending to, but not across, middorsum

of carapace; hepatic sulcus deep to shallow, hepatic carina sharp to weak, turning anteroventrally

from almost horizontal posterior part and reaching base of pterygostomian spine; orbito-antennal

sulcus usually well developed, branchiocardiac carina and sulcus well marked; submarginal

carina sharp. Telson with pair of conspicuous, subterminal, fixed lateral spines.

Prosartema moderately long, broad and flexible; antennular flagella similar, subcylindrical and

long, not less than 3 times carapace length. Mandibular palp triarticulate, proximal article short

and narrow, intermediate one larger, scalene triangular in shape, and distal article considerably

shorter and narrower than middle one and tapering to blunt apex. Palp of first maxilla entire,

gently narrowing to rounded apex, and bearing short proximal setae on lateral margin and

subproximal and subapical tufts of longer setae on mesial margin. Fourth and fifth pereopods not

flagelliform, about as thick as preceding ones. Basial spine present or lacking on first pereopod.

Exopods quite small. Lateral ramus of uropod armed with subterminal distolateral spine. Single

arthrobranch on somite VII rather well developed, with numerous branched filaments.

Petasma open, with ventral costa distally fused to flexible flap of ventrolateral lobule.

Appendix masculina and appendix interna present; base of endopod of second pleopod produced

into very short distolateral spur. Thelycum open.

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

167

Distribution of branchiae, epipods, and exopods in the genus Haliporoides.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

XII

(P3)

(P4)

xrv

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 114. — Haliporoides triarthrus vniroi : A, 6 36 mm cl, off Ilha do Bazaruto, Mozambique. Petasma,

ventral view of left half (extended). — B, 2 43 mm cl, off Ilha do Bazaruto, Mozambique. Thelycum.

Scales = 1 mm.

Species. — Haliporoides cristatus Kensley, Tranter, and Griffin, 1987. East coast of

Australia.

Haliporoides diomedeae (Faxon, 1893). Eastern Pacific Ocean from Panama to southern

Chile.

Haliporoides sibogae australiensis Kensley, Tranter, and Griffin, 1987. South-east and east

coast of Australia.

Haliporoides sibogae madagascariensis Crosnier, 1978. Madagascar; Reunion.

Haliporoides sibogae sibogae (De Man, 1907). Malaysia; Indonesia; South China Sea;

Taiwan; Philippines; Japan; Australia; New Zealand.

Haliporoides triarthrus triarthrus (Stebbing, 1914). Cape of Good Hope to Natal, South

Africa.

Haliporoides triarthrus vniroi Crosnier, 1978. East coast of South Africa; Mozambique.

Source:

168

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Genus Haliporus Bate, 1881

Figures 115-117

Haliporus Bate, 1881, Ann. Mag. nat. Hist., (5)8: 171, 185 [part]; 1888, Rep. scient. Results Voy. Challenger,

24: 288 [part]. — Faxon, 1893, Bull. Mus. comp. Zool. Harv., 24(7): 213 [part]; 1895, Mem. Mus. comp.

Zool. Harv., 18: 189. — Alcock and Anderson, 1899, Ann. Mag. nat. Hist., (7)3: 280. — Alcock, 1901,

Descr. Cat. Indian Deep-Sea Crust., : 22 [part]. — Bouvier, 1906b, Bull. Mus. oceanogr. Monaco, 81:1

[part]; 1908a, Result. Camp, scient. Prince Albert I, 33: 78 [part]. — Burkenroad, 1934b, Bull. Am. Mus.

Nat. Hist., 6(2): 64 [part]; 1936b, Bull. Bingham oceanogr. Coll., 5(2): 100 [part]. — P£rez Farfante, 1977a,

Fish. Bull., U. S., 75(2): 263 [part]. — Crosnier, 1978, Faune Madagascar, 46: 97 [part]. — Burkenroad,

1983, Crust. Issues, 1: 281. — De Freitas, 1985, Investl. Rep. oceanogr. Res. Inst., Durban, 57: 33. —

Crosnier, 1988, Bull. Mus. Hist. nat. Paris, (4)10A(3): 563.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1812, Bull. zool. Nomen., 25(4/5): 139.

Hymenopenaeus - Burkenroad, 1936b, Bull. Bingham Oceanogr. Coll., 5(2): 102 [part]. — Kensley, 1977, Ann.

S. Afr. Mus., 74(2): 27.

Fig. 115. — Haliporus taprobanensis , 5 45.5 mm cl, northeast of Sombrero Island, Balayan Bay, Philippines.

Lateral view (carapace displaced dorsally). Scale = 10 mm.

Type Species: By subsequent designation, by Fowler, 1912, Ann. Rep. N. J. State Mus.,

1911(2): 542, Haliporus curvirostris Bate, 1881, Ann. Mag. nat. Hist., (5)8: 185.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

169

Type Locality: South of Low Archipelago, Pacific Ocean, 32°36'S, 137°43'W, 4343 m.

Gender: Masculine.

Diagnosis. — Integument glabrous or pubescent.

Rostrum short or moderately long, not overreaching

third antennular segment, laterally compressed, with

dorsal teeth only; epigastric tooth separated from first

rostral by interval equal to or slightly longer than that

between first and second teeth. Carapace moderately

high, with long shallow dorsomedian depression at

level of cervical sulcus; antennal, postantennal,

hepatic, and pterygostomian spines present;

postcervical spine lacking; well marked cervical sulcus

crossing postrostral carina where forming deep notch;

postcervical sulcus absent, antennal and hepatic carinae

variable in length, long branchiocardiac carina

accompanied by well marked sulcus. Abdomen

slender, bearing dorsomedian carina on all somites, on

each one ending or not in spine. Telson tapering to

terminal point, armed with pair of subapical spines

preceded by three pairs of marginal, movable spinules

relatively widely spaced.

Eye with cornea broader than, or same width as,

optical calathus, only slightly pigmented, bearing

mesial tubercle; ocular scale indistinct or small.

Prosartema little or well developed, never foliaceaous;

stylocerite well developed; antennular flagella very

long, equal, filiform. Mandibular palp triarticulate,

first article very short, second article long and very

broad, third slender and shorter than second. Palp of first maxilla entire. Fourth pereopod

slightly shorter or longer than preceding one. Lateral ramus of uropod with strong distolateral

spine falling conspicuously short of distal margin of lamella.

Fig. 116. — Haliporus taprobanensis , 2

45.5 mm cl, northeast of Sombrero

Island, Balayan Bay, Philippines.

Thelycum. Scale = 5 mm.

Distribution of branchiae, epipods, and exopods in the genus Haliporus. *

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(PS)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

1 (-)

l(-)

Epipods

Exopods

* In H. curvirostris , podobranchia on first pereopod small, that on second and third pereopods vestigial,

exopods of pereopods rudimentary. In H. taprobanensis , podobranchia on third maxilliped usually small, but size

variations have been observed, as in those of the first and second pereopods (Crosnier, 1988). In H. thetis ,

podobranchia of first three pairs of pereopods small.

Source:

170

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Petasma open, symmetrical, dorsomedian lobule short or very short, 0.15-0.30 length of

petasma; ventromedian lobule elongate, ending in outwardly curved spatula, dorsolateral lobule

sinuous distally, ventrolateral lobule distally free from contiguous lobule, relatively narrow to

broadened and slightly to strongly curved. Endopod of second pleopod bearing appendix

masculina, latter longer than appendix interna, with two or three concave surfaces; appendix

interna subcylindrical or flattened; basis of pleopod produced laterally into foliaceous projection

or spur more or less curved distally. Thelycum open, with strong protuberance on sternite XIV

and smaller one on stemite XIII.

Fig. 117. — Haliporus taprobanensis, 3 23 mm cl, Diuata Point, Philippines. Petasma (extended), dorsal view.

Scale = 5 mm.

Species. — Haliporus curvirostris Bate, 1881. North-east of Mariana Islands; south-west of

Tuamotu Archipelago; north-west Pacific Ocean.

Haliporus taprobanensis Alcock and Anderson, 1899. Off Natal, South Africa; Madagascar

India; Indonesia; Philippines.

Haliporus thetis Faxon, 1893. North-west of Madagascar; Reunion; central Indian Ocean -

India; Galapagos Islands.

Genus Hymenopenaeus Smith, 1882

Figures 118-120

Haliporus - _Bate, 1881, Ann. Mag. nat. Hist., (5)8: 185 [part, excluding Haliporus curvirostris Bate, 18811 —

Bate, 1888, Rep scient. Results Voy. Challenger, 24: 284 [part], — Alcock, 1901, Descr. Cat. Indian Deep-

Sea Crust., : 22 [part], — Bouvier, 1906b, Bull. Mus. oceanogr. Monaco, 81: 1 [part]; 1908a, Result. Camp

scient Pnnce Albert I 33: 78 [part]. — A. Milne Edwards and Bouvier, 1909, Mem. Mus. comp. Zool.

Harv., 27: 206. — De Man, 1911, Siboga Exped., 39a: 31 [part].

Hymenopenaeus Smith, 1882, Bull. Mus. comp. Zool. Harv., 10(1): 91. — Burkenroad, 1936b, Bull Bingham

1967 n< RMl ’. 5 M ): 102 ,o [ ?oT ] ;n7 KUB °’ ^ 1 T ° ky ° C ° 11- F ‘ Sh- ’ 36(10): 212 [!»«*]■ ~ Holthu.S,

1962, Bull. zool. Norn., 19(2): 108. — PErez Farfante, 1977a, Fish. Bull., U. S„ 75(2): 263-266. —

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

171

Crosnier, 1978, Faune Madagascar, 46: 100. — De Freitas, 1985, Invest. Rep. oceanogr. Res. Inst. Durban,

57: 28. — Liu and Zhong, 1988, Pen. Shr. S. China Sea, : 179. — Hayashi, 1992, Dendr. Crust. Jap. Waters,

: 179.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1816, Bull. zool. Norn., 25(4/5): 139.

Hymenopeneus - Burkenroad, 1983, Crust. Issues, 1: 281.

Fig. 118. — Hymenopenaeus debilis , 3 8.5 mm cl, south of Grand Terre Island, Louisiana. Lateral view.

Scale = 5 mm. (From PErez Fare ante, 1977).

Type Species: By monotypy, Hymenopenaeus debilis Smith, 1882, Bull. Mus. comp. Zool.

Harv., 10(1): 91.

Type Locality: Syntypes from three localities, off Georgia and North Carolina, U.S.A. (type

locality not restricted).

Gender: Masculine.

Diagnosis. — Body slender, integument thin, flexible and glabrous, rarely with photophores.

Rostrum variable in length, reaching between distal 0.25 length of first antennular segment and

end of peduncle; ventral margin straight; usually armed with dorsal teeth only, occasionally also

with ventral teeth; epigastric and first rostral teeth separated from remaining teeth by relatively

Source:

172

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

long interval. Carapace short to moderately long; orbital spine lacking; postorbital, antennal,

hepatic, and branchiostegal spines present; pterygostomian, suprahepatic, and post-cervical

spines present or absent. Cervical sulcus deep, long, extending to, but not across, middorsum of

carapace, accompanying carina sharp; hepatic sulcus and carina well marked posteriorly, present

or absent anteriorly; branchiocardiac carina sharp, accompanying sulcus deep; posthepatic and

submarginal carinae present. Telson armed with pair of conspicuous, fixed, subapical spines.

Prosartema long, flexible; antennular flagella similar, filiform and longer than carapace.

Mandibular palp triarticulate, articles relatively narrow, proximal one very small, distalmost much

shorter than middle one and tapering to apex. Palp of first maxilla entire, gently narrowing to

blunt apex and bearing proximal, inconspicuous setose lobule on lateral margin and with mesial

margin broadly rounded or produced into lobule proximally and smoothly concave or bearing

small lobule distally, both lobules fringed with shorter setae than those of apical tuft. Fourth and

fifth pereopods extremely long and flagelliform. Basial spine and usually ischial spine on first

pereopod. Only one arthrobranchia on somite VII. Exopods small. Lateral ramus of uropod

armed with distolateral spine reaching distal margin of lamella (terminal spine).

Petasma semi-open, with distal part of ventral costa fused to flexible flap of ventrolateral

lobule; ventromedian lobule usually produced into paired conspicuous processes distally.

Appendix masculina usually foliaceous and produced proximolaterally into conspicuous lobe;

appendix interna elongate-ovate; basis of pleopod produced distally into ventrolateral spur.

Thelycum open, with stemite XIV raised in median protuberance or strong ridge and stemite XIII

often bearing conspicuous median plate.

Distribution of branchiae, epipods, and exopods in the genus Hymenopenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

xin

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

r or s

Podobranchiae

Epipods

Exopods

Species. — Hymenopenaeus aphoticus Burkenroad, 1936b. Western Atlantic Ocean from

Florida to Gulf of Mexico; Caribbean Sea; Azores; Morocco.

Hymenopenaeus chacei Crosnier and Forest, 1969. North-east Atlantic Ocean from Mauritania

to Angola.

Hymenopenaeus debilis Smith, 1882. North-western Atlantic Ocean from New Jersey to Gulf

of Mexico; West Indies; Belize; Nicaragua; Panama; Venezuela; Colombia; Guyana; north-east

Atlantic Ocean off Azores; Cape Verde Islands; Canary Islands; Morocco.

Hymenopenaeus doris (Faxon, 1893). Eastern Pacific Ocean from Baja California, Gulf of

California, Isla del Coco, Costa Rica to Guanape, Peru.

Hymenopenaeus equalis (Bate, 1888). East coast of Africa; Arabian Sea; Sri Lanka; Andaman

Islands; Indonesia; South China Sea; Philippines; Taiwan; Japan; Wallis and Futuna Islands;

Hawaii.

Hymenopenaeus fattahi Ramadan, 1938. Arabian Sea.

Hymenopenaeus furici Crosnier, 1978. Madagascar.

Source: MNHN . Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

173

Fig. 119. — Hymenopenaeus debilis : A-B. <? 13 mm cl, off Cape Canaveral, Florida. A, Petasma, dorsolateral

view of left half. B , Idem, ventral view. — C, 9 15.5 mm cl, off Dominica Island, Lesser Antilles.

Thelycum. Scales = 1 mm. (From P£rez Farfante, 1977).

Hymenopenaeus halli Bruce, 1966. Off Natal, South Africa; Madagascar; Indonesia; South

China Sea; Philippines; Japan; New South Wales, Australia; Wallis and Futuna Islands.

Hymenopenaeus laevis (Bate, 1881). Western Atlantic Ocean from Massachusetts to

Bahamas; Bermuda; eastern Atlantic Ocean from Mauritania to Equatorial Guinea; Arabian Sea;

Bay of Bengal; Philippines; north-west Pacific Ocean.

Hymenopenaeus neptunus (Bate, 1881). Bay of Bengal; Indonesia; Philippines; Wallis and

Futuna Islands.

Hymenopenaeus nereus (Faxon, 1893). Eastern Pacific Ocean from Costa Rica to Ecuador;

Galapagos Islands.

Hymenopenaeus obliquirostris (Bate, 1881). Indonesia; off Kermadec Islands; Hawaii.

Hymenopenaeus propinquus (De Man, 1907). Zanzibar; Gulf of Aden; Madagascar; Reunion;

Maidive Islands; Saya de Malha Bank; Indonesia; South China Sea; Philippines; New South

Wales, Australia; Wallis and Futuna Islands.

Hymenopenaeus sewelli Ramadan, 1938. Maidive Islands; central Indian Ocean.

Hymenopenaeus tuerkayi Crosnier, 1995. Red Sea.

Genus Mesopenaeus Perez Farfante, 1977

Figures 120-121

Parartemesia - Bouvier, 1905b, C. r. hebd. S6anc. Acad. Sci., Paris, 141: 747 [part, excluding Parartemesia

carinata Bouvier, 1905b = Pleoticus muelleri (Bate, 1888)].

174

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

Haliporus - Bouvier, 1906b, Bull. Mus. oc£anogr. Monaco, 81:1 [part]. — A. Milne Edwards and Bouvier,

1909, Mem. Mus. comp. Zool. Harv., 27: 206 [part].

Hymenopenaeus - Burkenroad, 1936b, Bull. Bingham oceanogr. Coll., 5(2): 102 [part].

Mesopenaeus Perez Farfante, 1977a, Fish. Bull., U. S„ 75(2): 331. — P6rez Farfante and Ivanov, 1982,

J. Crust. Biol., 2(2): 303. — Crosnier, 1986b, Indo-Malayan Zool., 3: 19. — Hayashi, 1992, Dendr. Crust.

Jap. Waters, : 184.

Fig. 120. — Mesopenaeus tropicalis, 9 25 mm cl, east of Cayos de Albulquerque, Colombia. Lateral view.

Scale = 10 mm. (From Perez Farfante, 1977).

Type Species: By original designation, Parartemesia tropicalis Bouvier, 1905b, C. r. hebd.

Seanc. Acad. Sci., Paris, 141: 748.

Type Locality: Gulf of Mexico, exact position indeterminable [see P6rez Farfante, 1977a

Fish. Bull., U. S„ 75(2): 341],

Gender: Masculine.

Diagnosis. — Body stout, integument firm. Rostrum short, not overreaching base of second

antennular segment, deep, with ventral margin pronouncedly convex; armed with dorsal teeth

only; epigastric and first rostral separated by interval similar to, or only slightly longer than that

between first and second rostral teeth. Carapace proportionately short; orbital, postorbital,

antennal, and hepatic spines present; pterygostomian, branchiostegal, and suprahepatic spines

absent. Cervical sulcus long, almost reaching middorsum of carapace; hepatic sulcus deep, nearly

horizontal posteriorly to level of hepatic spine, from there turning anteroventrally and reaching

pterygostomian region; hepatic carina sharp anteriorly, indistinct posteriorly; branchiocardiac

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

175

carina and sulcus, posthepatic and submarginal carinae lacking. Telson with pair of conspicuous,

fixed, posterolateral spines.

Prosartema long and flexible; antennular flagella shorter to slightly longer than carapace and

dissimilar; dorsal flagellum subcylindrical and slender, ventral flagellum conspicuously

depressed. Mandibular palp biarticulate, articles broad, distal nearly as long as penultimate one

and tapering to blunt apex. Fourth and fifth pereopods rather stout proximally, slender distally,

fifth usually moderately, instead of considerably longer than fourth. Basial and ischial spines on

first pereopod. Exopods well developed. Lateral ramus of uropod armed with terminal (reaching

distal margin of lamella) distolateral spine. Single arthrobranch on somite VII rudimentary.

Fig. 121. — Mesopenaeus tropicalis: A, 3 17.5 mm cl, east of Cayos de Albulquerque, Colombia. Petasma,

dorsal view. (From PfiREZ Farfante, 1977). — B, $ 25 mm cl, east of Cayos de Albulquerque, Colombia.

Thelycum. Scales = 1 mm. (From P£rez Farfante, 1977).

Distribution of branchiae, epipods, and exopods in the genus Mesopenaeus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

176

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Petasma semi-open, with ventral costa free or attached to adjacent membranous part and

bearing terminal flap. Appendix masculina and appendix interna present; basis of pleopod

produced distally into elongate ventrolateral spur. Thelycum open, with paired anterior

protuberances, more or less elevated, on stemite XIV.

Descriptions and illustrations of the complex spermatophores of M. tropicalis and M. mariae

are provided by Perez Farfante (1977a) and Perez Farfante and Ivanov (1982), respectively.

Species. — Mesopenaeus brucei Crosnier, 1986b. Japan; Queensland and north-western

Australia.

Mesopenaeus mariae Perez Farfante and Ivanov, 1982. Southern Mozambique; off Saya de

Malha Bank, Indian Ocean; Taiwan; Japan; Wallis and Futuna Islands.

Mesopenaeus tropicalis (Bouvier, 1905a). Western Atlantic Ocean from North Carolina to

southern Brazil; Gulf of Mexico.

Genus Pleoticus Bate, 1888

Figures 122-123

Hymenopenaeus - Smith, 1885b, Proc. U. S. natn. Mus., 8(11/12): 179 [part]. — Burkenroad, 1936b, Bull.

Bingham oceanogr. Coll., 5(2): 102 [part].

Philonicus Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 273 [part, excluding Philonicus lucasii Bate,

1881 = Hadropenaeus lucasii , and Philonicus pectinatus Bate, 1888 = Solenocera pectinata ]. Preoccupied by

Philonicus Loew, 1849: 144 (Diptera).

Pleoticus Bate, 1888, Rep. scient. Results Voy. Challenger, 24: xii [part]. Replacement name for Philonicus Bate,

1888. — P£rez Farfante, 1977a, Fish. Bull., U. S., 75(2): 295. — Squires, 1990, Can. Bull. Fish. Aq. Sci.,

221: 36.

Fig. 122. — Pleoticus robustus, 6 31 mm cl, east of Peninsula Valiente, Panama. Lateral view. Scale = 10 mm.

(From P£rez Farfante, 1977).

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

177

Faxonia Bouvier, 1905a, C. r. hebd. Seanc. Acad. Sci., Paris, 140: 981 [part, excluding Faxonia diomedeae

(Faxon, 1893) = Haliporoides diomedeae (Faxon, 1893)]. Type species: By subsequent designation, by Fowler,

1912, Ann. Rep. N. J. State Mus. for 1911: 543, Penaeopsis [sic] ocularis Faxon, 1895 [= Pleoticus robustus

Smith, 1885b], Mem. Mus. comp. Zool. Harv., 18: 187.

Parartemesia Bouvier, 1905b, C. r. hebd. Seanc. Acad. Sci., Paris, 141: 747 [part, excluding Parartemesia

tropicalis Bouvier, 1905b = Mesopenaeus tropicalis (Bouvier, 1905b)]. Type species: By subsequent

designation by Fowler, 1912, Ann. Rep. N. J. State Mus. for 1911: 543, Parartemesia carinata Bouvier,

1905b [= Pleoticus muelleri (Bate, 1888)].

Haliporus - Bouvier, 1905b, C. r. hebd. Seanc. Acad. Sci., Paris, 141: 1 [part], — A. Milne Edwards and

Bouvier, 1909, Mem. Mus. comp. Zool. Harv., 27: 206 [part], — De Man, 1911, Siboga Exped., 39a: 31. —

Balss, 1914b, Zool. Anz. Akad. Wiss. Wien, 9: 7. — Crosnier, 1989b, Sencken. biol., 69(4/6): 380.

Type Species: (for both Philonicus and Pleoticus), by subsequent designation by Fowler,

1912, Ann. Rep. N. J. State Mus. for 1911: 543, Philonicus miilleri Bate, 1888.

Type Locality: Off Montevideo, Uruguay, 35°02’N, 55°15'W, 24 m.

Gender: Masculine.

Diagnosis. — Body robust, integument rather thick and firm. Rostrum moderately long,

reaching at least midlength of second antennular segment or at most slightly overreaching

peduncle; ventral margin straight or concave; armed with dorsal teeth only; epigastric tooth and

first rostral separated by interval equal to, or only slightly greater than that between first and

second rostral teeth. Carapace moderately long; orbital, postorbital, antennal, and hepatic spines

present; pterygostomian spine absent; branchiostegal spine present or absent. Cervical sulcus

long, ending close or rather close to middorsum of carapace; hepatic carina well marked

anteriorly, indistinct posteriorly; hepatic sulcus nearly horizontal posteriorly, to level of hepatic

spine, from there turning anteroventrally and reaching pterygostomian region; branchiocardiac

carina and sulcus present or lacking; submarginal carina sharp. Telson armed with pair of

conspicuous, fixed, subapical spines.

Prosartema long or moderately so, flexible; antennular flagella similar, subcylindrical, longer

than carapace. Mandibular palp triarticulate (not biarticulate as stated by Pfrf? Farfante, 1977a,

and illustrated by Balss, 1914), distal articles broad, distalmost as long, or almost as long as

middle one, and tapering to blunt apex. Palp of first maxilla entire, narrowing distally to apex,

and bearing proximal tuft of relatively long setae on lateral margin and produced into broad

proximomesial and distomesial lobules. Fifth moderately to considerably longer than fourth.

Basial and ischial spines on first pereopod. Exopods well developed. Lateral ramus of uropod

armed with terminal distolateral spine. One or 2 rudimentary arthrobranchiae on somite VII.

Distribution of branchiae, epipods, and exopods in the genus Pleoticus.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

xin

(P4)

XfV

(P5)

Pleurobranchiae

Arthrobranchiae

r+r

Podobranchiae

Epipods

Exopods

Petasma semi-open, with distal part of ventral costa distally free from flexible terminal part of

ventrolateral lobule. Appendix masculina and appendix interna present, basis of pleopod

178

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

produced distally into elongate ventrolateral spur. Thelycum open; anterior part of sternite XIV

conspicuously modified, bearing paired prominences or strong plates.

An illustrated account of the spermatophore of P. robustus is provided by Perez Farfante

(1977a).

Fig. 123. — Pleoticus robustus : A, 6 32 mm cl, east of Peninsula Valiente, Panama. Petasma, dorsolateral view

of right half. Scale = 2 mm. (From P£rez Farfante, 1977). — B, $ 49 mm cl, south of Curasao. Thelycum.

Scale = 2 mm. (From P£rez Farfante, 1977).

Species. — Pleoticus muelleri (Bate, 1888). Southern Brazil to southern Argentina.

Pleoticus robustus (Smith, 1885b). Western Atlantic Ocean from Massachusetts to Gulf of

Mexico; Caribbean to French Guiana.

Pleoticus steindachneri (Balss, 1914b). Red Sea; Gulf of Aden.

Genus Solenocera Lucas, 1849

Figures 124-125

Penaeus - Fabricius, 1798, Suppl. Ent. Syst., : 408 [part].

Peneus - Risso, 1816, Hist. Nat. Crust. Nice, : 98 [part].

Solenocera Lucas, 1849a, Rev. Mag. Zool., (2)1: 159 [nomen nudum].

Solenocera Lucas, 1849b, Rev. Mag. Zool., (2)1: 300; 1850, Annls Soc. ent. Fr., (2)8: 219. — Alcock, 1901,

Descr. Cat. Indian Deep-Sea Crust., : 19. — Bouvier, 1908a, Result. Camp, scient. Prince Albert I, 33: 86.

— De Man, 1911, Siboga Exped., 39a: 45. — Burkenroad, 1936b, Bull. Bingham oceanogr. Coll., 5(2):

120. — Kubo, 1949, J. Tokyo Coll. Fish., 36(1): 223. — Zariquiey Alvarez, 1968, Investnes Pesq., 32: 49.

Source: MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

179

— P£rez Farfante and Bullis, 1973, Smithson. Contrib. Zool., 153: 2. — Crosnier, 1978, Faune

Madagascar, 46: 129. — Burkenroad, 1983, Crust. Issues, 1: 281. — De Freitas, 1985, Investl. Rep.

oceanogr. Res. Inst., Durban, 57: 54. — Liu and Zhong, 1988, Pen. Shr. S. China Sea, : 84 — Ivanov

1994, Arthrop. Sel., 3: 13.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1961, Opinion 611, Name No. 1444, Bull. zool. Norn., 18: 306.

Philonicus - Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 273 [part, excluding Philonicus mulleri Bate,

1888 = Pleoticus muelleri (Bate, 1888), and Philonicus lucasii Bate, 1881 = Hadropenaeus lucasii (Bate

1881)].

Pleoticus - Bate, 1888, Rep. scient. Results Voy. Challenger, 24: xii [part]. Replacement name for Philonicus

Bate.

Parasolenocera Wood-Mason, 1891, Ann. Mag. nat. Hist., (6): 276. Type Species: By monotypy, Parasolenocera

annectens Wood-Mason, 1891. Type locality: Andaman Sea, 11°25'5"N, 92°47'6"E, 652 m. Gender: Feminine.

Transolenocera Burkenroad, 1934b, Bull. Am. Mus. Nat. Hist., 68(2): 69.

Fig. 124. — Solenocera membranacea, 9 34 mm cl, southeast Atlantic Ocean off the Congo. Lateral view.

Scale = 10 mm.

Type Species: By monotypy, Solenocera philippii Lucas, 1849a, [= Penaeus siphonoceros

Philippi, 1840], [= Penaeus membranaceus Risso, 1816 = Solenocera membranacea (Risso,

1816), Ann. Soc. ent. Fr., 2: 8)].

Type Locality: Bay of Alger near Fort Bab-Azzoun, and that of Philippi's specimens; type

locality not restricted.

Gender: Feminine.

Diagnosis. — Body moderately slender, integument glabrous or sparsely setose on carapace,

and with dorsal patch of setae on rostrum and anterior part of carapace. Rostrum short, not

overreaching second antennular segment, strongly compressed laterally, deep; armed with dorsal

Source:

180

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

teeth only; epigastric and first rostral teeth variably separated. Carapace with orbital spine present

or absent; armed with postorbital, antennal and hepatic spines; postantennal spine lacking;

pterygostomian or branchiostegal spines present or absent. Cervical sulcus strongly marked,

long, reaching, or almost reaching, middorsum of carapace; hepatic sulcus deep, nearly

horizontal posterior to level of hepatic spine, from there turning anteroventrally and reaching

pterygostomian region; hepatic carina sharp anteriorly; branchiocardiac carina and sulcus present

or absent; posthepatic carina absent. Telson usually with pair of fixed posterolateral spines,

lacking movable spines.

Antennule with prosartema well developed, variable in length; antennular flagella longer than

carapace, lamellate, broad, ventral pair forming trough covered by narrower dorsal flagellum,

four together constituting respiratory siphon. Mandibular palp triarticulate, distal articles

subtriangular with contiguous bases. Palp of first maxilla entire, rather abruptly narrowing to

rounded apex, bearing setose proximolateral lobule (setae extending near to or beyond apical tuft)

and broad, setose proximomesial and small distomesial lobules, latter armed with 1-3 long,

spinous setae. Basial and ischial spines on first pereopod; ischial spine usually on second

pereopod. Exopods well developed, on all maxillipeds and pereopods. Lateral ramus of uropod

almost always lacking distolateral spine. Two arthrobranchiae on somite VII, both small or

anterior one rudimentary, consisting of small lamella lacking filaments.

Petasma semi-open, with dorsolateral lobule usually bearing terminal process and ventral costa

distally free from contiguous part of ventrolateral lobule. Appendix masculina and appendix

interna present, basis of pleopod produced distally into elongate distolateral spur.Thelycum open,

usually with conspicuous protuberances on anterior part of stemite XIV.

Distribution of branchiae, epipods, and exopods in the genus Solenocera.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

xrn

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

S or r+s

Podobranchiae

Epipods

Exopods

Species. — Solenocera acuminata Perez Farfante and Bullis, 1973. Bahamas; Caribbean Sea

to French Guiana.

Solenocera africana Stebbing, 1917. Eastern Atlantic Ocean from Morocco to Agulhas Bank,

South Africa; off Natal, South Africa.

Solenocera agassizii Faxon, 1893. Cabo Blanco, Costa Rica to northern Peru.

Solenocera alfonso Perez Farfante, 1981, forma typica Crosnier, 1989a. Philippines.

Solenocera alfonso Perez Farfante, 1981, forma inermis Crosnier, 1989a. Indonesia;

Philippines.

Solenocera algoensis Barnard, 1947. East coast of Africa from Algoa Bay, South Africa, to

Zanzibar; Madagascar.

Solenocera alticarinata Kubo, 1949. South China Sea; Taiwan; Hong Kong; Japan.

Solenocera annectens (Wood-Mason, 1891). Andaman Sea; Indonesia; Philippines.

Solenocera atlantidis Burkenroad, 1939. North Carolina to Florida, Gulf of Mexico; West

Indies; Caribbean coast of Central and South America to Brazil.

Source: MNHN. Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

181

Fig. 125. —Solenocera membranacea : A, 3 13.5 mm cl, off Sicily, Mediterranean. Peiasma, dorsal view

Scale = 1 mm. — B, 2 34 mm cl, southeast Atlantic Ocean off the Congo. Thelycum. Scale = 10 mm.

Solenocera australiana Perez Farfante and Grey, 1980. Northern Australia.

Solenocera barunajaya Crosnier, 1994c. Indonesia; Timor Sea off Northern Australia.

Solenocera bedokensis Hall, 1962. Sri Lanka; Singapore; Gulf of Tonkin; South China Sea;

Philippines; Hong Kong.

Solenocera choprai Nataraj, 1945. South-east coast of South Africa; Mozambique;

Madagascar; Gulf of Suez; Red Sea; Arabian Sea; Andaman Sea; Straits of Malacca; Indonesia;

South China Sea; Philippines; Taiwan; Western Australia and New South Wales, Australia.

Solenocera comata Stebbing, 1915. South West Indian Ocean off South Africa; Madagascar;

Reunion; Indonesia; Philippines; South China Sea; Japan.

Solenocera crassicornis (H. Milne Edwards, 1837). Persian Gulf; Pakistan; India; Burma;

Singapore; Malaysia; Gulf of Tonkin; North Borneo; South China Sea; Hong Kong; Taiwan;

Japan.

Solenocera faxoni De Man, 1907. Indonesia; South China Sea; Japan; New South Wales,

Australia.

Solenocera florea Burkenroad, 1938. West coast of Baja California, Gulf of California;

Mexico to northern Peru.

Solenocera geijskesi Holthuis, 1959. Western Caribbean; Surinam; French Guiana; northern

Brazil.

Solenocera gurjanovae Starobogatov, 1972. Gulf of Tonkin.

Solenocera halli Starobogatov, 1972. Singapore; Straits of Malacca; Gulf of Tonkin.

Solenocera hextii Wood-Mason and Alcock, 1891. Gulf of Aden; Arabian Sea; Bay of

Bengal.

182

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Solenocera koelbeli De Man, 1911. Arabian Sea; Indonesia; Philippines; Gulf of Tonkin;

South China Sea; Hong Kong; Taiwan; Japan; Korea.

Solenocera mascarensis Burukovsky, 1993. Saya de Malha Bank.

Solenocera melantho De Man, 1907. Indonesia; Philippines; Taiwan; Japan; Korea.

Solenocera membranacea (Risso, 1816). North-east Atlantic Ocean from Ireland to Morocco;

Azores; Mediterranean; Senegal; south-east Atlantic Ocean off Cape of Good Hope, Agulhas

Bank.

Solenocera moosai Crosnier, 1985. Indonesia; Philippines.

Solenocera mutator Burkenroad, 1938. Southern California, U.S.A.; West coast of Baja

California, Gulf of California, Mexico to Isla Lobos de Tierra, northern Peru.

Solenocera necopina Burkenroad, 1939. North Carolina to Florida; Gulf of Mexico; Bahamas;

Caribbean to Uruguay.

Solenocera pectinata (Bate, 1888). East African coast off Tanzania and Kenya; Madagascar;

Arabian Sea; Bay of Bengal; Burma; Malaysia; Indonesia; Gulf of Tonkin; South China Sea;

Philippines; Taiwan; Japan; Wallis and Futuna Islands.

Solenocera pectinulata Kubo, 1949. Off Kenya; Madagascar; Mauritius; India; Indonesia;

South China Sea; Philippines; Japan.

Solenocera phuongi Starobogatov, 1972. Indonesia; Gulf of Tonkin.

Solenocera rathbunae Ramadan, 1938. Gulf of Aden; Zanzibar; Madagascar; South China Sea;

Philippines; New South Wales, Australia; Wallis and Futuna Islands; Hawaii.

Solenocera spinajugo Hall, 1961. Straits of Malacca; Philippines.

Solenocera vioscai Burkenroad, 1934b. North Carolina to Florida and Gulf of Mexico.

Solenocera waltairensis George and Muthu, 1970. East coast of Africa off Kenya;

Madagascar; India; Sri Lanka; Philippines.

Solenocera zarenkovi Starobogatov, 1972. Gulf of Tonkin.

Superfamily SERGESTOIDEA Dana, 1852

Sergestoidea Dana, 1852a, Proc. Acad. nat. Sci. Philad., 6: 18 [as Sergestidae, name translated to superfamilial

status by Holthuis in Glaessner, 1969, Treatise Invert. Paleo., R. Arthropoda 4(2): R446, R450].

— Burkenroad, 1983, Crust. Issues, 1: 280, 281, 284. — Williams, 1984, Shr. Lob. Crabs Atl. Coast

U.S., : 50.

Diagnosis. — Carapace moderately to extremely compressed. Rostrum shorter than

eyestalk. Antennule with ventral flagellum modified or absent. Pereopods 4 and 5 reduced or

absent (except in Sicyonella). Pleurobranchia absent. Never more than 2 branchiae per thoracic

somite on each side. No more than 7 or 8 well developed branchiae on each side.

Key to the Families of the Superfamily SERGESTOIDEA

— Branchiae absent; body strongly compresssed .LUCIFERIDAE

— Branchiae present; body moderately compressed . SERGESTIDAE

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

183

Family LUCIFERIDAE De Haan, 1849

Luciferidae De Haan, 1849, Fauna Jap., Crust., : 242. — Dana, 1850, Am. J. Sci. Arts, (2)9: 5; 1852, U. S.

Expl. Exped., 13(1): 639; 1853, Class. Geog. Distrib. Crust., : 1435. — Crosnier and Forest, 1973, Faune

Trop., 19: 345. — Burkenroad, 1983, Crust. Issues, 1: 281, 283. — Williams, 1984, Shr. Lob. Crabs All.

Coast U.S., : 52. — Squires, 1990, Can. Bull. Fish. Aq. Sci., 221: 57.

Placed on the Official List of Family-Group Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Bull. zool. Norn., 25(4/5): 141. [In the Official List, the author of the

name Luciferidae is incorrectly given as Dana, 1850. The correct author of Luciferidae is De Haan, 1849],

Luciferinae - Bate, 1888, Rep. scient. Res. Voy. Challenger, 24: 443. — Ortmann, 1893, Ergebn. Plankton-

Exped. Humboldt-Stift., 2: 29. — Hansen, 1922, Result. Camp, scient. Prince Albert I, 64: 10, 198. —

Cecchini, 1933, Mem. R. Com. Talas. Ital., 200: 12. — Burkenroad, 1934b, Bull. Am. Mus. nat. Hist., 68:

132. — Gordon, 1956, Scient. Rep. Gt. Barrier Reef Exped., 6(5): 324.

Leuciferinae - Ortmann, 1898, Bronn's Kl. Ordn. Tierreichs, (5)2: 1121.

Leuciferidae - Barnard, 1950, Ann. Mag. nat. Hist., 38: 580, 644. — Balss, 1957, Bronn's Kl. Ordn. Tierreichs

5(1)7(12): 1521.

Diagnosis. — Rostrum short, acute. Carapace extremely laterally compressed, anteriorly

elongate, with mandibles widely separated from antennae and eyes. Sixth abdominal somite in

male bearing 2 ventral processes. Telson in male with strong protuberance on ventral surface.

Antennules lacking ventral flagellum in both sexes. Mandible lacking palp. Maxillae lacking

palp; with exopod in form of small plate. First maxilliped lacking epipod and exopod. Second

maxilliped lacking epipod. Chelae lacking, or imperfect chela having no fixed finger present only

on third pereopod. Fourth and fifth pereopods absent. Branchiae absent. Genital aperture single

in both sexes.

Petasma sessile, attached proximally to first pleopodal peduncle. Second pleopod in male with

unilamellate appendix masculina.

Genus Lucifer Thompson, 1829

Figures 126-127

Lucifer Thompson, 1829, Zool. Res., 3: 58, pi. 7, fig. 2. — Dana, 1850, Am. J. Sci. Arts, (2)9: 5. — Bate,

1888, Rep. scient. Results Voy. Challenger, 24: 443. — Hansen, 1919, Siboga Exped., 38: 48; 1922, Result.

Camp, scient. Prince Albert 1, 54: 198. — Bowman and Holthuis, 1968, Crustaceana, 14(2): 216. — Omori.

1977, Proc. Symp. Warm Water Zoopl. Spl. Publ. UNESCO/NIO, 1977: 2. — Burkenroad, 1983, Crust.

Issues, 1: 283. — Squires, 1990, Can. Bull. Fish. Aq. Sci., 221: 57.

Placed on Official List of Generic Names in Zoology, International Commission on Zoological Nomenclature,

1969, Opinion 864, Name No. 1817, Bull. zool. Norn., 25(4/5): 139.

Leucifer H. Milne Edwards, 1837, Hist. Nat. Crust., : 467. — Ortmann, 1898, Bronn's Kl. Ordn. Tierreichs, (5)2:

1122. [Dr. L. Holthuis (in litt., 1996) notes that the name Leucifer H. Milne Edwards is an incorrect

subsequent spelling of Lucifer , and is not an emendation, as nowhere in Milne Edwards' publication is the

spelling Lucifer cited, and no choice between the two names is mentioned. Leucifer therefore has no standing

and is unavailable.]

Type Species: By subsequent indication under Article 68b, International Code of Zoological

Nomenclature, 3rd ed., by H. Milne Edwards, 1837, Leucifer typus H. Milne Edwards, 1837,

Hist. Nat. Crust., : 469.

Type Locality: Atlantic Ocean.

Gender: Masculine.

Source:

184

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Remarks. — The position and relationships

of the Luciferidae are problematic, as was

pointed out by Burkenroad (1983: 284).

Although the latter included the group in the

Dendrobranchiata, they possess none of the

features that he used to characterize this

suborder, other than that they hatch as free

nauplii (generally agreed to be a primitive

feature; see Kaestner, 1970: 53). Traditionally

regarded as a subgroup of the Sergestoidea, the

placement of Lucifer within the Penaeidea was

based on the presence of a petasma in the male

(a feature also found in the eumalacostracan

order Euphausiacea), while the absence of

pereopods 4 and 5 was seen as indicating its

relationship to the sergestids: Acetes, for

example, also lacks pereopods 4 and 5. The

assumption implicit here is that loss of

pereopods 4 and 5 is a synapomorphic feature of

Fig. 127. — Lucifer typus, 6 , north-east Atlantic the Luciferidae and the Sergestidae, rather than

Ocean off Liberia. Petasma. Scale = 2 mm. two events that occurred independently.

However, the advanced features that characterize

the Luciferidae, namely the extreme compression of the body, the lack of chelae, the absence of a

Source . MNHN, Paris

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

185

ventral antennular flagellum, the absence of mandibular and maxillar palps, the retension of eggs

on the third pleopods of the female, and the absence of gills, suggest that loss of the two

posterior pereopods was part of the suite of extreme modifications for a highly specialized

planktonic existence. Further, the complete lack of gills in Lucifer negates the most basic feature

for inclusion in the Dendrobranchiata.

Species. — Lucifer chacei Bowman, 1967. East coast of South Africa; Madagascar; Indonesia

to Hawaii; Tahiti.

Lucifer faxoni Borradaile, 1915. Western Atlantic Ocean from Long Island Sound to Rio de

laneiro; Gulf of Mexico; Caribbean Sea; Bermuda; eastern Atlantic Ocean off Senegal and

Congo.

Lucifer hanseni Nobili, 1905. Madagascar; Red Sea; India; South China Sea; Victoria,

Australia.

Lucifer intennedius Hansen, 1919. Gulf of Oman; Indonesia; Malacca Straits to lapan.

Lucifer orientalis Hansen, 1919. East coast of South Africa; Red Sea; Indonesia; Malaysia to

South China Sea; Philippines; eastern Central Pacific Ocean.

Lucifer penicillifer Hansen, 1919. South-east coast of South Africa; east African coast; Bay of

Bengal; Malaysia; Indonesia; South China Sea; Philippines; Hong Kong; Japan; northern

Australia.

Lucifer typus H. Milne Edwards, 1837. North-east Pacific Ocean, off Baja California and

Gulf of California to north of 4 N; off New Foundland; North-west Atlantic Ocean off U.S.A.;

Sargasso Sea; Brazil; north-east Atlantic Ocean; Mediterranean; south-east Atlantic Ocean off

Cape of Good Hope; east coast of South Africa; Bay of Bengal; Philippines; Queensland,

Australia; eastern Central Pacific Ocean.

Family SERGESTIDAE Dana, 1852

Sergestidae Dana, 1852a, Proc. Acad. nat. Sci. Philad., 6: 18; 1852b, Am. J. Sci. Arts, (2)14: 123; 1853, Class.

Geog. Distrib. Crust., : 1434. — Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 345. — Ortmann,

1898, Bronn's Kl. Ordn. Tierreichs, (5)2: 1121. — Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 47.

— Senna, 1903, Boll. Soc. ent. ital., 34: 280. — Hansen, 1922, Result. Camp, scient. Prince Albert I. 64: 9.

— Cecchini, 1933, Mems. R. Com. Talas. Ital., 200: 4. — Barnard, 1950, Ann. S. Afr. Mus., 38: 637. —

Balss, 1957, Bronn's Kl. Ordn. Tierreichs, 5(1)7(12): 1520. — Crosnier and Forest, 1973, Faune Trop., 19:

305. — Omori, 1975, Bull. Oc. Res. Inst. Univ. Tokyo, 7: 5. — Williams, 1984, Shr. Lob. Crabs Atl. Coast

U.S., 50. — Bauer, 1986, J. Crust. Biol., 6(3): 318. — Squires, 1990, Can. Bull. Fish. Aq. Sci., 221: 46.

Placed on the Official List of Family-Group Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Bull. zool. Nom., 25(4/5): 141.

Sergestinae - Bate, 1888, Rep. scient. Results Voy. Challenger, 24: 345. — Ortmann, 1893, Ergebn. Plankton-

Exped. Humboldt-Stif., 2: 29; 1898, Bronn's Kl. Ordn. Tierreichs, (5)2: 1121. — Hansen, 1922, Result.

Camp, scient. Prince Albert I, 64: 10. — Balss, 1957, Bronn's Kl. Ordn. Tierreichs, 5(1)7(12): 1520.

Diagnosis. — Integument thin, often very soft, bearing photophores in two genera. Rostrum

shorter than eyestalks, often small to rudimentary. Carapace moderately compressed; supraorbital

spine and hepatic spine present in some species of some genera; antennal, branchiostegal, and

pterygostomian spines absent; cervical sulcus well marked, weak, or absent. First to fifth

abdominal somites dorsally rounded, sixth somite weakly carinate. Telson with no more than

3 pairs, or lacking lateral movable spines.

186

ISABEL PfiREZ FARFANTE & BRIAN KENSLEY

Ventral antennular flagellum modified in male to form clasping organ. Antennal flagellum

bipartite, consisting of stiff proximal portion and more flexible distal portion. Mandibular palp of

3 articles. First maxilliped with exopod and epipod; second maxilliped with epipod; second and

third maxillipeds and all pereopods lacking exopods. First to third pereopods (in Peisos) or

second and third pereopods with minute chela (Sergestes pectinatus lacking chela on third pereo-

pod). Fourth and fifth pereopods reduced (except in Sicyonella) or absent. Branchiae present.

Petasma variously composed of lobus accessorius, lobus armatus, lobus connectens,

lobus inermis, lobus terminalis, processus ventralis, processus uncifer; lobes often bear¬

ing hooks. Appendix masculina unilamellate. Thelycum with sternite XII and sometimes sternite

XIII and coxae of third pereopod modified; seminal receptacles present, small, varying from

simple shallow pockets to sac-like invaginations situated submesially at base of third pereopods.

Remarks. — Of all the sergestid genera, Sergestes H. Milne Edwards, 1830, has had the

most checkered history. Stimpson (1860) applied the name Sergia to a late mastigopus sergestid

larva, and even though Ortmann (1893) recognized Sergia as a full genus, this was not generally

accepted. Even Yaldwyn's landmark 1957 paper on New Zealand sergestids, while recognizing

a fundamental difference between those species possessing organs of Pesta ( Sergestes ) and those

with dermal photophores (Sergia), gave these taxa subgeneric status within Sergestes. Omori

(1974) finally reintroduced Sergia as a full genus. It has long been recognized that both genera

can be divided into groups of closely-related species (e.g. Yaldwyn, 1957; Judkins, 1978), but

apart from the unpublished work of Judkins' (1972), this division has never been formally pro¬

posed. Unfortunately, as the latter work does not constitute a publication under articles 8 and 9 of

the International Code of Zoological Nomenclature (1985), the seven new genera proposed

therein (and based primarily on petasmatal features) cannot be used. Foxton (1972) demonstrated

several groupings of species of Sergestes s.l. based on the structure and arrangement of the or¬

gans of Pesta, but, again, no formal names for these groupings were proposed.

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

187

Key to the genera of the family SERGESTIDAE

1. Fourth and fifth pereopods absent. Acetes

— Fourth pereopod present, fifth pereopod present or absent. 2

2. Fourth pereopod of 7 podomeres. Sicyonella

— Fourth pereopod of fewer than 7 podomeres . 3

3. Fourth pereopod of 5 podomeres; fifth pereopod vestigial in male, of

3 segments in female . Peisos

— Fourth and fifth pereopods of 6 podomeres. 4

4. Branchiae absent or rudimentary above fourth pereopod . Petalidium

— Branchiae present above fourth pereopod . 5

5. Dermal photophores present, organ of Pesta absent; ischiopodite of first

and second pereopods lacking subterminal spine. Sergia

— Dermal photophores absent, organ of Pesta present; ischiopodite of first

and second pereopods with well developed subterminal spine .. Sergestes

Genus Acetes H. Milne Edwards, 1830

Figures 128-130

Acetes H. Milne Edwards, 1830, Annls Sci. nat., 19: 350. — Bate, 1881, Ann. Mag. nat. Hist., (5)8: 172, 195.

— Ortmann, 1893, Ergebn. Plankton-Exped. Humboldt-Stift., 2: 38: 1898, Bronn's Kl. Ordn. Tierreichs, (5)2:

1122. — Hansen, 1919, Siboga Exped., 38: 31. — Burkenroad, 1934b. Bull. Am. Mus. nat. Hist., 68: 126.

— Colefax, 1940, Rec. Austr. Mus., 20(5): 341. — Barnard, 1950, Ann. S. Afr. Mus., 38: 822. —

Kensley, 1971a, Ann. S. Afr. Mus., 57(10): 226. — Omori, 1975, Bull. Ocean Res. Inst. Univ. Tokyo, 7: 5;

1977, Proc. Symp. Warm Water Zoopl. Spl. Publ. UNESCO/NIO, : 4. — Burkenroad, 1983. Crust. Issues,

1: 283. — Xiao and Greenwood, 1993, Oceanogr. Mar. Biol. Annu. Rev., 31: 259-444.

Sergestes - Kroyer, 1859, K. Dansk. Vidensk. Selsk. Skr., (5)4: 268 [part].

Type Species: By original designation Acetes indicus H. Milne Edwards, 1830, Annls Sci.

nat., 19: 351, pi. 1, 2.

Diagnosis. — Rostrum short, acute, lacking, or with

1-2 dorsal teeth. Carapace with supraorbital and hepatic

spines present; almost devoid of sulci. Abdominal

somites dorsally rounded. Telson lacking lateral movable

spines, tapering to subacute apex, tip bearing or lacking

minute spines.

Ventral antennular flagellum in male modified, short,

having one or more strong basal spines, flagellar portion

of few articles; dorsal flagellum elongate, slender, terete.

First maxilla lacking palp; second maxilla with single

lobe. First maxilliped lacking palp. First to third

pereopods each with minute chela. Fourth and fifth

pereopods absent; fifth pereopod in male represented by

Type Locality: Ganges River.

Gender: Masculine.

Fig. 129. — Acetes indicus , S 4.8 mm

cl, Chao Praya River at Pak Nam,

Thailand. Antennule.

Source:

188

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

coxal protuberances.

Petasma with processus ventralis of varying length, triangular, apically acute; capitulum

bearing hooks; processus uncifer broad. Thelycum with subspherical thin-walled seminal

receptacles opening submedially between bases of third pereopods; coxa of third pereopod with

mesially directed flange.

Distribution of branchiae, epipods, and exopods in the genus Acetes.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

xrn

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

r(-)

Podobranchiae

r(-)

Epipods

Exopods

Fig. 130. — Acetes indicus: A, 6 4.8 mm cl, Chao Praya River at Pak Nam, Thailand. Petasma, dorsal view.

Scale = 0.5 mm. — B. $ 6.0 mm cl, Chao Praya River at Pak Nam, Thailand. Seminal receptacles, dorsal

view. Scale = 1 mm.

Species. — Acetes americanus americanus Ortmann, 1893. Puerto Rico; Panama; Surinam;

French Guiana; Brazil (Amazon estuary).

Acetes americanus carolinae Hansen, 1933. North Carolina; Florida to Gulf of Mexico;

Panama; Surinam; French Guiana.

Acetes binghami Burkenroad, 1934b. Pacific Panama to Gulf of Guayaquil, Ecuador.

Acetes chinensis Hansen, 1919. Japan; China; Korea.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

189

Acetes erythraeus Nobili, 1905. East coast of South Africa to Red Sea; Madagascar; Indo-

West Pacific Ocean to Hong Kong and Philippines; northern Australia.

Acetes indicus H. Milne Edwards, 1830. India; Indonesia to South China Sea.

Acetes intermedius Omori, 1975. Indonesia; Philippines; Taiwan.

Acetes japonicus Kishinouye, 1905. India to Japan; Straits of Malacca; Java, Indonesia;

China; Korea.

Acetes johni Nataraj, 1947. Southern India.

Acetes marinus Omori, 1975. Surinam; Brazil (brackish water).

Acetes natalensis Barnard, 1955. Natal, South Africa.

Acetes paraguayensis Hansen, 1919. Amazon River basin; Parana River basin; Paraguay

River basin.

Acetes serrulatus (Krpyer, 1855). Singapore; Java, Indonesia; Sarawak.

Acetes sibogae australis Colefax, 1940. East coast of Australia.

Acetes sibogae sibogae Hansen, 1919. South India; Straits of Malacca; Singapore; Java;

Borneo; Thailand; Philippines.

Acetes sibogae sibogalis Achuthankutty and George, 1973. Cochin, India.

Acetes vulgaris Hansen, 1919. Singapore; Java, Indonesia; Flores Sea; Thailand to South

China Sea.

Genus Peisos Burkenroad, 1945

Figures 131-133

Peisos Burkenroad, 1945, Trans. Conn. Acad. Arts Aci., 36: 553. — Mallo and Boschi, 1982, Physis, B. Aires,

A41(100): 85. — Burkenroad, 1983, Crust. Issues, 1: 283.

Fig. 131. — Peisos petrunkevitchi, 9 10 mm cl, Rawson, Chubut, Argentina. Lateral view. Scale = 5 mm.

190

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Type Species: By original designation, Peisos petrunkevitchi Burkenroad, 1945, Trans. Conn.

Acad. Arts Sci., 36: 554.

Type Locality: Montevideo, Uruguay, 9-11 m.

Gender: Masculine.

Diagnosis. — Rostrum short, triangular, anterodorsally directed and followed by single

postrostral tooth. Carapace with strong supraorbital and hepatic spines; cervical sulcus feeble,

only laterally visible; branchiocardiac carina well marked. Sixth abdominal somite weakly

carinate, bearing small middorsal spine on posterior margin. Telson lacking lateral movable

spines, posteriorly trispinose.

Ventral antennular flagellum of male S-shaped, lacking basal spines, with 4 short spines

on segment 4. Maxilla with narrow blade-like palp. First to third pereopods each with small

chela; fourth and fifth pereopods reduced; fifth pereopod about one-sixth length of fourth

pereopod.

Distribution of branchiae, epipods, and exopods in the genus Peisos.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

xni

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

Podobranchiae

Epipods

Exopods

Fig. 132. — Peisos petrunkevitchi, 6 11.2 mm cl, Rawson, Chubut, Argentina. Petasma, dorsal view

Scale = 1 mm.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

191

Petasma with processus ventralis elongate narrow, C-shaped, rounded apex directed laterally,

armed with 6 claws; dorsodistal lobe at base of processus ventralis triangular, directed laterally,

bearing terminal spine and smaller subterminal spines. Thelycum consisting of curved transverse

plate on sternite XII having lateral depression to accommodate coxal process of third pereopod;

saclike seminal receptacles opening mesially at base of third pereopod, consisting of small

rounded anteromesial lobe and more elongate posterior lobe.

Fig. 133. — Peisos petrunkevilclii:, 9 10 mm cl, Rawson, Chubul, Argentina: A, Sternitcs XI-X1V. B, Seminal

receptacles, dorsal view. Scales = 1 mm.

Species. — Peisos petrunkevitchi Burkenroad, 1945. Rio Grande, Brazil, to Chubut,

Argentina.

Genus Petalidium Bate, 1881

Figures 134-136

Sergestes - Kroyer, 1855, Overs. K. Dansk. Vidensk. Selsk. Forh. Copenhagen, 1855: 31, 34 [part],

Petalidium Bate, 1881, Ann. Mag. nat. Hist., (5)8: 172, 194. — Ortmann, 1898, Bronn's Kl. Ordn. Tierreichs,

(5)2: 1122. — Hansen, 1903, Proc. zool. Soc. Loud., 1903(1): 54; 1922, Rdsult. Camp, scient. Prince Albert

I, 54: 189. — Burkenroad, 1937a, Zoologica, N. Y., 22(25): 324. — Kensley, 1971b, Ann. S. Afr. Mus.,

57(10): 228. — Wasmer, 1974, Crustaceana, 27(2): 159. — Burkenroad, 1983, Crust. Issues, 1: 283.

Type Species: By monotypy, Petalidium foliaceum Bate, 1881, Ann. Mag. nat. Hist.,

Source:

192

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Type Locality: Off Marion Island, 46°46'S, 45°31'E, 2516 m; South of Australia, 47°25'S,

130°22'E, 3935 m.

Gender: Masculine.

Fig. 134. — Petalidium foliaceum , $ 18 mm cl. South Pacific Ocean. Lateral view. Scale = 5 mm.

Diagnosis. — Integument fairly soft and thin. Rostrum short, triangular, with one small

dorsal tooth, latter sometimes lacking. Carapace with supraorbital spine absent; minute hepatic

spine present, or with low hepatic prominence; cervical sulcus reaching midline, demarked

laterally by distinct carina; postantennal and branchiocardiac carinae present; hepatic carina short,

directed anteroventrally; anterior branchiostegal carina setose. Abdominal somites dorsally

rounded. Telson lacking lateral movable spines, posterior margin trispinose with posterolateral

spines much smaller than median spine, or lacking posterolateral spines.

Eye pigmented; eyestalk reaching anteriorly well beyond rostral apex, with small mesiodistal

tubercle. Ventral antennular flagellum in male with segment 3 of peduncle elongate, tipped with

spine, segment 4 concave, with distal protuberance; short spines on inner surface of segments 3

and 4; flagellum of 5 slender articles. First to third pereopods slender, elongate, second and third

Distribution of branchiae, epipods, and exopods in the genus Petalidium.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

XII

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

lam

1+lam

s+r

Podobranchiae

Epipods

Exopods

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

193

Fig. 135. — Petalidium foliaceum, 6 13 mm

cl, Antarctic Ocean. Petasma, dorsal view.

Scale = 1 mm.

pereopods with minute chela; fourth and fifth

pereopods reduced, fourth longer than fifth.

Branchial lamellae as well as branchiae present;

branchiae sometimes having fewer branches than in

Sergestes.

Petasma with processus ventralis bifurcate;

lobus armatus bilobed; lobus terminalis bifurcate,

slender; lobus inermis short; pars externa with

small oblique tooth distally. Thelycum barely

differentiated, with shallow seminal receptacles

situated laterally beneath rounded lateral flap at base

of third pereopod.

Species. — Petalidium foliaceum Bate, 1881.

South-east Atlantic Ocean off Cape of Good Hope,

South Africa; south-west Indian Ocean off Natal,

South Africa; southern oceans to Antarctica.

Petalidium obesum (Krpyer, 1855). North-east

Atlantic Ocean off Azores, Canary Islands, Cape

Verde Islands; south-east Atlantic Ocean off

Namibia and Cape of Good Hope, South Africa;

south-west Indian Ocean off Natal, South Africa.

Petalidium suspiriosum Burkenroad, 1937a.

North-east Pacific Ocean off Oregon, and Baja

California to Isla Clarion, Mexico; Hawaii; north¬

west Pacific Ocean.

Fig. 136. — Petalidium foliaceum, $ 18 mm cl. South Pacific Ocean. A, Thelycum. B, Seminal receptacles,

dorsal view. Scales = 1 mm.

Source:

194

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Genus Sergestes H. Milne Edwards, 1830

Figures 137-139

Nika - Risso, 1816, Hist. nat. Crust. Nice, : 87 [part].

Sergestes H. Milne Edwards, 1830, Annls Sci. nat. Zool., 19: 346. — Kroyer, 1855, Vidensk. Selsk. Skr., (5)4:

219-234. — Bate, 1881, Ann. Mag. nat. Hist., (5)8: 171. — Ortmann, 1893, Ergebn. Plankton-Exped.

Humboldt-Stift., 2: 29; 1898, Bronn's Kl. Ordn. Tierreichs, (5)2: 1121. — Hansen, 1896, Proc. zool. Soc.

Lond., 1896: 936-970 [part]; 1903, Proc. zool. Soc. Lond., 1903(1): 53; 1919, Siboga Exped., 38: 2; 1922,

Result. Camp, scient. Prince Albert I, 64: 11. — Alcock, 1901, Descr. Cat. Indian Deep-Sea Crust., : 48. — ■

Kemp, 1910b, Scient. Invest. Fish. Brch Ire., 1: 24. — Burkenroad, 1937a, Zoologica, N. Y., 22(25): 316.

— Barnard, 1950, Ann. S. Afr. Mus., 38: 638. — Yaldwyn, 1957, Zoology Pubis Viet. Univ. Coll., 22: 7.

— Milne, 1968, Crustaceana, 14(1): 22. — Kensley, 1971a, Ann. S. Afr. Mus., 57(10): 230. — Crosnier

and Forest, 1973, Faune Trop., 19: 305. — Burkenroad, 1983, Crust. Issues, 1: 283. — Squires, 1990,

Can. Bull. Fish. Aq. Sci., 221: 47.

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1824, Bull. zool. Nom., 25(4/5): 139.

Acheles Cocco, 1832, Eff. Scient. Lett. Sicilia, 2: 204. — Burkenroad, 1983, Crust. Issues, 1: 288. Type

species: By monotypy, Acheles arachnipodus Cocco, 1832, Eff. Scient. Lett. Sicilia, 2: 204 [= Sergestes

corniculum Krpyer, 1855]. Type locality: Messina, Sicily. [See Holthuis, 1977, for a discussion of this

species]. Gender: Feminine.

Fig. 137. — Sergestes atlanticus, $ 5.8 mm cl, Northwest Atlantic Ocean. Lateral view. Scale = 1 mm.

Type Species: By monotypy, Sergestes atlanticus H. Milne Edwards, 1830, Annls Sci. nat.

Zool., 19: 349, pi. 10, figs. 1-9.

Type Locality: North Atlantic Ocean.

Gender: Masculine.

Diagnosis. — Integument moderately firm, lacking dermal photophores. Red subcuticular

chromatophores mainly concentrated in anterior part of body. Rostrum short, barely extending

beyond anterior margin of carapace, apically acute or rounded, with or without single dorsal

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

195

tooth. Carapace with supraorbital and hepatic spines present or absent in adults. Cervical sulcus

well marked, reaching dorsal midline; postcervical sulcus, if present, reaching dorsal midline;

rounded branchiocardiac carina usually present; rounded hepatic carina sometimes present.

Gastrohepatic gland modified to form luminescent Organ of Pesta. Ovary in female confined to

cephalothorax. Abdominal somites dorsally rounded, or sixth somite weakly carinate. Telson

lacking lateral movable spines, tapering to subacute apex.

Fig. 138. —Sergestes atlanticus, 6 5.8 mm cl, Northwest Atlantic Ocean: A, Dorsal

view of chela of pereopod 2. B, Ventral view of chela of pereopod 2. C, Antennule.

Eye with cornea well pigmented, wider than eyestalk. Third maxilliped either subequal in

length to third pereopod, or markedly more robust and longer than third pereopod. First to third

pereopods elongate, slender, bearing stiff setae; first pereopod lacking chela; second and third

pereopods with minute chelae; fourth and fifth pereopods bearing natatory setae, consisting of

6 podomeres, dactyls lacking; fifth pereopod shorter than fourth pereopod, lacking gills.

Petasma with processus ventralis not forked. Thelycum with small sac-like seminal receptacles

situated submesially at base of third pereopod, often connected by shallow transverse groove.

Distribution of branchiae, epipods, and exopods in the genus Sergestes.

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

1+lam

Podobranchiae

Epipods

Exopods

Source .

196

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Fig. 139. — Sergestes atlanticus, S 6.0 mm cl, Northwest Atlantic Ocean. A, Petasma, dorsal view. B, Idem,

lateral view. Scale = 0.5 mm.

Species. — Sergestes arachnipodus (Cocco, 1832). Bermuda; Mediterranean.

Sergestes arcticus Kr 0 yer, 1855. Mediterranean; south-west Atlantic Ocean; south-east

Atlantic Ocean; south-west Indian Ocean; south-east Australia; New Zealand.

Sergestes armatus Kr 0 yer, 1855. Mediterranean; north Atlantic Ocean off Bermuda, Azores

and Canary Islands; south-east Atlantic Ocean; south-west Indian Ocean; north-west Pacific

Ocean.

Sergestes atlanticus H. Milne Edwards, 1830. Sargasso Sea; north-east Atlantic Ocean off

Bermuda, Azores and Canary Islands; Mediterranean; south-east Atlantic Ocean; south-west

Indian Ocean; eastern Australia; north-west Pacific Ocean.

Sergestes brevispinatus Judkins, 1978. Eastern Central Pacific Ocean.

Sergestes coalitus Burkenroad, 1940. Sulu Sea.

Sergestes consobrinus Milne, 1968. Northeastern Pacific Ocean off California, U.S.A. and

Baja California, Mexico; north-west Pacific Ocean.

Sergestes comutus Kr 0 yer, 1855. Off Bermuda.

Sergestes curvatus Crosnier and Forest, 1973. North-east and south-east Atlantic Ocean,

south-west Indian Ocean.

Sergestes diapontius Bate, 1881. North-east and south-east Atlantic Ocean; Gulf of Aden;

Seychelles, Chagos Archipelago; Sri Lanka; Bay of Bengal.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS 197

Sergestes disjunctus Burkenroad, 1940. South-east Atlantic Ocean; south-west Indian Ocean;

off New Zealand.

Sergestes edwardsi Kr0yer, 1855. Off Bermuda; north-west Atlantic Ocean; Madeira; Cape

Verde Islands; eastern Atlantic Ocean from Sierra Leone to Zaire.

Sergestes erectus Burkenroad, 1940. Off Baja California; off Hawaii; off Rarotonga Island,

south-central Pacific Ocean; north-west Pacific Ocean.

Sergestes extensus Hanamura, 1983. Off Baja California.

Sergestes geminus Judkins, 1978. Eastern Central Pacific Ocean.

Sergestes gibbolobatus Judkins, 1978. Central equatorial Pacific Ocean.

Sergestes halia Faxon, 1893. Southern Baja California and Gulf of California; Gulf of

Panama.

Sergestes hamifer Alcock and Anderson, 1894. Laccadive Islands.

Sergestes henseni (Ortmann, 1893). North and South Atlantic Ocean; Mediterranean.

Sergestes incertus Hansen, 1896. South-central Atlantic Ocean off Tristan da Cunha.

Sergestes index Burkenroad, 1940. Off New Zealand; north-west Pacific Ocean.

Sergestes nipponensis Yokoya, 1933. Japan.

Sergestes nudus Illig, 1914. North-eastern Atlantic Ocean from Madeira to Congo; Gulf of

Aden; Seychelles; Chagos Archipelago; Cocos Islands; Bay of Bengal.

Sergestes orientalis Hansen, 1919. South-east Atlantic Ocean; south-west Indian Ocean; Red

Sea; Indonesia; South China Sea; north-west Pacific Ocean.

Sergestes paraseminudus Crosnier and Forest, 1973. South-east Atlantic Ocean off Gabon,

Congo, and Angola.

Sergestes pectinatus Sund, 1920. Off Baja California; Bermuda; Sargasso Sea; North-east

Atlantic Ocean off Azores and Canary Islands; south-east Atlantic Ocean, south-west Indian

Ocean; north-west Pacific Ocean.

Sergestes pediformis Crosnier and Forest, 1973. South-east Atlantic Ocean off Gabon,

Congo, and Angola.

Sergestes pestafer Burkenroad, 1937a. Eastern Pacific Ocean from southern California and

Gulf of California to Galapagos Islands.

Sergestes rubroguttatus Wood-Mason, 1891. Mediterranean; Arabian Sea; Bay of Bengal;

Andaman Sea.

Sergestes sargassi Ortmann, 1893. Off Baja California; Sargasso Sea; Bermuda; North-east

Atlantic Ocean off Azores and Canary Islands; Mediterranean; south-east Atlantic Ocean, south¬

west Indian Ocean; north-west Pacific Ocean.

Sergestes seminudus Hansen, 1919. Arabian Sea; Indonesia; eastern Australia; Japan; north¬

west Pacific Ocean.

Sergestes semissis Burkenroad, 1940. Northern Mozambique; Arabian Sea; Bay of Bengal.

Sergestes similis Hansen, 1903. North-east Pacific Ocean, Bering Sea; Baja California Sur

and Gulf of California; south Pacific Ocean off Chile; eastern South Atlantic Ocean in Benguela

Current.

Sergestes stimulator Burkenroad, 1940. Off New South Wales, Australia.

Sergestes tantillus Burkenroad, 1940. Off Galapagos Islands.

Sergestes verpus Burkenroad, 1940. Celebes Sea.

Sergestes vigilax Stimpson, 1860. Off Bermuda; Mediterranean; north-west Pacific Ocean.

Source:

198

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Genus Sergia Stimpson, 1860

Figures 140-141

Sergestes - Kroyer, 1859, Vidensk. Selsk. Skr., (5)4: 219-234 [part]. — Hansen, 1896, Proc. zool. Soc. Lond.,

1896: 936 [part]; 1919, Siboga Exped., 38: 2 [part]; 1922, Result. Camp, scient. Prince Albert I, 64: 11

[part]. — Barnard, 1950, Ann. S. Afr. Mus., 38: 638 [part],

Sergia Stimpson, 1860, Proc. Acad. nat. Sci. Philad., 1860: 46. — Ortmann, 1893, Ergebn. Plankton-Exped.

Humboldt-Stift., 2: 37. — Kemp, 1910b, Scient. Invest. Fish. Brch Ire., 1: 24. — Burkenroad, 1983, Crust.

Issues, 1: 283, 288. — Squires, 1990, Can. Bull. Fish. Aq. Sci., 221: 52. — Vereshchaka, 1994,

Steenstrupia, 20(3): 73-95.

Sergestes (Sergia ) - Yaldwyn, 1957, Zoology Pubis Viet. Univ. Coll., 22: 8. — Kensley, 1971a, Ann. S. Afr.

Mus., 57(10): 230. — Crosnier & Forest, 1973, Faune Trop., 19: 306.

Fig. 140. — Sergia robusta, 2 12.5 mm cl, North Atlantic Ocean. Lateral view. Scale = 10 mm.

Type Species: By monotypy, Sergia remipes Stimpson, 1860, Proc. Acad. nat. Sci. Philad.,

1860: 46.

Type Locality: Pacific Ocean, 27.5°N, 138.5°E. [The type species, Sergia remipes, was

determined to be a late mastigopus stage by Burkenroad (1945: 587, footnote). The latter author

identified this mastigopus with the group of species in what was then the subgenus Sergia (see

Yaldwyn, 1957: 8)].

Gender: Feminine.

Diagnosis. — Integument very soft and thin to fairly firm; lacking or bearing photophores

with or without cuticular lenses; red cuticular pigment distributed over entire body and

appendages. Rostrum variable, short, never reaching cornea, apically acute or rounded, with or

without single dorsal tooth. Carapace lacking supraorbital and hepatic spines in adult; secondary

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

199

North Atlantic Ocean. Petasma, dorsal view.

Scale = 1 mm.

base of third pereopods, sometimes connected 1

hepatic prominences sometimes developed.

Cervical sulcus usually well defined, reaching

midline; postcervical sulcus often well defined,

reaching midline; rounded branchiocardiac

carina usually well marked; rounded hepatic-

antennal carina sometimes present.

Gastrohepatic gland not modified to form

luminescent organ. Ovary in female sometimes

extending into abdomen. Abdominal somites

dorsally rounded. Telson lacking lateral

movable spines, tapering to subacute apex.

Eye with cornea well pigmented, wider than

eyestalk. First to third pereopods elongate,

slender, bearing stiff setae; first pereopod

lacking proper chela; second and third

pereopods with minute chela; fourth and fifth

pereopods of 6 segments bearing natatory

setae, dactyl lacking; fifth pereopod shorter

than fourth pereopod. Usually 7 branchiae per

side, 8 in S. grandis, latter having podo-

branchia on second maxilliped.

Petasma with processus ventralis not

forked. Thelycum having small, shallow

seminal receptacles situated submesially at

shallow groove across stemite XII.

Distribution of branchiae, epipods, and exopods in the genus Sergia

Somite

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

XIV

(P5)

Pleurobranchiae

Arthrobranchiae

1+r

Podobranchiae

Epipods

Exopods

Species. — Sergia bigemmea (Burkenroad, 1940). Baja California; north-west Pacific Ocean;

off Tahiti; Hawaii.

Sergia bisulcata (Wood-Mason, 1891). Arabian Sea; Bay of Bengal; Indonesia; Philippines.

Sergia challenged (Hansen, 1903). Seychelles; Indonesia; Fiji.

Sergia colosii (Cecchini, 1933). Red Sea.

Sergia extenuata (Burkenroad, 1940). Eastern Atlantic Ocean from Ivory Coast, Gabon, and

Angola.

Sergia fdicta (Burkenroad, 1940). Baja California; Gulf of Panama; north-west Pacific Ocean.

Sergia fulgens (Hansen, 1919). Indonesia.

Sergia gardineri (Kemp, 1913). Natal, South Africa; Seychelles; Indonesia; central Pacific

Ocean; north-west Pacific Ocean.

Source:

200

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Sergia grandis (Sund, 1920). North-west Atlantic Ocean from Grand Banks, Bermuda,

Sargasso Sea, Surinam; North-east Atlantic Ocean to Cape Verde Islands; Azores; south-east

Atlantic Ocean; southern Indian Ocean.

Sergia hansjacobi Vereshchaka, 1994. North Atlantic: Caribbean region and South America.

Sergia inequalis (Burkenroad, 1940). South-west Indian Ocean off Natal, South Africa; Java

Sea; Japan; north-west Pacific Ocean.

Sergia inoa (Faxon, 1893). Indonesia; north-west Pacific Ocean.

Sergia japonica (Bate, 1881). Off Bermuda; Seychelles; Sri Lanka; Philippines; Japan; north¬

west Pacific Ocean; eastern Australia; New Zealand.

Sergia laminata (Burkenroad, 1940). Southern California, U.S.A.; Baja California; south-east

Atlantic Ocean; south-west Indian Ocean; Madagascar; north-west Pacific Ocean.

Sergia lucens (Hansen, 1922). Taiwan; Japan.

Sergia maxima (Burkenroad, 1940). Baja California; northern Madagascar; Hawaii; New

Zealand; north-west Pacific Ocean.

Sergia plumea (Illig, 1927). North-west Indian Ocean off Somalia.

Sergia potens (Burkenroad, 1940). South-east Atlantic Ocean; south-west Atlantic Ocean off

Brazil and Argentina; south-west Indian Ocean; New Zealand.

Sergia prehensilis (Bate, 1881). South-east Atlantic Ocean off South Africa; Antarctic; south¬

west Indian Ocean; Japan; eastern Australia; north-west Pacific Ocean.

Sergia profunda (Bate, 1888). South-east Atlantic Ocean; south-east Pacific Ocean off

Valparaiso.

Sergia regalis (Gordon, 1939). South-east Atlantic Ocean; south-west Indian Ocean; Japan;

north-west Pacific Ocean.

Sergia robusta (Smith, 1882). Gulf of Mexico; Caribbean Sea; Surinam; North Sea; Faroe

Islands; off Bermuda; Mediterranean; Cape Verde Islands to Congo and Angola.

Sergia scintillans (Burkenroad, 1940). Baja California; south-east Atlantic Ocean; south-west

Indian Ocean; Sumatra; Hawaii; north-west Pacific Ocean.

Sergia splendens (Sund, 1920). Off Bermuda; Mediterranean; north-east and south-east

Atlantic Ocean.

Sergia stellata (Burkenroad, 1940). South of Sri Lanka, Indian Ocean.

Sergia talismani (Barnard, 1947). Off Bermuda; north-east Atlantic Ocean from Cape Verde

Islands to Portuguese Guinea, Gabon, Congo, Cabinda; south-west Indian Ocean; Japan.

Sergia tenuiremis (Krpyer, 1855). Off Bermuda; north-west Pacific Ocean.

Sergia wolffi Vereshchaka, 1994.Caribbean and Northeastern Atlantic.

Genus Sicyonella Borradaile, 1910

Figures 142-143

Aphareus Paulson, 1875, [Stud. Crust. Red Sea, 1: 123]. [Invalid junior homonym of Aphareus Cuvier and

Valenciennes, 1830, Hist. Nat. Poissons, 6: 485]. Type species: By monotypy, Aphareus inermis Paulson,

1875, [Stud. Crust. Red Sea, 1: 123]. Type locality: Red Sea. Gender: Masculine.

Sicyonella Borradaile, 1910, Trans. Linn. Soc. Lond., (2) zool., 13(2): 259. — Hansen, 1919, Siboga Exped., 38:

27. — Burkenroad, 1937b, Ann. Mag. nat. Hist., (10)19: 505; 1983, Crust. Issues, 1: 283.

Source:

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

201

Placed on the Official List of Generic Names in Zoology, International Commission on Zoological

Nomenclature, 1969, Opinion 864, Name No. 1825, Bull. zool. Norn., 25(4/5): 139.

Aphareocaris Caiman, 1913, J. Linn. Soc. Lond., zool., 32: 219. Type species: Aphareus inermis Paulson, 1875,

Izs. Rak. Kras. Mor. Chast. Kiev, 1: 117, by designation. [Name to replace Aphareus Paulson, 1875,

preoccupied]. Type locality: Red Sea. Gender: Masculine.

Fig. 142. — Sicyonella maldivensis, 9 8 mm cl, Bikini Atoll, Marshall Islands. Lateral view. Scale = 5 mm.

Type Species: By monotypy, Sicyonella maldivensis Borradaile, 1910, Trans. Linn. Soc.

Lond., (2) zool. 13(2): 259.

Type Locality: Maidive Islands, Indian Ocean.

Gender: Feminine.

Diagnosis. — Integument moderately firm. Rostrum short, triangular, apically acute, with

2 dorsal teeth. Carapace with strong supraorbital and hepatic spines; cervical sulcus poorly

defined, only laterally visible; rounded branchiocardiac carina well marked. Abdominal somites

dorsally rounded. Telson with 4 pairs of lateral movable spines; apically spiniform. acute.

Eye with cornea with well developed pigment, wider than relatively elongate eyestalk. Third

maxilliped more robust and longer than pereopods, armed with strong spines on posterior margin

Distribution of branchiae, epipods, and exopods in the genus Sicyonella

Somile

(Mxpl)

(Mxp2)

(Mxp3)

(PI)

(P2)

(P3)

(P4)

xrv

(P5)

Pleurobranchiae

Arthrobranchiae

1+r

Podobranchiae

Epipods

Exopods

Source:

202

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

dorsal view. Scale = 0.5 mm.

of segments. First to third pereopods slender,

increasing in length posteriorly, each bearing

small distinct chela; fourth and fifth pereopods

flattened, of 7 segments, somewhat broader than

preceding pereopods, posterior margin of propod

and carpus setose, anterior and posterior margins

of merus and ischium setose; fifth pereopod

shorter than fourth pereopod.

Petasma with lamellar processus uncifer, broad

lobus armatus, strong processus ventralis distally

divided into 2 lobes. Seminal receptacles broad,

shallow, situated laterally at base of third

pereopods, beneath broad subcircular flaps.

Stemites of somites XIII and XIV in female gently

convex.

Species. — Sicyonella antennata Hansen,

1919. Indonesia; Thailand.

Sicyonella elegans Caiman, 1913. Torres

Straits.

Sicyonella inermis (Paulson, 1875). Red Sea.

Sicyonella maldivensis Borradaile, 1910. Red

Sea; Cargados Carajos Shoals; Maidive Islands;

Indonesia; Palau; Bikini Atoll.

Source:

REFERENCES

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Zariquiey Alvarez, R., 1968. — Crustaceos Decdpodos Ibericos. Investigacion Pesquera , 32: i-xv + 1-510.

Source: MNHN, Paris

INDEX TO TAXA

Valid taxa are printed in bold. In the pagination the major entry for a taxon is in bold. Pages

having figures of the taxa are given in italics.

acclivirostris, Parapenaeopsis . 120

acclivis, Metapenaeopsis . 104

Acetes . 12, 184, 187

Acheles . 194

acuminata, Solenocera . 180

aegyptia, Metapenaeopsis . 104

affiniSy Hadropenaeus . 163, 164, 165

affiniSy Metapenaeus . 109, 110, 111

affinis, Penaeus . 109

affiniSy Sicyonia . 155

africana, Solenocera . 180

africana, Macropetasma . 90, 91, 92

africanus, Parapeneus . 91

agassizii, Solenocera . 180

akayebi, Erythropenaeus . 102

akayebi, Parapenaeus . 102

albicoma, Trachysalambria . 148, 149

alcockiy Aristeus . 40

alcocki, Metapenaeus . 110

alfonso, Solenocera . 180

algoensiSy Solenocera . 180

aliaffiniSy Sicyonia . 155

alticarinata, Solenocera . 180

altuSy Benthesicymus . 60

Amalopenaeus . 64

americanuSy Acetes americanus . 188

americanuSy Parapenaeus . 123

anchistusy Metapenaeus . 110

anchoralis, Penaeus . 144

anchoraliSy Trachypenaeus .... 144, 145, 146

andamanensis, Metapenaeopsis . 104

angusta, Metapenaeopsis . 104

annectens, Parasolenocera . 179

annectens, Solenocera . 180

antennata, Sicyonella . 202

antennatus, Aristeus . 40, 41

antennatus, Penaeus . 39

antillensis, Aristeus . 41

Aphareocaris . 201

Aphareus . 200

aphoticus, Hymenopenaeus . 172

arachnipodus, Acheles . 194

araclinipoduSy Sergestes . 196

arafurica, Parapenaeopsis . 120

Archipenaeopsis . 102

arcticuSy Sergestes . 196

Aristaeidae. 32

Aristaeina. 32

Aristaeinae. 32

Aristaeomorpha . 12, 32, 33, 34

Aristaeopsis . 33, 36, 43

Aristaeus . 36, 39, 46, 50, 52

Aristeae. 32

Aristeidae. 12, 32, 33

Aristeinae. 32

Aristeomorpha . 34

Aristeopsis . 36, 51

Aristeus . 12, 33, 36, 39, 50, 83

armatus, Aristeus . 51

armatus, Benthesicymus . 60

armatus, Plesiopenaeus . 38, 50, 52

armatus, Sergestes . 196

aroaensis, Parapenaeopsis . 120

Artemesia . 12, 67, 69, 71, 127

Artemisia . 71

aspera, Trachysalambria . 149

assimilis, Metapenaeopsis . 104

atlantica, Parapenaeopsis . 118, 779, 120

atlanticuSy Sergestes . 194, 195, 196

atlantidis, Solenocera . 180

Atypopenaeus . 12, 70, 73, 74, 114

Atypopeneus . 73

australiana, Solenocera . 181

australiensis, Haliporoides sibogae . 167

australiensis, Parapenaeus . 123

australis, Acetes sibogae . 189

Austropenaeus . 12, 33, 41

aztecus, Farfantepenaeus . 77, 79

aztecus, Penaeus brasiliensis var. 78

Source: MNHN. Paris

226

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

balboae, Pelagopenaeus . 125, 126

balboae, Penaeus . 124

balli, Parapenaeopsis . 116, 118, 120

balssi, Penaeopsis . 130

barbari , Gennadas . 65

barbata , Metapenaeopsis . 104

bartlettiy Benthesicymus .59, 60

barunajaya, Solenocera . 181

bedokensiSy Solenocera . 181

beebeiy Metapenaeopsis . 104

bennettae , Metapenaeus . 110

Bentheogennema . 12,56,57

Benthesicymae. 56

Benthesicymidae. 12, 32, 56

Benthesicymina. 56

Benthesicyminae. 56

Benthesicymini. 56

Benthesicymus . 12, 56, 59

Benthoecetes . 59

Benthonectes . 12, 56, 61

benthophila, Sicyonia . 155

bicorniSy Atypopenaeus.. 73, 75, 76 , 77, 116

bigemmeay Sergia . 199

binghamiy Acetes . 188

bispinosa, Sicyonia . 155

bisulcatay Sergia . 199

borealis , Bentheogennema . 58

bouvieriy Gennadas . 65

brasiliensiSy Benthesicymus . 60

brasiliensis, Farfantepenaeus .78, 79

brevicorniSy Metapenaeus . Ill

brevirostrisy Benthesicymus . 60

brevirostris, Cryptopenaeus . 211

brevirostrisy Farfantepenaeus .78, 79

brevirostris, Gennadas . 66

brevirostris, Sicyonia . 155

brevispinatuSy Sergestes . 196

brevisuturae f Trachysalambria . 148, 149

bruceiy Mesopenaeus . 176

burkenroadiy Bentheogennema . 58

burkenroadiy Sicyonia . 155

byrdiy Rimapenaeus . 136, 138

californiensiSy Farfantepenaeus . 79

canaliculatuSy Melicertus . 68, 100, 101

Cancer . 79, 87, 99, 153

capensiSy Gennadas . 66

caribaeuSy Tanypenaeus . 139, 140 , 141

carinata, Parartemesia . 173, 177

carinata , Sicyonia . 153, 154, 755, 155

carinatuSy Benthesicymus . 60

carinatus, Cancer . 153, 154

carolinae } Acetes americanus . 188

carpenteri, Hemipenaeus . 46

catherinae , Cryptopenaeus.... 158, 759, 160

Ceratopenaeus . 102

cereuSy Benthesicymus . 60

ceylonicay Metapenaeopsis . 104

chaceiy Hymenopenaeus . 172

chaceiy Lucifer . 185

challengeri, Penaeopsis . 130

challenged, Sergia . 199

chinensiSy Acetes . 188

chinensiSy Fenneropenaeus . 82

chopraiy Solenocera . 181

clavicarpuSy Gennadas . 66

clevaiy Cryptopenaeus . 160

coalituSy Sergestes . 196

colosiiy Sergia . 199

comatay Solenocera . 181

commensalis, Metapenaeopsis . 104

complanata, Metapenaeopsis mogiensis .

. 107

compressipesy Atypopenaeus . 77, 114

compressipes, Penaeus . 74

coniger, Metapenaeopsis . 106

conjunctuSy Metapenaeus . Ill

consobrina y Metapenaeopsis mogiensis.. 107

consobrinuSy Sergestes . 196

constrictus, Penaeus . 135

constrictuSy Rimapenaeus . 138

corniculum, Sergestes . 194

cornuta, Parapenaeopsis . 120

cornutuSy Sergestes . 196

coromandelicay Parapenaeopsis stylifera .

. 116, 118, 119 , 121

coruscans, Plesiopenaeus . 52

costata, Metapenaeopsis . 106

crassicornisy Solenocera . 181

Crassipenaeus . 158

erassipes, Aristeus . 52

crassipesy Pseudaristeus . 54, 55

crassissima , Metapenaeopsis . 106

crass us, Gennadas . 66

crenatuSy Benthesicymus . 60

cristata , Sicyonia . 155

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

227

cristatus, Haliporoides . 167

crosnieri, Cryptopenaeus . 160

Cryptopenaeus . 12,158

cultirostrisy Parapenaeopsis . 120

curvatuSy Sergestes . 196

curvirostrisy Haliporus . 168, 169, 170

curvirostris, Penaeus . 146, 147

curvirostris, Sicyonia . 155

curvirostris, Trachysalambria .

.98, 147, 148, 149, 151

daleiy Metapenaeopsis . 106

dalliy Metapenaeus . Ill

danae, Funchalia . 85

dearmatusy Atypopenaeus . 74, 77, 114

debilis, Hymenopenaeus . 777, 172, 173

demaniy Metapenaeus demani . Ill

danae, Funchalia . 85

Decapoda. 31

Dendrobranchiata . 15, 31

denticulata } Parapenaeus longipes forma....

diapontiuSy Sergestes . 196

difficilis, Metapenaeopsis . 106

diomedeae, Faxonia . 177

diomedeae y Haliporoides . 167, 177

disdorsalisy Sicyonia . 155

disedwardsiy Sicyonia . 755, 156

disjunctuSy Sergestes . 197

disparriy Sicyonia . 156

distinctay Metapenaeopsis . 106

dobsoni, Mangalura . 108

dobsoniy Metapenaeus . 108, 111

dons, Hymenopenaeus . 172

dorsalis, Sicyonia . 156

duorarumy Farfantepenaeus .78, 79

dura 9 Metapenaeopsis . 106

eboracensis, Metapenaeus . Ill

eduardoiy Penaeopsis . 729, 130

edwardsiy Sergestes . 197

edwardsiana, Aristaeopsis . 37, 38 , 39

edwardsianus, Penaeus . 37

elegans, Amalopenaeus . 64

eleganSy Gennadas . 64 , 66

eleganSy Metapenaeus . Ill

elegans, Sicyonella . 202

endeavouri, Metapenaeus . Ill

ensisy Metapenaeus . 110, 111

equaliSy Hymenopenaeus . 172

erectuSy Sergestes . 197

erythraea y Metapenaeopsis . 106

erythraeus, Acetes . 189

erythraeuSy Parapenaeus fissuroides . 123

Erythropenaeus . 102

esculentusy Penaeus . 133

Eusicyonia . 153

evermanniy Metapenaeopsis . 106

extensusy Sergestes . 197

extenuata, Sergia. . 199

fallaxy Sicyonia .156

faoe y Rimapenaeus . 135, 136, 137, 138

faoe, Trachypenaeus . 135

faouziiy Metapenaeopsis . 106

Farfantepenaeus . 12, 68, 77, 101, 131

fattahiy Hymenopenaeus . 172

Faxonia . 165, 177

faxoni, Lucifer . 185

faxoni, Solenocera . 181

Fenneropenaeus . 12, 68, 79, 80, 131

filicta, Sergia . 199

filipes, Benthonectes . 62, 63

fissuroides, Parapenaeus fissuroides . 123

fissurus, Parapenaeus . 123

florea, Solenocera . 181

foliacea, Aristaeomorpha . 34, 35, 36

foliaceum , Petalidium . 191, 792, 193

formosa, Sicyonia . 156

formosus, Atypopenaeus . 75, 77

fulgens, Sergia . 199

fulva, Trachysalambria . 149

Funchalia . 12, 67, 68, 82, 124

Funchaliae. 67

Funchaliinae. 67

Funchalis . 83

furcata, Sicyonia . 156

fund, Hymenopenaeus . 172

fusca, Metapenaeopsis . 106

fuscina, Rimapenaeus . 137, 139

fuscina, Trachy penaeus . 69

Source: MNHN. Paris

228

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

gaillardi , Metapenaeopsis . 106

galeata, Sicyonia . 156

gallensis, Metapenaeopsis . 106

gardineri, Sergia . 199

geijskesi , Solenocera . 181

geminus, Sergestes . 197

Gennadas . 12,56,57,59,63

gerardoiy Metapenaeopsis . 106

gibbolobatus, Sergestes . 197

gilchristiy Gennadas . 66

glacialiSy Hepomadus . 47, 48

gonospinifer, Megokris . 96, 99

goodeiy Metapenaeopsis . 102, 104, 106

Gordonella . 12, 157, 160

gracilirostris, Panaeus . 88

gracilisy Pseudaristeus .54, 55

gracillima, Parapenaeopsis . 120

grandiSy Sergia . 200

granulosuSy Megokris . 95, 96, 97 , 99

granulosus, Penaeus . 95

Grimaldiella . 83

gurjanovae, Solenocera . 181

Hadropenaeus .

. 12, 158, 163

halia, Sergestes .

. 197

Haliporae.

. 67

Hal in nr aides .

. 17. 1 58. 165

Halioorus ...

.. 12, 157, 160, 163, 165, 168,

170, 174, 177

halliy Hymenopenaeus .

. 173

halliy Solenocera .

. 181

hamifer , Sergestes .

. 197

hanseni, Lucifer .

. 185

hansjacobiy Sergia .

. 200

Hapalopoda .

hardwickiiy Parapenaeopsis ....

. 118, 120

hartii, Xiphopenaeus .

. 150

hat ho r , Melicertus .

. 102

Hernipenaeopsis .

. 83

Hemipenaeus . 12,

33, 43, 45, 52

Hemipeneus .

. 44

Hemypenaeus .

.. 44

henseniy Sergestes .

. 197

Hepomadus . 12, 33, 46

Heteropenaeus . 12, 68, 85

Heteropeneus . 85

hextiiy Solenocera . 181

hilarula, Metapenaeopsis . 106

Hippolyte . 153

hjorti, Benthesicymus . 61

hobbsiy Metapenaeopsis . 106

hungerfordi, Parapenaeopsis . 120

Hymenopenaeus .

. 12, 158, 163, 165, 168, 170, 174, 176

Hymenopeneus . 171

incertuSy Gennadas . 66

incertuSy Sergestes . 197

incisa, Metapenaeopsis . 106

incisa , Parapenaeopsis . 120

index, Sergestes . 197

indicusy Acetes . 186, 187, 188 , 189

indicus, Fenneropenaeus . 80, 81, 82

indicusy Parapenaeus fissuroides . 123

indicus, Penaeus . 80

inequaliSy Sergia . 200

inermis, Aphareus . 200, 201

inermis, Hepomadus . 48

inermis, Sicyonella . 202

inermis, Solenocera alfonso . 180

inflexa, Sicyonia . 156

ingentiSy Sicyonia . 156

inoa, Sergia . 200

insolituSy Metapenaeus . Ill

intermedia , Bentheogennema . 57, 58, 61

intermedia , Metapenaeopsis mogiensis .

. 107

intermediuSy Acetes . 189

intermedius, Gennadas . 57

intermedius, Lucifer . 185

intermedius , Metapenaeus . 111

investigator, Hapalopoda . 62

investigatoriSy Benthesicymus . 61

investigatoriSy Parapenaeus . 124

iridescenSy Benthesicymus . 61

ivanoviy Metapenaeopsis . 106

japonicay Sergia . 200

japonica, Sicyonia . 156

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

229

japonicuSy Acetes . 189

japonic us, Marsupenaeus . 93, 94

japonicus, Penaeus canaliculatus var. 92

jerryi, Penaeopsis . 130

johni, Acetes . 189

joyneri, Metapenaeus . 112

kathleenae, Pseudaristeus . 53, 54, 55

kernpi, Gennadas . 66

kensleyi, Gordonella . 161, 162

kerathurus, Cancer . 100

kerathurus, Melicertus . 15, 100, 101 , 102

kishinouyei, Metapenaeopsis . 106

koelbeli, Solenocera . 182

krishnatrii, Metapenaeus . 112

kroyeri, Xiphopenaeus . 150, 151 , 152

kutchensis, Metapenaeus . 112

kyushuensis, Metapenaeopsis . 106

laciniatus, Benthesicymus . 61

laevigata , Sicyonia . 156

laeviSy Hymenopenaeus . 173

laevis, Sicyonia . 156

lamellata, Metapenaeopsis . 106

laminata> Sergia . 200

lanceolatuSy Parapenaeus . 124

lancifer } Sicyonia . 156

latay Metapenaeopsis . 106

latisulcatusy Melicertus . 102

laubieriy Metapenaeopsis . 106

Leptopenaeus . 102

Leucifer . 183

Leuciferidae. 183

Leuciferinae. 183

lindae, Metapenaeopsis . 106

Litopenaeus . 12, 15, 16, 68, 87, 89, 128

liuiy Metapenaeopsis . 106

longicauda , Sicyonia . 156

longimanus, Heteropenaeus . 85, 86 , 87

longinariSy Artemesia . 71, 72 , 73

longipes, Parapenaeus . 121, 124

longipes, Trachysalambria . 149

longirostriSy Metapenaeopsis provocatoria...

.107

longirostris, Parapenaeus .

. 121, 722, 123 , 124

longirostris, Penaeus . 121

longistyliSy Melicertus . 102

lucasiiy Hadropenaeus . 164, 176, 179

lucasii, Philonicus . 176, 179

lucenSy Sergia . 200

Lucifer . 12, 183

Luciferidae. 12, 182, 183

Luciferinae. 183

lysianassa f Metapenaeus . 112

mabahissae , Aristeus . 41

macleayiy Metapenaeus . 112

Macropetasma . 12, 67, 69, 90

madagascariensiSy Haliporoides sibogae .

.167

Malacostraca. 31

rnalaianay Trachysalambria . 148, 149

maldivensiSy Sicyonella . 201, 202

Mangalura . 108

mannarensis, Metapenaeopsis . 107

manningi, Metapenaeopsis . 107

marginatus, Melicertus . 100, 102

mariae , Mesopenaeus . 176

marinuSy Acetes . 189

marquesasy Metapenaeopsis . 107

Marsupenaeus . 12, 68, 82, 92

martinella, Metapenaeopsis . 107

martiniy Sicyonia . 156

mascarensis , Solenocera . 182

maxillipedOy Parapenaeopsis . 120

maximay Sergia . 200

Megokris . 12, 70, 95, 99, 138, 146

melanthOy Solenocera . 182

Melicertus ... 12, 15, 68, 77, 92, 99, 100, 131

membranacea , Solenocera .... 179, 181 , 182

membranaceous, Penaeus . 179

menouiy Metapenaeopsis . 107

merguiensisy Fenneropenaeus . 81, 82

Mesopenaeus . 12, 158, 173

Metapenaeopsis . 12, 69, 102

Metapenaeus .

. 12,69,76, 102,108, 127, 141, 146

Metapeneopsis . 102

Source:

230

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

Metapeneus . 102, 109, 117

miersi, Metapenaeopsis . 103, 105, 107

miersi, Penaeopsis . 103

mineri, Metapenaeopsis . 107

minicoyensisy Trachypenaeopsis . 143

mixtay Sicyonia . 156

Miyadiella . 12, 70, 112

mobilispinis, Metapenaeus . 142

mobilispiniSy Trachypenaeopsis .

. 141, 142, 143

modestuSy Hadropenaeus . 165

modestus, Hymenopenaeus . 163

mogiensis, Ceratopenaeus . 102

mogiensis, Metapenaeopsis mogiensis.... 1 07

mogiensis, Parapenaeus . 102

monoceroSy Metapenaeus . 112

monodotiy Penaeus . 130, 131, 132, 133

moosaiy Solenocera . 182

moyebiy Metapenaeus . 112

muelleriy Pleoticus . 163, 173, 177, 178, 179

mulleri, Philonicus . 163, 177, 179

murrayi, Parapenaeus . 124

mutator , Solenocera . 182

nana , Parapenaeopsis . 121

nasica y Sicyonia . 156

natalensiSy Acetes . 189

nebulosay Sicyonia . 156

necopitta, Solenocera . 182

Neopenaeopsis . 121

neptunusy Hymenopenaeus . 173

nereuSy Hymenopenaeus . 173

Nika . 194

nipponensis, Sergestes . 197

nitiduSy Austropenaeus . 42, 43

nitidus, Plesiopenaeus .38, 41, 42

nodosa, Parapenaeus australiensis forma....

notialiSy Farfantepenaeus . 79

novaeguineae , Metapenaeopsis . 107

nuduSy Sergestes . 197

obesum y Petalidium . 193

obliquirostriSy Hymenopenaeus . 173

ocellatay Sicyonia . 156

occidentalis, Litopenaeus . 90

ocularis, Penaeopsis . 165, 177

olgae f Sicyonia . 156

ommanneyiy Sicyonia . 156

orientalis, Lucifer . 185

orientalis, Sergestes . 197

ornata, Miyadiella . 113, 114, 115

pacificuSy Rimapenaeus . 139

Palaemon . 153

pallidicauda y Aristeus . 41

palmensiSy Metapenaeopsis . 107

Panaeus . 88

Paneus . 77

papuensis, Metapenaeus . 112

paradoxus, Neopenaeopsis . 121

parafallaxy Sicyonia . 156

ParaguayensiSy Acetes . 189

Parahaliporus . 165

Parahepomadus . 12, 33, 48

parahilarula , Metapenaeopsis . 107

parapalmensiSy Metapenaeopsis . 107

Parapenaeinae.67, 157

Parapenaeopsis . 12, 16, 70, 73, 102, 117

Parapenaeus . 12, 69, 102, 121, 127, 135

Parapeneopsis . 117

Parapeneus . 90, 121, 127

Parartemesia . 173, 177

paraseminuduSy Sergestes . 197

Parasolenocera . 179

paravillosay Gordonella . 162

parriy Sicyonia . 156

parvula , Sicyonia . 156

parvus, Gennadas . 64, 65, 66

Pasiphodes . 64

pasithea , Bentheogennema . 59

paulensiSy Farfantepenaeus . 79

pectinata, Solenocera . 163, 176, 182

pectinatuSy Gennadas . 66

pectinatus, Philonicus . 163, 176

pectinatuSy Sergestes . 186, 197

pectinulatay Solenocera . 182

pediformisy Sergestes . 197

pedunculata, Miyadiella . 112

Peisos . 12, 186, 187, 189

Pelagopenaeus . 12, 68, 124

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

231

Penaeidae. 12, 32, 56, 67, 152, 157

Penaeidea. 184

Penaeinae. 67

Penaeoidea. 12, 31

Penaeopsis .

. 12, 69, 102, 109, 117, 126. 141, 165

Penaeus .... 12, 33, 36, 39, 68, 73,77, 80, 83,

87, 92, 95, 99, 102, 108, 112, 117, 121,124,

126, 130, 135, 141, 144, 146, 149, 178

Penedia. 67

Peneinae. '57

Peneopsis . 1-7

Peneits . 77, 79, 88,92,99, 109, 117,

121, 124, 127, 130, 135, 144, 146, 178

penicillata , Sicyonia . 156

penicillatus, Fenneropenaeus . 82

penicillifer, Lucifer . 185

Penoeus . 102, 130

perezfarfantae , Parapenaeus . 124

persica, Metapenaeopsis . 107

pescadoreensiSy Megokris . 98 , 99

pestafer , Sergestes . 197

Petalidium . 12, 187, 191

petrunkevitchi, Peisos . 189, 190, 191

philippii, Metapenaeopsis . 107

philippi, Solenocera . 179

philippii, Penaeus . 102

Philippinensis, Leptopenaeus . 102

Philonicus . 163, 176, 177, 179

phuongiy Solenocera . 182

picta y Sicyonia . 156

plebejusy Melicertus . 102

Pleoticus . 12, 128, 158, 163, 176, 179

Plesiopenaeus . 12, 33, 38, 41, 43, 50, 51

Plesiopeneus . 36,51

plumea, Sergia . 200

Plyonuria. 67

podophthalmuSy Miyadiella... 113 , 114, 115

podophthalmus, Penaeus . 112

polituSy Parapenaeus . 122, 123 , 124

polyarthra, Gordonella . 160 , 161

potenSy Sergia . 200

precipua, Protrachypene . 133, 134, 135

prehensiliSy Sergia . 200

profunda y Sergia . 200

propinqua y Metapenaeopsis . 107

propinquuSy Gennadas . 66

propinquus, Hymenopenaeus . 173

protensiSy Pseudaristeus . 55

Protrachypene . 12, 67, 69, 133

provocatoria y Metapenaeopsis provocatoria

. 107

proximo , Metapenaeopsis . 107

Pseudaristeus . 12,33,52

pubescens, Metapenaeopsis . 103

pubescens, Penaeus . 103

purpureus, Pasiphodes . 64

quadrilobatay Metapenaeopsis . 107

quinquedentatay Metapenaeopsis . 107

rathbunae , Solenocera . 182

rectacutay Penaeopsis . 128, 130

rectirostrisy Sicyonia . 156

regalis, Sergia . 200

remipeSy Sergia . 198

richardi, Grimaldiella . 83

richeriy Metapenaeopsis . 107

richtersiiy Trachypenaeopsis . 144

Rimapenaeus .

robusta, Sergia . 198, 199 , 200

robustuSy Pleoticus . 165, 176 , 177, 178

rosea, Metapenaeopsis . 107

rostridentatus, Aristeus . 34

ruberoculatuSy Parapenaeus . 124

rubroguttatuSy Sergestes . 197

Ruvulus . 153

sagamiensiSy Funchalia .

Salambria .

sargassiy Sergestes .

schmittiy Litopenaeus .

scintillans, Sergia .

scottiy Metapenaeopsis .

sculptay Sicyonia ..

sculptiliSy Parapenaeopsis.

scutatus, Gennadas .

sediliy Megokris .

semidentatus, Aristeus .

seminuduSy Sergestes .

semis sis, Sergestes .

.. 84, 85

. 99

. 197

. 90

. 200

. 107

153, 154

. 121

. 66

... 98 , 99

. 41

. 197

Source:

232

ISABEL PEREZ FARFANTE & BRIAN KENSLEY

semisulcatus, Penaeus . 133

Sergestes . 12, 186, 187, 191, 194, 198

Sergestidae. 12, 182, 185, 187

Sergestinae. 185

Sergestoidea. 12, 31, 182

Sergia . 12, 186, 187, 198

serrata, Penaeopsis . 127, 128, 129 , 130

serratus, Penaeopsis . 127

serrulatus, Acetes . 189

setiferuSy Litopenaeus . 15, 88, 89 , 90

sewelli, Hymenopenaeus . 173

sextuberculatuSy Parapenaeus . 124

seymouriy Benthesicymus . 61

sibogae, Acetes sibogae . 189

sibogae , Haliporoides sibogae . 167

sibogae, Haliporus . 165

sibogae y Metapenaeopsis . 107

sibogae, Parahaliporus . 165

sibogaey Pseudaristeus . 55

sibogalis, Acetes sibogae . 189

Sicyonella . 12, 186, 187, 200

Sicyonia . 12, 83, 153

Sicyonidae. 152

Sicyoniidae. 12, 32, 152

Sicyoninae. 152

silasiy Fenneropenaeus . 82

similis, Rimapenaeus . 137, 139, 151

similiSy Sergestes . 197

sinensis, Crassipenaeus . 158

sinensis , Cryptopenaeus . 160

sinica y Metapenaeopsis . 108

sinica, Parapenaeopsis . 121

sinuosa, Metapenaeopsis . 108

siphonoceros, Penaeus . 179

smithiy Metapenaeopsis . 108

Solenocera . 12, 158, 163, 178

Solenoceridae. 12, 32, 157

Solenocerina. 157

Solenocerinae. 157

sordiduSy Gennadas . 66

spatulata y Metapenaeopsis . 108

speciosusy Pseudaristeus . 55

spinajugOy Solenocera . 182

spinicauda, Hadropenaeus . 165

spinidorsalisy Hemipenaeus . 44, 45, 46

spiridonoviy Metapenaeopsis . 108

splendenSy Sergia . 200

stebbingiy Metapenaeus . 112

steindachneriy Pleoticus . 178

stellata y Sergia . 200

stenodactylus, Atypopenaeus .

. 74, 75, 76, 11, 114

stenodactylus, Penaeus . 114

stephani, Metapenaeus demani . 111

stephenseniy Bentheogennerna . 59

stimpsoniy Sicyonia . 156

stimulator, Sergestes . 197

stokmaniy Metapenaeopsis . 108

strabus, Benthesicymus . 61

stridulanSy Metapenaeopsis . 108

stylifera y Parapenaeopsis . 69

stylifera, Parapenaeopsis stylifera . 121

sty life rus, Penaeus . 117

stylirostrisy Litopenaeus .89, 90

subtiliSy Farfantepenaeus . 79

suluensis, Metapenaeus . 112

suspiriosumy Petalidium . 193

Synhimantites... . 153

taaningi, Funchalia . 85

talismaniy Gennadas . 66

talismaniy Sergia . 200

tanneriy Benthesicymus . 61

tantilluSy Sergestes . 197

Tanypenaeus . 12, 70, 139,

taprobanensis, Haliporus . 168, 169, 170

tarawensiSy Metapenaeopsis . 108

tchekunovae, Metapenaeopsis . 108

tenella , Metapenaeopsis . 108, 121

tenella, Parapenaeopsis . 121

tener , Hepomadus . 46, 47, 48

tenuipes, Metapenaeus . 112

tenuiremiSy Sergia . 200

thetiSy Haliporus . 169, 170

tigrinuSy Melicertus . 100

tinayreiy Gennadas . 66

tirmiziae y Benthesicymus . 61

toloensis, Metapenaeopsis . 108

Trachypenaeopsis . 12, 69, 141

Trachypenaeus .

... 12, 70, 95, 97, 99, 117, 135, 138, 144, 146

Trachypeneopsis . 141

Trachypeneus . 95, 135, 144, 146

Trachy salambria .

. 12, 70, 97, 99, 135, 138, 146

Transolenocera . 179

triarthruSy Haliporoides . 165

Source

FAMILIES AND GENERA OF THE PENAEOID AND SERGESTOID SHRIMPS

233

triarthrus , Haliporoides triarthrus . 167

trispinosa , Sicyonia . 156

trisulcatus, Penaeus . 101

tropicalis f Mesopenaeus . 174, 175 , 176, 177

tropicalis, Parartemesia . 174, 177

truncata , Sicyonia . 157

tuerkayi, Hymenopenaeus . 173

typica y Sicyonia . 157

typicus, Synhimantites . 153

typus, Leucifer . 183

typus , Lucifer . 184 , 185

uncta, Parapenaeopsis . 121

urinator, Benthesicymus . 61

vaillantiy Metapenaeopsis . 108

valens, Gennadas . 64, 66

vannamei, Litopenaeus . 89, 90

vannamei, Penaeus . 88

varidens, Aristeus . 41

vaubani, Parahepomadus . 48, 49 , 50

velutina , Metapenaeopsis . 108

venusta , Parapenaeopsis . 121

verpus, Sergestes . 197

vestitus, Archipenaeopsis . 102

vigilax y Sergestes . 197

villaluzi, Trachysalambria . 149

villosa, Funchalia . 85

villosa, Gordonella . 160 , 162

villosus, Haliporus . 161

villosus, Hemipenaeopsis . 83

vioscaiy Solenocera . 182

viriliSy Aristeus . 41

vitulansy Sicyonia . 157

vniroi, Haliporoides triarthrus . 166, 167

vulgaris , Acetes . 189

waltairensiSy Solenocera . 182

wellsiy Metapenaeopsis . 108

wheeleriy Sicyonia . 157

wolffiy Sergia . 200

woodmasoniy Aristaeomorpha . 36

woodwardiy Funchalia . 82, 83, 84 , 85

Xiphopenaeus . 12, 70, 140, 149, 150

Xiphopeneus . 149

zarenkoviy Solenocera . 182

BIBL QU

RAP4S

* 1

Source:

Remerciements aux rapporteurs / Acknowledgements to referees

La Redaction tient a rcmercier ies experts extdrieurs au Museum national d’Histoire naturelle dont les noms suivent, d’avoir bien voulu contribuer,

avec les rapporteurs de 1’Etablissement, a revaluation des manuscrits (1994/1997) :

The Editorial Board acknowledges with thanks the following referees who. with Museum referees, have reviewed papers submitted to the Memoires

du Museum (1994/1997):

Adkison D.

Macon

Afzelius B

Stockholm

AkamM.

Cambridge

Akesson B.

Goteborg

Amlard J.C.

Nantes

Andersen N.

Copenhague

Andres H.

Hambourg

Baba K.

Kumamoto

Bachelet G.

Arcachon

Bachmann G. H.

Halle-Wittenberg

Bally A. W.

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BellanG.

Marseille

Bell ido A.

Paimpont

Ben-Eliahu N.

Jerusalem

Berggren M.

Fiskebackskil

Bernoulli D.

Zurich

Bertotti G.

Amsterdam

Bessereau G.

Rueil-Malmaison

BestM.

Leiden

Bhaud M.

Banyuls-sur-Mer

Blake J. A.

Woods Hole

Bourdon R.

Roscoff

Bourseau J.P.

Villeurbanne

Bruce J.

Helensvale

Butman C. A.

Woods Hole

Carpenter J.

New York

Cassagneau P.

Toulouse

Castelli A.

Modena

Chace F. A.

Washington

Charest P.

Quebec

Cherix D

Lausanne

Clark P

Londres

ClobertJ.

Paris

Cloetingh S.

Amsterdam

Cohen D.

Los Angeles

CoRNUDELLA L.

Barcelone

CUZIN-ROUDY J.

Villefranchc-sur-Mer

DarluP.

Paris

Danchin E.

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Davie P.

Brisbane

De B ROYER C.

Bruxelles

Dejean A.

Villetaneuse

Deleporte P.

Paimpont

DesbruyEres D.

Brest

Dhainaut A.

Villeneuve d’Ascq

Dietrich C.

Champaign

Dorresteun A.

Mayence

DuchEne J.C.

Banyuls-sur-Mer

Duffels J. P.

Amsterdam

Eibye-Jacobsen D.

Copenhague

Eldredge L. L.

Hawaii

Fahay M.

Highlands

Fauchald K.

Washington

Fischer A.

Mayence

Fleury A.

Orsay

Fournier J.

Ottawa

Fransen C.

Leiden

Gagne R

Washington

Gambi M. C.

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G until F.

Roscoff

George D.

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Kensley B.

Kielan-Jaworowska Z.

Knight-Jones P

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KomaiT.

Kristens en N.

Kudenov J. D.

USA

LagardEre J.P.

Suide

Lana P. C.

Grande-Bretagne

LaubierL.

Suide

Laubscher H P

France

Laverde-Castillo J. J. A.

Danemark

LemattreR.

Allemagne

Levin L. A.

Japon

Lovelock P. E. R.

France

Machida Y.

Allemagne

Mack ie A.

USA

MacPherson E.

France

M ADDISON D.

France

Manning R.

Israel

Markle D.

Suede

Masaki S.

Suisse

MascleA.

Pays-Bas

Mauchline J.

France

Maurer D.

Pays-Bas

McLaughlin P.

France

McLennan D.

USA

MerrettN.

France

Messing C.

France

Mettam C.

Australie

Morand S.

USA

Muir A. I.

USA

Nakamura I.

France

Naumann C.

Italie

Newman W. A.

USA

Oliva R.

Canada

Olive P. J W.

Suisse

Orousset J.

Grande-Bretagne

Packer L.

France

Paterson G. L. J.

Pays-Bas

Paxton H

USA

PEREZ FaRFANTE I.

Espagne

Perkins T. H.

France

Peter NG

France

Petersen M. E.

France

Pettibone M. H.

Australie

Peyrot-Clausade M.

Belgique

Plateaux C.

France

PleuelF

France

Poccia D. L.

France

POCKLINGTON P.

France

Poore G.

USA

Purschke G.

Allemagne

Raikova 0.

France

Reish D. J.

Pays-Bas

Rente D. C. R.

Danemark

Richards W.

USA

Richer de Forges B.

USA

RI EM AN F.

USA

Roberts C.

Allemagne

Roure F.

France

Rouse G.

Canada

Salomon M.

Pays-Bas

San Martin G.

USA

Sard a R.

Italie

Scholtz C.

France

Schmid S. M.

Grande-Bretagne

Schroeder P.

Italie

Schwander M.

Grandc-Bretagne

SibuetM.

France

Sigvaldadottir E.

Australie

Simon J. L.

France

Spiridonov V.

Allemagne

Stefanescu M. 0.

Grande-Bretagne

Stewart A.

Suisse

Takeda M.

Canada

Tan C. G. S.

Canada

Tassy P

France

Taylor P. D.

Italie

Thorne B

Australie

Thurston M. H.

Suisse

Toulmond A.

Allemagne

Tudge C.

Australie

Van Ameron H. W. J.

Afrique du Sud

VanBaaren J.

USA

Van Soest R. W. M.

USA

Vernon P.

Grande-Bretagne

Vickery Vernon R.

Norvfcge

Vovelle J.

Pays-Bas

Vul M. A.

Australie

Wagele J. W

Hongrie

Warren L.

Pays-Bas

Watson N.

Australie

Wenzel J.

Allemagne

Westheide W.

Suisse

Wiegmann B.

Grandc-Bretagne

Wilson M.

USA

Wilson R.

Norvfcge

Wilson S.

Grande-Bretagne

Yeates D.

Grande-Bretagne

Zevina G B.

Japon

Zibrowius H.

Danemark

USA

Ziegler P. A.

La Rochelle

France

Parana

Brasil

Paris

France

Bale

Suisse

Bogota

Colombic

Washington

USA

La Jolla

USA

La Haye

Pays-Bas

Kochi

Japon

Cardiff

Grande-Bretagne

Barcelona

Espagne

Tucson

USA

Washington

USA

Oregon

USA

Hirosaki

Japon

Rueil-Malmaison

France

Oban

Grande-Bretagne

Long Beach

USA

Washington

USA

Toronto

Canada

Londres

Grande-Bretagne

Dania

USA

Cardiff

Grande-Bretagne

Perpignan

France

Londres

Grande-Bretagne

Kyoto

Japon

Bonn

Allemagne

San Diego

USA

Barcelone

Tyne

Espagne

Grande-Bretagne

Paris

France

York

Canada

Londres

Grandc-Bretagne

North Ryde

Australie

Washington

USA

Saint Petersburg

USA

Singapore

Singapour

Copenhague

Danemark

Washington

USA

Marseille

France

Nancy

France

Stockholm

Sufcde

Amherst

USA

Halifax

Canada

Victoria

Australie

Osnabruck

Allemagne

Saini-P£tersbourg

Russie

Long Beach

USA

Canberra

Australie

Miami

USA

Noum6a

Nouvelle-Cal6donie

Bremerhaven

Allemagne

Wellington

Nouvelle-Z^lande

Rueil-Malmaison

France

Washington

USA

Marseille

France

Madrid

Espagne

Blanes

Espagne

Pretoria

Afrique du Sud

Bale

Suisse

Pullmann

USA

La Haye

Pays-Bas

Brest

France

Stockholm

Sufcde

Tampa

USA

Moscou

Russie

Bucarest

Roumanie

Wellington

Nouvelle-Z61ande

Tokyo

Japon

Singapore

Singapour

Paris

France

Londres

Grande-Bretagne

Maryland

USA

Wormley

Grande-Bretagne

Paris

France

Brisbane

Australie

Krefcld

Allemagne

Rennes

France

Amsterdam

Pays-Bas

Paimpont

France

Ste-Anne-de-Bellevue

Canada

Paris

France

Lvov

Ukraine

Bielefeld

Allemagne

Cardiff

Grande-Bretagne

Armidale

Australie

Colombus

USA

Osnabriick

Allemagne

Maryland

USA

Cardiff

Grande-Bretagne

Victoria

Australie

Warrensburg

USA

Brisbane

Australie

Moscou

Russie

Marseille

France

Bale

Suisse

Plymouth

Angers

Canberra

Brest

Bremerhavcn

Wormlcy

Geneve

New Brunswick

Nantes

Messina

Canberra

Zurich

Oldenburg

Brisbane

Grahamstown

Puerto Rico

Massachusetts

Ashford

Tromsoe

Leiden

Brisbane

Budapest

Amsterdam

Sydney

Frankfurt

Solothurn

Plymouth

Washington

Oslo

Swansea

Chiba

Copenhague

Alaska

Achcvc d’imprimer cn novembre 1997 sur les presses dc I’lmprimerie de Montligeon

61400 La Chapelle Montligeon - D6p6t legal 4' trimestre 1997 - N° Imp. 18467 - Distribue le 28 novembre 1997

5 DEC. 1997

Source: MNHN. Paris

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This work presents a study of the taxonomy of the penaeidean shrimps of the world.

This group of shrimp includes many of the commercially important marine species of the

tropics and subtropics, as well as many of the species that constitute a major link in the food

chain of the oceans. The works present keys for the identification of seven families (the

Aristeidae, Benthesicymidae, Penaeidae, Sicyoniidae, Solenoceridae, Luciferidae, and Ser-

gestidae) and 56 genera, as well as diagnoses that attempt to define these taxa with scientific

clarity. Also included are lists of the species and subspecies currently included in these

genera, along with their geographic distribution, and a full bibliography.

Dr Isabel PEREZ Farfante is an Emeritus Research Scientist with the U.S. National

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Institution, Washington, D.C. She has published extensively on the systematics of penaeoid

shrimps of the world.

Dr Brian Kensley is a curator in the Department of Invertebrate Zoology, U.S. Natio¬

nal Museum of Natural History, Smithsonian Institution, Washington D.C. For eleven years,

prior to 1978, he was a Scientific Officer in the Department of Marine Biology at the South

African Museum, Cape Town. His principal research interests are in the systematics of iso¬

pod and decapod crustaceans of the world.

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