chapter 1 general introduction -...

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CHAPTER 1

GENERAL INTRODUCTION

Family Atyidae De Haan, 1849 under the order Decapoda Latreille, 1802 is

one of the species rich families in the infraorder Caridea Dana, 1852. The order

Decapoda comes under superorder Eucarida Calman (1904), subclass

Eumalacostraca Grobben (1892) and Class Malacostraca Latreille (1802).

The species coming under Class Malacostraca are characterized by:

crustaceans generally with a “5-8-6 (7)” body plan: 5 cephalic, 8 thoracic and 6

abdominal segments (7 somites in some), most of them bear paired appendages (7th

devoid of appendages); a pre-antennular region bearing compound eyes and a post-

anal region, the telson; the cephalic and thoracic somites covered by carapace;

antennule, antenna, mandible, maxillula and maxilla comprise cephalic appendages;

1st to 3rd maxillipeds, 1st to 5th pereiopods comprise thoracic; 1st to 5th pleopods and

uropods constitute abdominal appendages; male gonadial openings on the inner

aspect of coxa of 5th pereiopods whereas the female genital openings on the inner

region of coxa of 3rd pereiopods.

The Class Malacostraca Latreille (1802) is divided into three subclasses,

namely, Phyllocarida Packard, 1879, Hoplocarida Calman, 1904 and

Eumalacostraca Grobben, 1892. The Phyllocarida contains Nebaliids, Nebaliopsids

and Paranebaliids. The Hoplocarida includes Stomatopodids (Squilla).

Eumalacostraca is a large group containing diverse groups of organisms like

Bathynellaceans, Anaspidaceans, Spelaeogriphaceans, Thermosbaenaceans,

Lophogastrids, Mysidaceans, Mictaceans, Amphipods, Isopods, Tanaidaceans,

Cumaceans and Eucarids. Among these, Eucarids are highly advanced forms with 3

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orders, namely, Euphausiacea Dana, 1852, Amphionidacea Williamson, 1973 and

Decapoda Latreille, 1802. Since the crustaceans under investigation belong to the

order Decapoda, a widely accepted broad outline classification is given below in

order to get the exact taxonomic position of the family Atyidae (based on Martin &

Davis, 2001).

Order Decapoda Latreille, 1802

Suborder Dendrobranchiata Bate, 1888 (2 superfamilies & 7

families)

Suborder Pleocyemata Burkenroad, 1963

Infraorder Stenopodidea Claus, 1872 (2 families)

Infraorder Caridea Dana, 1852

Superfamily Procaridoidea Chace & Manning, 1972

Family Procarididae Chace & Manning, 1972

Superfamily Galatheacaridoidea Vereshchaka, 1997

Family Galatheacarididae Vereshchaka, 1997

Superfamily Pasiphaeoidea Dana, 1852

Family Pasiphaeidae Dana, 1852

Superfamily Oplophoroidea Dana, 1852

Family Oplophoridae Dana, 1852

Superfamily Atyoidea De Haan, 1849

Family Atyidae De Haan, 1849

Superfamily Bresilioidea Calman, 1896

Family Agostocarididae Hart & Manning, 1986

Alvinocarididae Christoffersen, 1986

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Bresiliidae Calman, 1896

Disciadidae Rathbun, 1902

Mirocarididae Vereshchaka, 1997

Superfamily Nematocarcinoidea Smith, 1884

Family Eugonatonotidae Chace, 1937

Nematocarcinidae Smith, 1884

Rhynchocinetidae Ortmann, 1890

Xiphocarididae Ortmann, 1895

Superfamily Psalidopodoidea Wood-Mason & Alcock,

1892

Family Psalidopodidae Wood-Mason & Alcock,

1892

Superfamily Stylodactyloidea Bate, 1888

Family Stylodactylidae Bate, 1888

Superfamily Campylonotoidea Sollaud, 1913

Family Bathypalaemonellidae de Saint Laurent,

1985

Campylonotidae Sollaud, 1913

Superfamily Palaemonoidea Rafinesque, 1815

Family Anchistioididae Borradaile, 1915

Desmocarididae Borredaile, 1915

Euryrhynchidae Holthuis, 1950

Gnathophyllidae Dana, 1852

Hymenoceridae Ortmann, 1890

Kakaducarididae Bruce, 1993

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Palaemonidae Rafinesque, 1815

Typhlocarididae Annandale & Kemp, 1913

Superfamily Alpheoidea Rafinesque, 1815

Family Alpheidae Rafinesque, 1815

Barbouriidae Christoffersen, 1987

Hippolytidae Dana, 1852

Ogyrididae Holthuis, 1955

Superfamily Processoidea Ortmann, 1890

Family Processidae Ortmann , 1890

Superfamily Pandaloidea Haworth, 1825

Family Pandalidae Haworth, 1825

Thalassocarididae Bate, 1888

Superfamily Physetocaridoidea Chace, 1940

Family Physetocarididae Chace, 1940

Superfamily Crangonoidea Haworth, 1825

Family Crangonidae Haworth, 1825

Glyphocrangonidae Smith, 1884

Infraorder Astacidea Latreille, 1802 (5 superfamilies & 7

families)

Infraorder Thalassinidea Latreille, 1831 (3 superfamilies & 11

families)

Infraorder Palinura Latreille, 1802 (2 superfamilies & 4

families)

Infraorder Anomura MacLeay, 1838 (4 superfamilies & 13

families)

Infraorder Brachyura Latreille, 1802 (2 sections, 24

superfamilies & 71 families)

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The order Decapoda Latreille, 1802 is characterized by: presence of

cephalothorax; carapace covers all thoracic somites and extends to the sides

covering gills in the branchial chamber as branchiostegite; exopodite of the 2nd

maxilla is a well developed scale; 1st to 3rd thoracic appendages are modified as

maxillipeds, and the rest are pereiopods, some of which may be chelate; pleopods

biramous; three kinds of branchiae present, namely, podobranchiae, arthrobranchiae

and pleurobranchiae; statocyst on the proximal most segment of the antennule.

It has 2 suborders viz., Dendrobranchiata Bate, 1888 and Pleocyemata

Burkenroad, 1963. The 2 suborders are distinguished based on the form of gill and

incubatory habit of females.

The suborder Dendrobranchiata Bate, 1888 is characterized by: presence of

dendrobranchiate gills; females are free spawners, releasing eggs into water and

hatch out as nauplius larva. Suborder Dendrobranchiata Bate, 1888 contains 2

subfamilies and 7 families. The characteristics of suborder Pleocyemata

Burkenroad, 1963 include Phyllobranchiate (caridean shrimps and brachyuran crabs

except dromiids) and trichobranchiate (lobsters and cray fishes) gills and incubatory

nature of females.

The suborder Pleocyemata Burkenroad, 1963 is further subdivided into 7

infraorders (see outline classification). The infraorder Caridea Dana, 1852 is

diagnosed by: abdomen generally with a sharp bent at the 3rd segment; pleurae of

2nd abdominal somite overlapping those of 1st and 3rd segments; mandibular palp, if

present, straight; third pereiopods never chelate; gills phyllobranchiate; eggs

attached to the pleopods of female.

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The infraorder Caridea Dana, 1852 contains 16 superfamilies, of which

Superfamily Atyoidea De Haan, 1849 is one among them and is characterized by:

first and second pairs of pereiopods with fingers of chelae usually terminating in

tufts of setae; second maxilliped with terminal segment broadly crescent shaped or

sub-semicircular, attached by curved margin to slender, sickle-shaped extension

from preceding segment; pereiopods with or without exopods; second pair of

pereiopods with carpus entire, not articulate; third maxilliped slender, pereiopod-

like, not broad or operculate, with exopod; first maxilliped with endite overreaching

exopod proper, exopod with lash, caridean lobe prominent; second maxilla with

distal endite incised near distal margin, palp not vestigial, scaphognathite narrowly

produced proximally; first maxilla with distal endite not unusually large, proximal

endite not reduced or absent; mandible without palp, incisor and molar processes

separated by sinuous, setose margin.

Atyidae De Haan, 1849 is the only family reported under the Superfamily

Atyoidea De Haan, 1849. The transfer of the Genus Xiphocaris Von Martens, 1872

to Xiphocarididae under the superfamily Nematocarcinoidea Smith, 1884 paved

way to establish Atyidae as a homogenous assemblage identifiable by anterior

pereiopods and mouthparts (Chace, 1992). Family Atyidae De Haan, 1849 is

characterized by: rostrum inflexibly attached to rest of carapace; carapace without

longitudinal lateral ridges or suture and without cardiac notch in posterior margin;

eyes well developed or reduced without pigments; antennule with 2 flagella, neither

with accessory branch; mandible without palp, with sub-truncate molar process not

distinctly separated from incisor process; pereiopods usually with strap-like epipods

on at least three anterior pairs or rarely absent; two anterior pairs of pereiopods

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similar, with fingers of chela usually terminating in tufts of setae; 2nd pereiopod

with carpus usually not articulated; pereiopods with or without exopods.

Holthuis (1986) divided the family Atyidae into 4 subfamilies namely

Atyinae De Haan, 1849; Caridellinae Holthuis, 1986; Paratyinae Holthuis, 1986

and Typhlatyinae Holthuis, 1986. However, recent studies by Sket & Zakšek

(2009) have synonymized the subfamily Typhlatyinae Holthuis, 1986 with

Paratyinae Holthuis, 1986 and hence resulted in three valid subfamilies which are:

Atyinae De Haan, 1849; Paratyinae Holthuis, 1986 and Caridellinae Holthuis,

1986.

The characters mostly used to separate genera are branchial complement,

presence or absence of exopods and extent of eye development. Both Chace (1992)

and Holthuis (1993) used the presence of exopods as distinguishing character to

separate genera in several families of caridean shrimps. The subfamilies coming

under the family Atyidae can be identified based on the key provided below.

Global distribution pattern of Atyidae is given in Fig. 1 and the symbols used to

represent genera are given on the left side of each genus for clarity, and legend to

symbols is appended along with the figure.

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1.1 Key for the identification of subfamilies under the family

Atyidae De Haan, 1849

1. Arthrobranchiae absent from at least the last four pereiopods, variable on 1st

pereiopod. Pereiopods with or without exopods. --------------------2

Pereiopods without exopods. First pereiopod with an arthrobranch. --------

Atyinae De Haan, 1849

2. Exopods present on some or all pereiopods. Eyes well developed or reduced

without or with a little pigment in the cornea. ----------------------------------------

Paratyinae Holthuis, 1986

Pereiopods without exopods (except Caridinides Calman, 1926) and

arthrobranchiae. Eyes generally well developed and with pigment. ---------------

Caridellinae Holthuis, 1986

1.2 Subfamily Paratyinae Holthuis, 1986

Paratyinae Holthuis, 1986, Zool. Med., 60 (7): 103

Synonym: Série Paratyienne Bouvier, 1925; Typhlatyinae Holthuis, 1986

Diagnosis:

• Supraorbital spines either present or absent

• Exopodites on some or all pereiopods.

• One or two lateral spines on uropodal diaeresis.

Remarks: As on the date 11 genera have been reported under the subfamily

Paratyinae Holthuis, 1986. Essential details of each genus, such as, diagnostic

characters, synonyms, type species, remarks and distribution are given below. This

subfamily has not been represented in Indian waters.

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1.2.1 Diagnosis of genera recorded under the subfamily Paratyinae

Holthuis, 1986

Genus: Atyaephyra De Brito Capello, 1867

Type species: Atyaephyra rosiana De Brito Capello, 1867

Gender: Feminine

Atyaephyra De Brito Capello, 1867, Mem. Acad. R. Sci. Lisboa, Sci. math. phys.

nat. (n. ser.), 4(1) (7): 5

Synonyms: Symethus Rafinesque, 1814; Hemicaridina Ortmann, 1890;

Symathus Rafinesque, 1815 (erroneous spelling); Acilius Rafinesque, 1815

(substitute name); Symaethus Agassiz, 1846 (invalid emendation); Atyephyra Von

Martens, 1868 (erroneous spelling); Atyephira Ishikawa, 1885 (erroneous spelling);

Lemicaridina Matzdorff, 1894 (erroneous spelling); Atyaëphyra Ortmann, 1895

(erroneous spelling); Athejaephira Magri, 1923 (erroneous spelling); Athejëphira

Magri, 1923 (erroneous spelling); Atyaephira Ferrer Galdiano, 1924 (erroneous

spelling); Atyaephrya Hertzog, 1930 (erroneous spelling); Athyaephyra Werner,

1938 (erroneous spelling); Atyalphyra Birstein, 1939 (erroneous spelling);

Athyaepora Sterk, 1950 (erroneous spelling) (Holthuis, 1955 for complete

synonymy).

Diagnosis:

• Eyes well developed, with pigment.

• Carapace without pterygostomian spine.

• Supraorbital spines present.

• Third maxilliped without arthrobranch.

• Pereiopods I, II with exopodites.

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Remarks: Atyaephyra is related to Dugastella Bouvier, 1912; Palaemonias

Hay, 1901; Paratya Miers, 1882; Syncaris Holmes, 1900; Troglocaris Dormitzer,

1853 and Gallocaris Sket & Zakšek, 2009 in the presence of a supraorbital spine on

the carapace. So far 3 species have been reported under this genus.

Distribution: Coimbra, Portugal; southern and western France- Corsica

Island, Sicily, Italy; Dalmatia, Croatia; Iran; western and northern Greece; Belgium;

Alabama, U.S.A.; Turkey and Czech Republic.

Genus: Dugastella Bouvier, 1912

Type species: Dugastella marocana Bouvier, 1912

Gender: Feminine

Dugastella Bouvier, 1912, C. R. Acad. Sci. Paris, 155: 993

Synonym: Nil

Diagnosis:

• Eyes well developed, with pigment.

• Carapace with supraorbital, suborbital and pterygostomian spines.

• Carpus of both first and second legs anteriorly hollowed.

• Pereiopods I to III or more with exopodites, 5th leg generally with a

rudimentary exopod.

Remarks: Resembles Antecaridina Edmondson, 1954; Palaemonias Hay,

1901; Stygiocaris Holthuis, 1960; Typhlatya Creaser, 1936; Typhlocaridina Liang

& Yan, 1981; Paracaridina Liang, Guo & Tang, 1999; Halocaridina Holthuis,

1963 and Halocaridinides Fujino & Shokita, 1975 in having a deep excavation in

the carpus of 1st and 2nd pereiopods. It also resembles Gallocaris Sket & Zakšek,

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2009 in having supraorbital and suborbital spines on the carapace. Two species

have been reported under this genus so far.

Distribution: Morocco and Spain.

P Genus: Palaemonias Hay, 1901

Type species: Palaemonias ganteri Hay, 1901

Gender: Masculine

Palaemonias Hay, 1901, Proc. Biol. Soc. Wash., 14: 179

Synonyms: Palemonias J. Roux, 1915 (erroneous spelling); Palaemonies

Giovannoli, 1933 (erroneous spelling); Poiotmonias Birstein, 1939 (erroneous

spelling); Pataemonias – Anonymous, 1904 (erroneous spelling); Paleomonetes

Poulson, 1964 (lapsus for palaemonetes).

Diagnosis:

• Eyes generally reduced, without pigment.

• Carapace with supraorbital and pterygostomian spines.

• Carpus of both first and second legs anteriorly hollowed.

• Exopods present on 1-4 pereiopods.

Remarks: Resembles Antecaridina Edmondson, 1954; Dugastella Bouvier,

1912; Stygiocaris Holthuis, 1960; Typhlatya Creaser, 1936; Typhlocaridina Liang

& Yan, 1981; Paracaridina Liang, Guo & Tang, 1999; Halocaridina Holthuis,

1963 and Halocaridinides Fujino & Shokita, 1975 in having a deep excavation in

the carpus of 1st and 2nd pereiopods. It shows similarities with Syncaris Holmes,

1900 in having supraorbital and pterygostomian spines on the carapace. Two

species have been reported under this genus.

Distribution: Kentucky and Alabama, U.S.A.

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Genus: Paratya Miers, 1882

Type species: Ephyra compressa De Haan, 1844

Gender: Feminine

Paratya Miers, 1882, Ann. Mag. nat. Hist., (5) 9: 194

Synonyms: Xiphocaridina Bouvier, 1909; Xiphatyoida J. Roux, 1915;

Xyphatyoida J. Roux, 1926 (erroneous spelling); Xiphicaridina Edmondson, 1935

(erroneous spelling).

Diagnosis:

• Eyes well developed, cornea pigmented.

• Carapace without pterygostomian spine, supraorbital spines present.

• All pereiopods with exopods, no arthrobranchiae on pereiopods.

• 3rd maxilliped with a single arthrobranch, epipods on 1-4 pereiopods.

Remarks: It shows similarity to Antecaridina Edmondson, 1954;

Stygiocaris Holthuis, 1960 and Typhlatya Creaser, 1936 in the presence of exopods

on all pereiopods. So far 15 species have been reported under this genus.

Distribution: Australia; eastern Siberia; Korea to the Lesser Sunda Islands

(southern Indonesia); New Zealand and nearby Islands; Ryukyu Islands and

southern Japan.

Genus: Syncaris Holmes, 1900

Type species: Miersia pacifica Holmes, 1895

Gender: Feminine

Syncaris Holmes, 1990, Occ. Pap. Calif. Acad. Sci., 7: 211

Synonym: Nil

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Diagnosis:

• Eyes well developed and pigmented.

• Rostrum long, slender with at least one ventral tooth; with or without

dorsal teeth and terminally appearing minutely bifid.

• Carapace with well developed supraorbital, antennal and

pterygostomian spines.

• Carpus of second legs without, that of first legs with an anterior

excavation.

• Carpus and merus of pereiopods 3-5 each with a pair of strong spines on

ventral or ventrolateral border.

• Exopods present on maxillipeds 2 and 3, variable on pereiopods 1-4,

always absent on pereiopod 5.

• Epipods present on pereiopods 1-4.

• Uropodal diaeresis weakly developed, terminating in one movable spine

and one immovable lateral point.

Remarks: Syncaris is related to Dugastella Bouvier, 1912; Palaemonias

Hay, 1901; Paratya Miers, 1882; Atyaephyra De Brito Capello, 1867; Troglocaris

Dormitzer, 1853 and Gallocaris Sket & Zakšek, 2009 in the presence of a

supraorbital spine on the carapace. Two species have been recorded under this

genus.

Distribution: California.

Genus: Troglocaris Dormitzer, 1853

Type species: Palaemon anophthalmus Kollar, 1848 (= Troglocaris schmidti

Dormitzer, 1853)

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Gender: Feminine

Troglocaris Dormitzer, 1853, Lotos Prague, 3: 85

Synonyms: Xiphocaridinella Sadovsky, 1930; Troglocaridella Babiě, 1930.

Diagnosis:

• Eyes greatly reduced, without pigment.

• Carapace without pterygostomian spine.

• Supraorbital spines present.

• Adult specimens with exopods on the first four or five legs. Third

maxilliped with arthrobranch.

• Appendix masculina having spines or setae much shorter than appendix

diameter.

Remarks: It shows affinities with Palaemonias Hay, 1901; Gallocaris Sket

& Zakšek, 2009; Antecaridina Edmondson, 1954; Stygiocaris Holthuis, 1960 and

Typhlatya Creaser, 1936 in having strongly reduced eyes with a little or without

pigmentation. Three species are present in this genus.

Distribution: Southern Europe from France and Italy to the vicinity of the

Caspian and Black Seas; Caves of Carniola; Caves of Kumpole and Gurk, southern

Slovania; Croatia; Bosnia and Herzegovina.

Genus: Gallocaris Sket & Zakšek, 2009

Type species: Troglocaris schmidti inermis Fage, 1937

Gender: Feminine

Gallocaris Sket & Zakšek, 2009, Zool. J. Linn. Soc., 2009: 786

Synonyms: Troglocaris schmidti inermis Fage, 1937; Troglocaris inermis

Holthuis, 1956.

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Diagnosis:

• Eyeless.

• Very short rostrum lacking teeth.

• Supra and suborbital spines present on the carapace.

• Maxilliped I exopodital lobe gradually narrowed distally, flagellum

short (shorter than width of lobe).

• Pereiopods I-III or more with exopodites.

• Dactylus of pereiopod V not comb-like.

• Males having appendix masculina shorter than appendix interna in

pleopod II.

Remarks: It is related to Antecaridina Edmondson, 1954 in having reduced

eyes and appendix interna of male 2nd pleopod longer than appendix masculina.

Only one species has been reported under this genus.

Distribution: Caves of southern France.

Genus: Antecaridina Edmondson, 1954

Type species: Mesocaris lauensis Edmondson, 1935

Gender: Feminine

Antecaridina Edmondson, 1954, Pacific Sci., 8: 368

Synonym: Mesocaris Edmondson, 1935

Diagnosis:

• Eyes reduced without or with a little pigment in the cornea.

• Rostrum unarmed.

• Pterygostomian and antennal spines present.

• Third maxilliped with 2 arthrobranchiae.

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• Epipods on 1st four pereiopods.

• 1st and 2nd pereiopods with carpus anteriorly excavated.

• Exopods present on all the legs; appendix interna of male 2nd pleopod

longer than appendix masculina.

• Live body colour usually bright red.

Remarks: This genus shows affinities with Dugastella Bouvier, 1912;

Palaemonias Hay, 1901; Stygiocaris Holthuis, 1960; Typhlatya Creaser, 1936;

Typhlocaridina Liang & Yan, 1981; Paracaridina Liang, Guo & Tang, 1999;

Halocaridina Holthuis, 1963 and Halocaridinides Fujino & Shokita, 1975 in

having a deep excavation in the carpus of 1st and 2nd pereiopods. It also resembles

Paratya Miers, 1882, Stygiocaris Holthuis, 1960 and Typhlatya Creaser, 1936 by

the presence of exopods on all pereiopods. Only one species has been reported.

Distribution: Hawaiian Islands; Ryukyu Islands, Kuroshima Island, Japan;

Lau Islands, Namuka Island, Wangava Island, Fiji; Europa Island; Sinai Peninsula;

Red Sea; Solomon Islands; Bohol, Philippines and Guam.

$ Genus: Stygiocaris Holthuis, 1960

Type species: Stygiocaris lancifera Holthuis, 1960

Gender: Masculine

Stygiocaris Holthuis, 1960, Crustaceana, 1: 47

Synonym: Nil

Diagnosis:

• Eyes strongly reduced, bullet shaped and without any pigment.

• Carapace without supraorbital spines.

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• 3rd maxilliped with a single arthrobranch.

• A single pleurobranch at the base of each of the pereiopods, which bear

no arthrobranchiae.

• Exopods on all pereiopods.

• The carpus of the 1st and 2nd pereiopods deeply excavated.

• The diaeresis of the uropodal exopod bears no spinules.

Remarks: It is closely related to Typhlopatsa Holthuis, 1956 from which it

differs in the presence of a podobranch on the 2nd maxilliped and in the presence of

an epipod on the 4th pereiopod. Two species have been reported.

Distribution: N. W. Australia.

Genus: Typhlatya Creaser, 1936

Type species: Typhlatya pearsei Creaser, 1936

Gender: Feminine

Typhlatya Creaser, 1936, Publ. Carnegie Inst. Wash., 457: 128

Synonyms: Typhlata Cárdenas F., 1950:157 (erroneous spelling); Typhatya

Monod, 1975 (erroneous spelling).

Diagnosis:

• Eyes reduced, with or without pigment in the cornea.

• Rostrum unarmed.

• Carapace without spines.

• 3rd maxilliped with a single arthrobranch.

• 1st to 4th pereiopods with epipods.

• 1st and 2nd pereiopods with carpus anteriorly excavated.

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• Exopods present on all the legs, that on 5th sometimes rudimentary or

absent.

Remarks: It shows affinities with Dugastella Bouvier, 1912; Palaemonias

Hay, 1901; Stygiocaris Holthuis, 1960; Antecaridina Edmondson, 1954;

Typhlocaridina Liang & Yan, 1981; Paracaridina Liang, Guo & Tang, 1999;

Halocaridina Holthuis, 1963 and Halocaridinides Fujino & Shokita, 1975 in having

a deep excavation in the carpus of 1st and 2nd pereiopods. It also resembles Paratya

Miers, 1882; Stygiocaris Holthuis, 1960 and Antecaridina Edmondson, 1954 in the

presence of exopods on all pereiopods. Seventeen species have been reported under

this genus.

Distribution: Anchialine caves around the Caribbean, Bermuda, Galapagos

Islands, Ascension Island; Spain; Mexico; Bahamas; Cuba; Dominican republic;

Honduras; Puerto Rica; Caico Islands and France.

T Genus: Typhlopatsa Holthuis, 1956

Type species: Typhlopatsa pauliani Holthuis, 1956

Gender: Masculine

Typhlopatsa Holthuis, 1956, Vie et Milieu, 7 (1): 43

Synonym: Nil

Diagnosis:

• Eyes strongly reduced, without pigment.

• No podobranch on the 2nd maxilliped.

• Third maxilliped with an arthrobranch.

• Epipods on 1st to 3rd pereiopods.

• Exopods on all pereiopods.

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• Carpus of 1st and 2nd pereiopods deeply excavated.

• Uropodal diaeresis bears no spinules.

Remarks: The genus shows affinities to Limnocaridella Bouvier, 1913;

Limnocaridina Calman, 1899; Halocaridina Holthuis, 1963 and Halocaridinides

Fujino & Shokita, 1975 in lacking a podobranch on the 2nd maxilliped. It is a

monotypic genus.

Distribution: S. W. Madagascar

1.3 Subfamily Atyinae De Haan, 1849

Atyinae Bouvier, 1925, Encycl. entomol. (A) 4: 1-370

Synonym: Série Caridinienne Bouvier, 1925

Diagnosis:

• Eyes generally well developed and with pigment.

• First pereiopod with an arthrobranch.

• Exopods absent on pereiopods.

Remarks: Subfamily Atyinae De Haan, 1849 was dealt by Bouvier under

the name Série Caridinienne. Later it became the nominate subfamily Atyinae De

Haan, 1849, as it contains the genus Atya Leach, 1816. Among the three subfamilies, only

the subfamily Atyinae De Haan, 1849 has been reported from India. So far 11

genera have been reported under this subfamily.

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1.3.1 Diagnosis of genera recorded under the subfamily Atyinae De

Haan, 1849

Ω Genus: Atya Leach, 1816

Type species: Atys scaber Leach, 1815

Gender: Feminine

Atya Leach, 1816, Suppl. 4th – 6th eds. Encycl. Brit., 1: 421

Synonyms: Atys Leach, 1815 (invalid junior homonym); Atia Latreille, 1817

(erroneous spelling); Atyoida Randall, 1839; Evatya Smith, 1871; Euatya Koelbel,

1884 (invalid emendation); Ortmannia Rathbun, 1902; Alya Bouvier, 1904

(erroneous spelling); Pseudatya J. Roux, 1928; Vanderbiltia Boone, 1935;

Orthmannia Sawaya, 1946 (erroneous spelling).

Diagnosis:

• 3rd maxilliped with 2 arthrobranchiae.

• 1st pereiopod with a single arthrobranch.

• 1st and 2nd pereiopods with chelae completely divided, lacking palm;

carpus of both appendages excavated distally, much shorter than broad.

• 3rd pereiopods are longer than the 4th and 5th pairs; merus of the 3rd

pereiopod is longer than the combined lengths of carpus, propodus and

dactylus (to base of claw).

• Pereiopods without exopods.

Remarks: The genus resembles Atyopsis Chace, 1983, Atyella Calman,

1906, Micratya Bouvier, 1913, and Australatya Chace, 1983 in having

monomorphic chela and an excavation in the carpus of 1st and 2nd pereiopods.

Thirteen species have been recorded under this genus.

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Distribution: Indonesia (Sumatra, Java, Bacan, Bali, Celebes, Saleyer,

Flores, Amboina, Timor, Halmahera); Samar, Philippine Islands; Caroline Islands;

Fiji Islands; New Calidonia; western Nicaragua; Hayti, West Indies; Cuba;

Republic of Trinidad and Tobago; Brazil; Baja California, Mexico to the north of

Peru and Republic of Colombia; Republic of Ecuador; Philippines; New Guinea;

Japan; Myanmar; Sri Lanka; Ghana; Republic of Cameroun; Democratic Republic

of Congo; Republic of Angola; Bolivia; Pacific coast of America (California,

Oregon, Washington, Alaska and Hawaii) and Martinique (eastern Caribbean Sea).

♥ Genus: Potimirim Holthuis, 1954

Type species: Caridina mexicana De Saussure, 1857

Gender: Feminine

Potimirim Holthuis, 1954, Zool. Verh. Leiden, 23: 2

Synonym: Nil

Diagnosis:

• Rostrum short, unarmed above, but with teeth below.

• Carpus of 1st pereiopod excavated anteriorly.

• Arthrobranch only at the base of the 1st pereiopod.

• Epipods at the bases of 1st to 3rd or 4th pereiopods.

• Palmar portion of chelae very small; dactylus of these chelae almost as

long as the propodus.


Remarks: With respect to the very small palmar portion of the chelae of 1st

and 2nd pereiopods Potimirim can be easily separated from other atyid genera. So

far four species have been reported worldwide.

22

Distribution: Jamaica; West Indies; Brazil; Cuba; Republic of Trinidad; El

Salvador; Nicaragua; north-eastern Mexico and Florida, U.S.A.

Genus: Atyoida Randall, 1840

Type species: Atyoida bisulcata Randall, 1840

Gender: Feminine

Atyoida Randall, 1840, J. Aca. Nat. Sci. Phil., 8: 106

Synonyms: Atyoidea Gibbes, 1850 (erroneous spelling); Ortmannia

Rathbun, 1901; Pseudatya Roux, 1928; Vanderbiltia Boone, 1935.

Diagnosis:

• 1st and 2nd pereiopds with chelae heteromorphic; carpus of both

appendages excavated anteriorly.

• Pereiopod 3 elongate, merus equal in length or shorter than combined

lengths of carpus, propodus and dactylus (to base of claw).


• Branchial complement consists of 5 pleurobranchiae, 3 arthrobranchiae,

1 podobranch, 5 epipods and 5 mastigobranchs.

• Endopod of 1st pleopod of male tapers gradually to form distally a

narrow appendix interna with retinacula; possibly protandrous.

Remarks: This genus is related to Australatya Chace, 1983 and Archaeatya

Villalobos, 1960 in having protandrous development. It also resembles Atya Leach,

1816; Atyopsis Chace, 1983; Australatya Chace, 1983; Typhlocaridina Liang &

Yan, 1981; Atyella Calman, 1906; Micratya Bouvier, 1913; Paracaridina Liang,

Guo & Tang, 1999; Halocaridina Holthuis, 1963 and Halocaridinides Fujino &

23

Shokita, 1975 in having an excavation in the carpus of 1st and 2nd pereiopods. Two


Distribution: Itajahy, Brazil; Oahu, Hawaiian Islands; Tahiti, eastern

Australia; New Zealand; Madagascar; Mauritius; Seychelles; Indonesia (Flores,

Timor, Sumatra); Philippines; New Guinea; Mariana Islands; Caroline Islands;

Solomon Islands; New Calidonia; Fiji Islands and French Polynesia (Mangareva,

Marquesas, Gambier Islands).

Genus: Atyopsis Chace, 1983

Type species: Atya spinipes Newport, 1847

Gender: Feminine

Atyopsis Chace, 1983, Smith. Contri. Zoo., No. 384: 1-54

Synonyms: Altya Roux, 1971 (erroneous spelling); Atyia Roux, 1932

(erroneous spelling).

Diagnosis:

• Rostrum without dorsal spines.

• Pterygostomian margin dentate or angular.

• Third maxilliped terminating in numerous stout setae, lacks the single

terminal spine.


• 1st and 2nd pereiopods with chelae monomorphic (without palm), carpus

of both appendages excavated distally.


1 podobranch, 5 epipods (reduced on 3rd and 4th pereiopods).

24

• 2nd pleopod of male with appendix masculina spinose over entire length

distal to base of appendix interna.

• Telson with posterolateral angles overreaching setigerous posterior

margin.

Remarks: The genus shows affinities with Atya Leach, 1816; Atyella

Calman, 1906; Micratya Bouvier, 1913 and Australatya Chace, 1983 in having

monomorphic chela and an excavation in the carpus of 1st and 2nd pereiopods. Two


Distribution: Fiji Islands; Philippines; Taiwan; Ryukyu Islands, Japan;

Samoa Islands; Sri Lanka; Indonesia (Sumatra, Java, Bali, Sulawesi, Borneo); New

Guinea; New Calidonia and Malasia.

Genus: Australatya Chace, 1983

Type species: Atya striolata McCulloch & McNeill, 1923

Gender: Feminine

Australatya Chace, 1983, Smith. Contri. Zool., No.384: 1-54

Synonyms: Atya McCulloch & McNeill, 1923; Atyoida Smith & Williams,

1982.

Diagnosis:

• Rostrum without dorsal spines; pterygostomian margin broadly

rounded.


• 1st and 2nd pereiopods with chelae monomorphic; carpus of both

appendages excavated distally.

25


1 podobranch, 5 epipods (reduced on 3rd and 4th pereiopods).

• 2nd pleopod of male with appendix masculina spinose on less than distal

third of length, possibly protandrous.

Remarks: It is related to Atyoida Randall, 1840 and Archaeatya Villalobos,

1960 in having protandrous development. It also resembles Atya Leach, 1816;

Atyopsis Chace, 1983; Atyoida Randall, 1840; Typhlocaridina Liang & Yan, 1981;

Atyella Calman, 1906; Micratya Bouvier, 1913; Paracaridina Liang, Guo & Tang,

1999; Halocaridina Holthuis, 1963 and Halocaridinides Fujino & Shokita, 1975 in

having an excavation in the carpus of 1st and 2nd pereiopods. Only one species has

been reported under this genus.

Distribution: Rivers and streams along the east coast of Australia.

Genus: Sinodina Liang & Cai, 1999

Type species: Caridina gregoriana Kemp, 1923

Gender: Feminine

Sinodina Liang & Cai, 1999, Raff. Bull. Zoo., 47 (2): 577-590

Synonym: Caridina Kemp, 1923

Diagnosis:

• One arthrobranch each on 3rd maxilliped and 1st pereiopod.

• 3rd and 4th pereiopods sexually dimorphic, male with propodus dilated

distally at flexor margin, armed with numerous small spinules, dactylus

long, about half as long as propodus, with 15-30 spines at flexor

margin; female with propodus not dilated, dactylus long, flexor margin

26

with 10-21 spines. 3rd pereiopod of male distinctly longer than that of

female.

• Podobranch only a single lamella.

• No exopods on pereiopods.

• Epipods on 3rd maxilliped and 1st to 4th pereiopods.

Remarks: With respect to the branchial complement this genus resembles

Caridina H. Milne Edwards, 1837 and Neocaridina Kubo, 1938. Twelve species

have been reported in this genus.

Distribution: Southern China.

Genus: Caridina H. Milne Edwards, 1837

Type species: Caridina typus H. Milne Edwards, 1837

Gender: Feminine

Caridina H. Milne Edwards, 1837, Hist. nat. Crust., 2: 362

Synonyms: Caradina Bate, 1863 (erroneous spelling); Carinida Filhol,

1886 (erroneous spelling); Caridine Urita, 1921 (erroneous spelling); Cardina

Hora, 1933 (erroneous spelling); Caradrina Carvalho, 1936 (erroneous spelling);

not Caridina Rathbun, 1906; Edmondson, 1929; Banner &Banner, 1960.

Diagnosis:

• Eyes well developed or reduced with a little pigment.

• Rostrum generally serrated on both the margins.

• Carapace with antennal spine, pterygostomian spine in a few species.

• Carpus of 2nd pair of legs longer than broad, generally not very deeply

excavated anteriorly.

27

• Palmar portion of chelae distinct, dactylus of these chelae much shorter

than the propodus.

• Epipods on 1st to 4th pereiopods.

• 3rd maxilliped with 2 arthrobranchiae.

• 1st pereiopod with a single arthrobranch.


Remarks: This is the only genus represented in India and is distributed

widely. So far 279 species (De Grave et al., 2009) have been reported worldwide

and from India 25 species have been recorded. During the present study 15 species

were collected and presented, of which three are new to science.

Distribution: Islands of Indian Ocean and Indo-Malaysia; Mauritius;

Seychelles; Indonesia (Saleyer, Celebes, Amboina, Flores, Sumatra, java); Loo-

choo islands; Commonwealth of Dominica; Japan (Bonin Islands, Ryukyu Islands);

Zulu Island, South Africa; Iran; east Africa to eastern Australia; Dar es Salaam,

Tanzania; Queensland; Egypt; Auckland, New Zealand; Sudan; Uganda;

Democratic Republic of Congo; Hong Kong; China; Taiwan; Fiji Islands; Santo

Island, Republic of Vanuatu; Sri Lanka; India; Myanmar; Singapore and Vietnam.

♣ Genus: Typhlocaridina Liang & Yan, 1981

Type species: Typhlocaridina lanceifrons Liang & Yan, 1981

Gender: Feminine

Typhlocaridina Liang & Yan, 1981, Acta. Zootaxon. Sin., 6 (1): 31

Synonym: Nil

Diagnosis:

• Eyes without pigment.

28

• Rostrum compressed, with dorsal teeth concentrated on anterior part of

carapace behind the orbit.

• Carpus of 1st and 2nd chelipeds deeply excavated anteriorly.

• No exopods on pereiopods.

• Endopod of male 1st pleopod ends in a slender appendix interna.

Remarks: This genus shows similarities with Caridinopsis Bouvier, 1912,

in common both the genera possess dorsal rostral teeth which are concentrated on

the proximal carapace behind the orbit. Three species have been recorded

worldwide.

Distribution: Guangxi, China.

N Genus: Neocaridina Kubo, 1938

Type species: Caridina denticulata sinensis Kemp, 1918

Gender: Feminine

Neocaridina Kubo, 1938, J. Imp. Fish. Inst. Tokyo, 33

Synonym: Caridina Kemp, 1918

Diagnosis:

• Endopod of male 1st pleopod pear-shaped or palm shaped.

• Appendix masculina of male 2nd pleopod dilated.

Remarks: Resembles Caridina H. Milne Edwards, 1837 in branchial

formula and general characteristics. So far 23 species have been reported under this

genus.

Distribution: Oahu, Hawaiian Islands; Korea; China; Ryukyu Islands, Japan

and Taiwan.

29

Genus: Jonga Hart, 1961

Type species: Ortmannia serrei, 1961; Bouvier, 1909

Gender: Feminine

Jonga Hart, 1961, Notulae Naturae (Philadelphia), 342: 3

Synonyms: Ortmannia Bouvier, 1909; Ortmannia Schmidt, 1935;

Potimirim Holthuis, 1951.

Diagnosis:

• Rostrum without dorsal teeth, ventral margin dentate but not finely

serrate.

• Orbital margin armed dorsally with row of denticles.


Remarks: The row of denticles on the dorsal rostral margin distinguishes it

from other atyid genera. So far only one species has been reported under this

genus.

Distribution: Jamaica.

Genus: Atydina Cai, 2010

Type species: Caridina atyoides, Nobili, 1900

Gender: Feminine

Atydina Cai, 2010, Zootaxa, 2372: 75

Synonym: Caridina atyoides, Nobili, 1900

Diagnosis:

• Robust 3rd and 4th pereiopods.

• Inflated cylindrical appendix masculina on the male second pleopod.

Remarks: Atydina is related to Atyopsis Chace, 1983 and Caridina H. Milne

Edwards, 1837. So far only one species has been reported under this genus.

30

Distribution: Indonesia.

1.4 Subfamily Caridellinae Holthuis, 1986

Caridellinae Holthuis, 1986, Zool. Med., 60 (7): 103

Synonym: Série Caridellienne Bouvier, 1925

Diagnosis:

• Eyes generally well developed and with pigment.

• Pereiopods without exopods and arthrobranchiae.

Remarks: So far 20 genera have been reported under this subfamily.

1.4.1 Diagnosis of genera recorded under the subfamily

Caridellinae Holthuis, 1986

A Genus: Atyella Calman, 1906

Type species: Atyella brevirostris Calman, 1906

Gender: Feminine

Atyella Calman, 1906, Proc. Zool. Soc. Lond., 1906 (1): 201

Synonym: Nil

Diagnosis:

• Anterolateral angle of basal segment of antennular peduncle with a

slender tooth, reaching midlength of 2nd segment. Several teeth of

dorsal rostral series placed behind the orbit.

• 3rd maxilliped with a rudimentary arthrobranch.

• Chelae monomorphic, carpus of the 1st and 2nd pereiopods anteriorly

excavated.


• Pleurobranchiae on somites bearing 4 anterior pairs of pereiopods.

• Epipods on 1st to 3rd pairs of pereiopods.

31

Remarks: The genus resembles Caridella Calman, 1906 in having a

rudimentary arthrobranch at the base of the 3rd maxilliped and pleurobranchiae only

on the 1st to 4th pereiopods. Three species have been reported under this genus.

Distribution: Lake Tanganyika, Zambia.

Genus: Caridella Calman, 1906

Type species: Caridella cunningtoni Calman, 1906

Gender: Feminine

Caridella Calman, 1906, Proc. Zool. Soc. Lond., 1906 (1): 198

Synonym: Nil

Diagnosis:

• A rudimentary arthrobranch at the base of the 3rd maxilliped.

• No pleurobranch on the 5th somite.

• Carpus of only the 1st pereiopod anteriorly excavated.

• Chelae with a distinct palmar portion.


Remarks: It shows similarities to Atyella Calman, 1906 in branchial

complement but differs in having a chela with a distinct palmar portion. Three

species are present under this genus.


♠ Genus: Caridinides Calman, 1926

Type species: Caridinides wilkinsi Calman, 1926

Gender: Masculine

Caridinides Calman, 1926, Ann. Mag. nat. Hist., (9) 17: 242

32

Synonym: Nil

Diagnosis:

• Exopod present at the base of the 1st pereiopod.

• Carpus of 2nd pair of legs longer than broad, generally not very deeply

excavated anteriorly.

• Epipods only on the 1st and 2nd pereiopods.

Remarks: It is the single genus in the subfamily Caridellinae having an

exopod on the 1st pereiopod. So far only one species has been reported under this

genus.

Distribution: Australia.

§ Genus: Caridinopsis Bouvier, 1912

Type species: Caridinopsis chevalieri Bouvier, 1912

Gender: Masculine

Caridinopsis Bouvier, 1912, Bull. Mus. Hist. nat. Paris, 18: 300

Synonym: Nil

Diagnosis:

• Rostrum rather long, but the dorsal spines concentrated behind the orbit,

no spines or teeth on the dorsal margin of the rostrum proper.

• Carpus of the 1st pereiopod not excavated anteriorly.

• Epipods present on 1st to 4th pereiopods.

• All pereiopods with pleurobranchiae; 3rd maxilliped with two

arthrobranchiae.


33

Remarks: Resembles Typhlocaridina Liang & Yan, 1981 in the position of

the rostral teeth, but can be distinguished by the branchial complement. Only one

species under this genus.

Distribution: French Guinea and West Africa.

Genus: Edoneus Holthuis, 1978

Type species: Edoneus atheatus Holthuis, 1978

Gender: Masculine

Edoneus Holthuis, 1978, Zoo. Mede. Uit. door het. Rij. van Nat. Hist. Lei.,

53 (19): 209

Synonym: Nil

Diagnosis:

• Eyes developed or reduced.

• Carapace without supraorbital or any other spines, pterygostomian

margin broadly rounded.

• No arthrobranch on pereiopods, no arthrobranch or a rudimentary

arthrobranch on 3rd maxilliped; podobranch vestigial or less developed.

• Epipods on 1-4 pereiopods; pleurobranchiae on all pereiopods.


• The outermost spine of uropodal diaeresis distinctly longer than lateral

angle.

Remarks: It is closely allied to Limnocaridella Bouvier, 1913 in its

branchial complement. Four species have been reported under this genus.

Distribution: Philippines.

34

≬ Genus: Lancaris Cai & Bahir, 2005

Type species: Caridina singhalensis Ortmann, 1894

Gender: Feminine

Lancaris Cai & Bahir, 2005, Raff. Bull. Zoo., 12: 39

Synonym: Caridina Ortmann, 1894

Diagnosis:

• Rostrum depressed, sharp, unarmed ventrally.

• Second maxilliped with a laminar shaped podobranch.

• 3rd maxilliped with an arthrobranch.

• Pereiopods without arthrobranchiae.


• Diaeresis with 22-35 movable spinules, lateral angle much shorter than

mesial spines.

• Distal margin of telson with intermediate pairs of setae much longer

than lateral pairs of spines.

Remarks: It shows similarities with Caridina H. Milne Edwards, 1837 but

can be differentiated by the branchial complement and the laminar shaped

podobranch on the 2nd maxilliped. This species is related to Parisia Holthuis, 1956

under the subfamily Caridellinae in having a vestigial podobranch and a single

arthrobranch on the 3rd maxilliped. Two species have been recorded under this

genus.

Distribution: Sri Lanka.

35

Limnocaridella Bouvier, 1913

Type species: Limnocaridina alberti Lenz, 1910

Gender: Feminine

Limnocaridella Bouvier, 1913, Bull. Soc. Entom. France, 1913: 180

Synonym: Nil

Diagnosis:

• Carpus of the 1st pereiopod not or not noticeably excavated anteriorly.

• A rudimentary arthrobranch present at the base of the 3rd maxilliped.


• Epipods present on 1st to 3rd pereiopods.

• Pleurobranch on the 5th leg generally absent; no podobranch on the 2nd

maxilliped.

Remarks: It shows similarities to Limnocaridina Calman, 1899 in lacking a

podobranch on the 2nd maxilliped and having pleurobranchiae only on the 1st to 4th

pereiopods. Only one species has been reported under this genus.

Distribution: Central Africa.

Genus: Limnocaridina Calman, 1899

Type species: Limnocaridina tanganyikae Calman, 1899

Gender: Feminine

Limnocaridina Calman, 1899, Proc. Zool. Soc. Lond., 1899: 704

Synonym: Nil

Diagnosis:

• No podobranch on the 2nd maxilliped.

• No arthrobranch on the 3rd maxilliped.

36

• Carpus of the 1st pereiopod not or not noticeably excavated anteriorly.


• No epipods on any of the pereiopods.

• Pleurobranch on the 5th leg generally absent.

Remarks: This genus resembles Limnocaridella Bouvier, 1913, but can be

distinguished by the absence of epipods on the pereiopods and arthrobranch on the

3rd maxilliped. So far eight species have been reported under this genus.


Z Genus: Marosina Cai & Ng, 2005

Type species: Marosina brevirostris Cai & Ng, 2005

Gender: Feminine

Marosina Cai & Ng, P.K.L., 2005, Crustaceana, 78 (2): 129

Synonym: Nil

Diagnosis:

• Eyes strongly reduced, without pigment.

• No arthrobranch at the base of the 1st pereiopod.

• Epipods present on 1-4 pereiopods.

• 3rd pereiopod terminating in one long claw, with one or two accessory

spines.

• Appendix interna of male 2nd pleopod reaching to end of appendix

masculina.

37

Remarks: It shows similarities with Parisia Holthuis, 1956 in the form of

endopod, telson and branchial formula. Two species have been reported under this

genus.

Distribution: Sulawesi, Indonesia.

U Genus: Micratya Bouvier, 1913

Type species: Atya poeyi Guérin Méneville, 1856

Gender: Feminine

Micratya Bouvier, 1913, Bull. Soc. Entom. France, 1913: 181

Synonyms: Calmania Bouvier, 1909 (invalid junior homonym of Calmania

Laurie, 1906); Balssiola Strand, 1922 (substitute name for Calmania Bouvier,

1909).

Diagnosis:

• All dorsal rostral teeth confined to the rostrum proper.

• Anterolateral angle of basal segment of antennular peduncle without

tooth.

• 3rd maxilliped with one well developed and one rudimentary

arthrobranch.

• Chelae monomorphic, carpus of the 1st and 2nd pereiopods excavated

anteriorly.


• Pleurobranchiae on somites bearing 4 anterior pairs of pereiopods,

mastigobranchs on 4 anterior pairs of pereiopods.

• Epipods on 1_ 3 pairs of pereiopods.

38

Remarks: It resembles Atyopsis Chace, 1983; Atyella Calman, 1906, Atya

Leach, 1861 and Australatya Chace, 1983 in having monomorphic chela and an

excavation in the carpus of 1st and 2nd pereiopods. So far only one species has been

recorded under this genus.

Distribution: Jamaica; Costa Rica and Panama.

β Genus: Archaeatya Villalobos, 1960

Type species: Archaeatya chacei Villalobos, 1960

Gender: Feminine

Archaeatya Villalobos, 1960, Ana. del Insti. de Biol. Uni. Nac. Auto. Mex.,

30 (1-2): 31

Synonym: Nil

Diagnosis:

• Rostrum without median dorsal spines.

• Pterygostomian margin of carapace rounded.

• Third maxilliped with 2 arthrobranchiae.

• Epipods on 4 anterior pairs of pereiopods, mastigobranchs on all 5

pereiopods, pleurobranchiae on all 5 somites bearing pereiopods.

• No exopods and arthrobranchiae on pereiopods.

• Propodus and carpus of 3rd pereiopod are densely covered by small,

strong spines.

• Endopod of 1st pleopod of male tapering from base to tip, possibly

protandrous.

39

Remarks: It is related to Australatya Chace, 1983 and Atyoida Randall,

1840 in having protandrous development. Only one species has been reported

under this genus.

Distribution: Costa Rica; Cocos Island; Perlas Archipelago and Gulf of

Panama.

s Genus: Puteonator Gurney, 1987

Type species: Puteonator iraqiensis Gurney, 1987

Gender: Masculine

Puteonator Gurney, 1987, Crustaceana, 53 (2): 160

Synonym: Nil

Diagnosis:

• Eyes reduced; rostrum short, unarmed.

• Antenna 2 with finger-like process arising from inner margin of basal

segment.

• 2 arthrobranchiae on 3rd maxilliped.

• Propodus of male pereiopods 3 and 4 widened distally with inner

margin convex and many marginal and sub-marginal spines.

• Pleurobranchiae present on all pereiopods.

• Epipods on all maxillipeds and 1-4 pereiopods.

• Appendix interna of male pleopod 1 arising at apex of endopod, with

retinacula at tip.

Remarks: With respect to branchial formula, Puteonator shows affinities

with Archaeatya Villalobos, 1960; Potimirim Holthuis, 1954 and Caridinopsis

Bouvier, 1912. So far 4 species have been recorded under this genus.

40

Distribution: Southern Iraq.

D Genus: Paracaridina Liang, Guo & Tang, 1999

Type species: Caridina longispina Guo et He, 1992

Gender: Feminine

Paracaridina Liang, Guo & Tang, 1999, J. Fish. China, 23: 69

Synonym: Caridina Liang & Yan, 1986

Diagnosis:

• Supraorbital spine absent.

• Two arthrobranchiae on 3rd maxilliped.

• No arthrobranch on the first pereiopod.

• 1st and 2nd pereiopods with carpus anteriorly excavated, fingers with

brushes of setae.

• Epipods on 1- 4 pereiopods.

Remarks: Resembles Caridina H. M. Edws., 1837 in general characteristics

but differs in lacking an arthrobranch in the 1st pereiopod. So far 5 species have

been recorded under this genus.

Distribution: Southern China.

Genus: Mancicaris Liang, Guo & Tang, 1999

Type species: Mancicaris sinensis Liang, Guo & Tang, 1999

Gender: Feminine

Mancicaris Liang, Guo & Tang, 1999, J. Fish. China, 23: 69

Synonym: Nil

41

Diagnosis:

• Eyes without pigment.

• Supraorbital spine absent.

• Exopod of the 1st maxilliped has no flagellum.

• No arthrobranch on the first pereiopod, two on the 3rd maxilliped.


Remarks: The absence of flagellum in the exopod of the 1st maxilliped

distinguishes it from other atyid genera. So far only one species has been reported.

Distribution: China.

Genus: Parisia Holthuis, 1956

Type species: Parisia macrophthalma Holthuis, 1956

Gender: Feminine

Parisia Holthuis, 1956, Vie et Milieu 7: 43

Synonym: Nil

Diagnosis:

• Eyes sometimes reduced, if reduced with or without a small pigment

spot.

• Carapace without supraorbital or pterygostomian spines.

• 2nd maxilliped with a vestigial podobranch.

• 3rd maxilliped with a distinct arthrobranch and sometimes with

rudiments of a second.



Remarks: Eight species have been reported under this genus so far.

42

Distribution: Australia; Madagascar; Zanzibar Island; Tanzania; Luzon and

Philippines.

Ψ Genus: Pycneus Holthuis, 1986

Type species: Pycneus morsitans, 1986

Gender: Masculine

Pycneus Holthuis, 1986, Zool. Med., 60 (7): 103

Synonym: Nil

Diagnosis:

• Eyes degenerated, without pigment.

• Rostrum very short, wider than long, depressed, unarmed.

• Carapace without spines.

• 3rd maxilliped with a small arthrobranch.

• Pleurobranchiae on 1-4 pereiopods.



• Fingers of chelipeds with sharp hooked tips and relatively few hairs.

Remarks: Resembles Atyella Calman, 1906; Caridella Calman, 1906;

Halocaridina Holthuis, 1963; Halocaridinides Fujino & Shokita, 1975;

Limnocaridella Bouvier, 1913 and Limnocaridina Calman, 1899 in having

pleurobranchiae at the base of the 1st to 4th pereiopods. Only one species has been

reported under this genus.

Distribution: Western Australia.

43

Genus: Halocaridina Holthuis, 1963

Type species: Halocaridina rubra Holthuis, 1963

Gender: Masculine

Halocaridina Holthuis, 1963, Zool. Med., 38 (16): 261

Synonyms: Caridina Rathbun, 1906; Caridina Edmondson, 1929; Caridina

Banner & Banner, 1960.

Diagnosis:

• Eyes reduced, with pigment.

• Rostrum unarmed, carapace without spines.

• Pleurobranchiae on pereiopods 1 to 4, no other branchiae present.


• Carpus of 1st and 2nd chelipeds deeply excavated anteriorly.

• Exopods absent from all pereiopods.

• 1st pleopod of male ending in a narrow appendix interna.

• Diaeresis with 1 or 2 spines.

Remarks: In the reduced number of branchiae Halocaridina resembles

Limnocaridina Calman, 1899; Limnocaridella Bouvier, 1913; Atyella Calman, 1906

and Caridella Calman, 1906. Two species have been recorded under this genus.

Distribution: Hawaiian Islands.

H Genus: Halocaridinides Fujino & Shokita, 1975

Type species: Halocaridina (Halocaridinides) trigonophthalma Fujino & Shokita,

1975

Gender: Masculine

Halocaridinides Fujino & Shokita, 1975, Bull. Sci. Eng. Div., Uni. Ryukyu, 18: 93

44

Synonyms: Palauatya Hart, 1980; Caridina Rathbun, 1906; Caridina

Edmondson, 1929; Caridina Banner & Banner, 1960.

Diagnosis:

• Cornea very small, pigmented.

• Rostrum unarmed, carapace without spines.

• Pleurobranchiae on pereiopods 1 to 4, no other branchiae present.

• Epipods on 1-3 pereiopods; exopods absent on pereiopods.

• Carpus of 1st and 2nd pereiopods excavated anteriorly.

• No appendix interna on the endopod of male 1st pleopod.

• Exhibits sexual dimorphism, in males propodus of the 3rd and 4th legs

widened distally and forms a subchela with the dactylus.

Remarks: Fujino & Shokita created a subgenus Halocaridinides under the

genus Halocaridina. Later Holthuis (1982) revised the subgenus Halocaridinides

and elevated to the genus level. He has also synonymized the genus Palauatya

Hart, 1980 to this genus. Two species have been recorded under this genus.

Distribution: Ryukyu Islands, Okinawa Island, Japan; Palau district, West

Caroline Islands; Guam and Philippines.

W Genus: Pycnisia Bruce, 1992

Type species: Pycnisia raptor Bruce, 1992

Gender: Feminine

Pycnisia Bruce, 1992, Invertebrate Taxonomy, 6 (3): 553

Synonym: Nil

Diagnosis:

• Eyes small with reduced pigmentation.

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• Rostrum extremely small, acute, much wider than long.

• 3rd and 4th pereiopods developed into massive raptorial appendages.

• Exopodites absent from all pereiopods.

Remarks: Two species have been reported worldwide.

Distribution: Australia.

∞ Genus: Jolivetya Cals, 1986

Type species: Jolivetya foresti Cals, 1986

Gender: Feminine

Jolivetya Cals, 1986, P. Acad. Sci., 303(10): 387

Synonym: Nil

Diagnosis:

• Eyes degenerated and non-pigmented.

Remarks: Only one species has been reported.

Distribution: Australia and Madagascar.

Delclosia Rabadá, 1993

Delclosia, the only fossil genus under the family Atyidae, has been reported

from Spain. So far two species namely, Delclosia martinelli Rabadá, 1993 &

Delclosia roselli (Via, 1971) have been reported.

1.5 Global distribution pattern of genera under the Family Atyidae

The global distribution pattern of the 43 genera discussed above are shown

in Fig. 1. It reveals that wide range of distribution has been noticed in Caridina and

Atya and narrow distributional range has been reported in the cases of Pycnisia,

Halocaridina, Pycneus, Mancicaris, Paracaridina, Puteonator, Marosina,

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Limnocaridina, Limnocaridella, Lancaris, Edoneus, Caridinopsis, Caridinides,

Caridella, Atyella, Atydina, Jonga, Typhlocaridina, Sinodina, Australatya,

Typhlopatsa, Stygiocaris, Gallocaris and Syncaris. With respect to species richness

Caridina contains the maximum number of species whereas Gallocaris,

Antecaridina, Typhlopatsa, Australatya, Jonga, Atydina, Caridinides, Caridinopsis,

Limnocaridella, Micratya, Archaeatya, Mancicaris, Pycneus and Jolivetya are

monotypic genera. Maximum congregation of genera has been reported between

30°N and 30°S and the distributional range extends up to 47°N and 47°S.

Legend to Symbols

Global distribution pattern of genera under the family Atyidae

1 Atyaephyra 23 A Atyella

2 Dugastella 24 Caridella

3 P Palaemonias 25 ♠ Caridinides

4 Paratya 26 § Caridinopsis

5 Syncaris 27 Edoneus

6 Troglocaris 28 ≬ Lancaris

7 Gallocaris 29 Limnocaridella

8 Antecaridina 30 Limnocaridina

9 $ Stygiocaris 31 Z Marosina

10 Typhlatya 32 U Micratya

11 T Typhlopatsa 33 β Archaeatya

12 Ω Atya 34 S Puteonator

13 ♥ Potimirim 35 D Paracaridina

14 Atyoida 36 Mancicaris

15 Atyopsis 37 Parisia

16 Australatya 38 Ψ Pycneus

17 Sinodina 39 Halocaridina

18 Caridina 40 H Halocaridinides

19 ♣ Typhlocaridina 41 W Pycnisia

20 N Neocaridina 42 ∞ Jolivetya

21 Jonga 43 Delclosia

22 Atydina

chapter 1 general introduction -...

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