Abbreviated larval development of Macrobrachium hainanense (Parisi, 1919) reared in the laboratory (Decapoda, Caridea, Palaemonidae)

ABBREVIATED LARVAL DEVELOPMENTOF MACROBRACHIUM TUXTLAENSE VILLALOBOS amp ALVAREZ 1999

REARED IN THE LABORATORY

BY

FERNANDO ALVAREZ JOSEacute LUIS VILLALOBOS and RAFAEL ROBLES

Coleccioacuten Nacional de Crustaacuteceos Instituto de Biologiacutea Universidad Nacional Autoacutenomade Meacutexico Apartado Postal 70-153 Mexico 04510 DF Mexico

ABSTRACT

The abbreviated larval development of the freshwater prawn Macrobrachium tuxtlaense isdescribed The rst stage of M tuxtlaense is similar to those of other Mexican and South Americanspecies of Macrobrachium such as M vicconi M reyesi and M nattereri sharing uniramouspereiopods biramous pleopods and a rounded telson Newly hatched M tuxtlaense undergo 5moults before reaching the juvenile stage Abbreviated development has allowed M tuxtlaense toget established in Lake Catemaco Veracruz Mexico where no other of the common coastal plainspecies of Macrobrachium occur

RESUMEN

Se describe el desarrollo larval abreviado del langostino Macrobrachium tuxtlaense La primeraetapa larval de M tuxtlaense es similar a aquellas de otras especies mexicanas y sudamericanasde Macrobrachium como M vicconi M reyesi y M nattereri con quien comparte pereioacutepodosunirraacutemeos pleopodos birraacutemeos y un telson redondeado Los M tuxtlaense recien eclosionadospasan por 5 mudas antes de alcanzar la etapa juvenil El desarrollo abreviado le ha permitido aM tuxtlaense establecerse en el Lago de Catemaco Veracruz Meacutexico en donde no se encuentraninguna otra de las especies de Macrobrachium que son comunes en la planicie costera

INTRODUCTION

The diversity of reproductive strategies found in the species of the genus Ma-crobrachium is a key attribute of their success in invading tropical and temperatecoastal systems around the world Among these abbreviated development has al-lowed many species to colonize freshwater habitats at considerable distances fromthe sea Abbreviated development however is a general term that encompasses avariety of patterns of larval development (Jalihal et al 1993)copy Koninklijke Brill NV Leiden 2002 Crustaceana 75 (5) 717-730Also available online wwwbrillnl

718 F ALVAREZ J L VILLALOBOS amp R ROBLES

Until recently species of Macrobrachium with abbreviated development on theAmerican continent were known only from South America (Rodriacuteguez 1982Pereira amp Garciacutea 1995) Recent ndings including the description of two newspecies suggest that this type of development is also widespread in southernMexico (Villalobos amp Alvarez 1999 Romaacuten et al 2000 Signoret et al 2000)In this paper we present the larval development of M tuxtlaense Villalobos ampAlvarez 1999 reared in the laboratory

MATERIALS AND METHODS

Ovigerous Macrobrachium tuxtlaense were collected near the type locality ofthe species in a small tributary of Lake Catemaco 1 km south from Las MargaritasVeracruz (Villalobos amp Alvarez 1999) All material studied was collected inMarch 1997 Captured individuals were transported to the laboratory at theInstitute of Biology UNAM in Mexico City in 20 l plastic containers withconstant aeration Once in the laboratory all organisms were placed in 40 l aquariaat 25plusmnC

The process of egg maturation was constantly monitored The eggs graduallychanged from black to transparent over a two-month period When the eyes inthe embryos were noticeable the egg-carrying female was placed in a 500 mljar inside the aquarium closed with nylon mesh and with constant aeration untilthe larvae hatched After hatching the larvae were placed individually in 45 mljars in the same way as the parental female The larvae were fed daily ad libitumwith a ne powder obtained from commercial shrimp pellet food Every jar wasexamined daily all exuvia were removed and xed in 70 alcohol and laterdissected to illustrate each stage Cephalothorax (CL) and total (TL) lengths wereobtained from each exuvium using a dissecting microscope Five to 10 exuvia weremeasured for each stage

Following Pereira amp Garciacutea (1995) we consider that an organism has reachedthe juvenile stage once the telson has adopted its nal shape and has the completeset of spines

RESULTS

The size of ovigerous females was 35 sect 2 mm TL (N D 15) mean sizeof recently spawned eggs was 153 pound 223 mm while that of mature eggs was156 pound 25 mm The hatching process lasted from 1 day for a clutch of 4 to 4days for a clutch of 15 Fecundity was unrelated to female size varying between2 and 20 eggs Three clutches of M tuxtlaense were followed through eight

ABBREVIATED DEVELOPMENT MACROBRACHIUM TUXTLAENSE 719

Fig 1 Numbers of eggs (E) and specimens in successive stages (S1-S7) from one female ofMacrobrachium tuxtlaense Alvarez amp Villalobos 1999 Day 1 marks start of hatching

stages in one case for 47 days ( g 1) and in the other two for 41 and 42 daysrespectively Stages 1 and 2 lasted for three days stage 3 six days stage 4 eightdays stages 5-6 nine days and stage 7 lasted for at least 9 days Stage 1 larvaedo not feed feeding starts during stage 2 In all stages studied the organismsremained immobile on the bottom of the jars for most of the time only reactingwhen disturbed or fed with quick avoidance movements

Stage 1 ( g 2)

Size mdash CL 22 mm range 21-25 mm TL 70 mm range 67-77 mmRostrum mdash Long slender distal tip sharp oriented downwards dorsal and

ventral margins smooth ( g 2a)Carapace mdash Shorter than abdomen integument membranous with small

antennal spine on anterior margin ( g 2a) Spot of vitellum visible in dorsal viewin gastric region

Abdomen mdash Somites 2-5 of approximately same length rst somite abouthalf this length First two somites with ventral border of pleura broadly roundedposterior angle with setae posterior four somites with ventral border formingsubacute angle ( g 2a)

Telson mdash Rounded posterior margin with 32-34 plumose setae ( g 2j)Uropods mdash Not developedEyes mdash Sessile ocular peduncle short as wide as the corneaAntennule mdash Peduncle not articulated inner agellum short half or less than

half length of the external one with thick apical seta ( g 2h) Outer agellumcomposed of a long segment with four apical setae


Fig 2 Macrobrachium tuxtlaense Alvarez amp Villalobos 1999 Stage 1 larva a lateral view(pereiopods omitted) b mandible c maxillule d maxilla e rst maxilliped f second maxillipedg third maxilliped h antennule i antenna j telson k rst pereiopod l second pereiopod m thirdpereiopod n fourth pereiopod o fth pereiopod p rst pleopod q second pleopod Scale bars

indicate 1 mm


Antenna mdash Biramous endopodite with three basal segments Inner and distalmargins of scaphocerite with setae becoming longer distally lateral margin withrobust acute distal spine not reaching distal margin Basicerite with small acutespine located ventrally next to anterior border Flagellum longer than body withdistal tuft of setae ( g 2i)

Mandible mdash Without palp molar and incisor processes fused distal teethabsent ( g 2b)

Maxillule mdash Endites and palp well developed devoid of setae anterior enditerobust longer than posterior one ( g 2c)

Maxilla mdash With two endites divided by cleft as deep as their length Palpslender with rounded tip endites and palp devoid of setae Scaphognathite welldeveloped bordered by 42 long plumose setae anterior lobe wide roundedposterior lobe slender ( g 2d)

First maxilliped mdash With two endites divided by shallow notch devoid of setaeon gnathal border anterior endite 15 times as long as posterior one ( g 2e)Exopodite slender four times as long as palp apex with four long plumose setaecaridean lobe discernible bordered by four long plumose setae Palp robust apexrounded Epipodite single-lobed elongated reaching beyond base of exopodite

Second maxilliped mdash Endopodite slender ( g 2f) Exopodite longer andslimmer than endopodite bearing four long apical setae Epipodite slender shortwith rounded apex

Third maxilliped mdash Endopodite slender 3-segmented middle segment shortest( g 2g) Exopodite reaching distal third of second segment of endopodite bearingfour long plumose apical setae Epipodite at shorter and wider than that ofsecond maxilliped

Pereiopods 1-2 mdash Chelate similar in shape rst one shorter chelae non-functional cutting edges smooth ( g 2k-l)

Pereiopods 3-5 mdash Simple slender with scattered setae and some tufts in thearticulations dactyli elongated ending in sharp tip ( g 2m-o)

Pleopods mdash Biramous well developed non-functional second to fth pairswith internal ramus well developed First pair with reduced endopodite Non-functional

Stage 2 ( g 3)

Size mdash CL 257 mm range 22-26 mm TL 73 mm range 72-76 mmRostrum mdash Slender oriented downwards ending in acute tip reaching beyond

rst antennular segment ( g 3a) Dorsal margin with seven teeth regularlydistributed along rostral length ventral margin smooth

Carapace mdash Shorter than abdomen integument membranous inferior orbitalangle subacute antennal spine small ( g 3a)


Fig 3 Macrobrachium tuxtlaense Alvarez amp Villalobos 1999 Stage 2 larva a lateral view(pereiopods and pleopods omitted) b mandible c maxillule d maxilla e rst maxilliped f secondmaxilliped g third maxilliped h antennule i antenna j telson k rst pereiopod l second

pereiopod m third pereiopod n fourth pereiopod o fth pereiopod Scale bars indicate 1 mm


Abdomen mdash Relative lengths of somites unchanged ( g 3a) Ventral border ofpleuron of second somite broadly rounded posterior angle of pleura of somites 3-5subacute with tufts of setae pleuron of sixth somite ending posteriorly in smallspine

Telson mdash Rounded posterior margin bearing 32-35 plumose setae ( g 3j)Uropods mdash Not developedEyes mdash With articulated peduncleAntennule mdash Peduncle 3-segmented basal segment longer than other two

segments combined ( g 3h) Basal segment with triangular spine on distolateralangle and small spine on ventral surface Stylocerite with acute apex Inner agellum short 3-segmented distal segment with four setae Outer agellumwith two branches beyond second segment inferior branch 1-segmented superiorbranch 3-segmented both branches bearing apical setae

Antenna mdash Unchanged ( g 3i)Mandible mdash Without palp molar and incisor processes well developed Incisor

process with small triangular teeth ( g 3b) Molar process with three low roundedprojections

Maxillule mdash Endites and palp devoid of setae ( g 3c) Anterior enditerobust with minute spines along gnathal border longer than posterior enditePosterior endite tapering distally Palp thick with rounded apical projectionoriented mesially

Maxilla mdash Endites devoid of setae separated by deep cleft anterior one longerpalp slender distal margin rounded with single seta on anterior margin ( g 3d)Scaphognathite same as in previous stage

First maxilliped mdash Endites divided by shallow cleft devoid of setae on gnathalborder anterior one 15 times as long as posterior one Exopodite slender 3 timesas long as palp apex with four long plumose setae caridean lobe well formedapex with ve setae Palp elongated apex rounded Epipodite divided by shallowcleft anterior lobe tapering distally reaching beyond base of exopodite posteriorlobe short with rounded apex ( g 3e)

Second maxilliped mdash Endopodite elongated 3-segmented distal segmentscurved mesially with few setae ( g 3f) Exopodite and endopodite as in previousstage

Third maxilliped mdash Endopodite elongated 4-segmented with scattered tufts ofsetae proximal segment longest Exopodite thinner longer than proximal segmentof endopodite with ve long plumose apical setae Epipodite blade-like shortrounded ( g 3g)

Pereiopods 1-2 mdash Chelate similar in shape different in size rst one shorterchelae functional with short setae on ngers Merus-carpus articulation with twotufts of short setae ( g 3k-l)


Pereiopods 3-5 mdash Unchanged ( g 3m-o)Pleopods mdash Functional well developed biramous First pair with small

endopodite appendix interna absent Second to fth pairs with well developedendopodite shorter than exopodite both rami bordered with long setae appendixinterna present

Stage 3 ( g 4)

Size mdash CL 285 mm range 27-31 mm TL 72 mm range 69-81 mmRostrum mdash Slender oriented slightly downwards ending in sharp tip reaching

second antennular segment Dorsal margin with 7-8 small teeth regularly distrib-uted in proximal 45 distal 15 smooth one long seta in spaces between teethVentral margin smooth ( g 4a)

Carapace mdash Shorter than abdomen integument more rigid than in previousstages suborbital angle with antennal spine Small hepatic spine below antennalspine reaching beyond margin of carapace ( g 4a)

Abdomen mdash Relative lengths of somites unchanged Ventral margin of pleuronof second somite broadly rounded devoid of setae ( g 4a) Pleura of fourth and fth somites with posterior margin subacute bearing pair of setae Pleuron of sixthsomite with posterior spine

Telson mdash Lateral margins slightly convergent posterior third becoming widerposterior margin rounded with 36 plumose setae ( g 4j) Dorsal surface with tworows of setae along central portion lateral margins with small movable spine onposterolateral angle

Uropods mdash Fully developed both rami similar in shape with internal andposterior margins bearing plumose setae Exopod longer and larger movable spineon posterolateral angle ( g 4j)

Eyes mdash UnchangedAntennule mdash Peduncle 3-segmented basal segment longer than other two seg-

ments combined with distolateral angle projected as a sharp spine small spine onventral surface next to internal margin Stylocerite with subacute apex Flagellaof same length inner 5-segmented with apical tuft of setae outer divided be-yond second segment inferior branch with single segment bearing setae api-cally superior branch 4-segmented distal segment with apical tuft of short setae( g 4h)

Antenna mdash Unchanged ( g 4i)Mandible mdash Without palp incisor process with three sharp teeth molar process

with four sharp projections ( g 4b)Maxillule mdash Endites and palp well developed Anterior endite longer than

posterior one bearing spines on gnathal border Posterior endite slender tapering





distally with setae on distal margin Palp robust apex mesially oriented ending intwo rounded projections distal one longer ( g 4c)

Maxilla mdash Unchanged except for appearance of apical setae on endites( g 4d)

First maxilliped mdash As in the previous stage except for appearance of shortthick setae on the gnathal border of the anterior endite ( g 4e)

Second maxilliped mdash Endopodite 5-segmented distal two segments wide andcurved mesially distal segment with short setae on gnathal border Exopoditeslender longer than endopodite with ve apical long plumose setae Epipoditeshort rounded ( g 4f)

Third maxilliped mdash Endopodite 3-segmented proximal segment the longest rst two segments with scattered setae third segment with ve rows and an apicaltuft of setae Exopodite shorter than proximal segment of endopodite bearing fourlong apical setae Epipodite attened short rounded ( g 4g)

Pereiopods mdash Unchanged ( g 4k-o)Pleopods mdash Unchanged

Stage 4 ( g 5)

Similar to stage 3 except for the following charactersSize mdash CL 31 mm range 30-33 mm TL 77 mm range 76-80 mmRostrum mdash Slender slightly oriented downwards reaching middle portion of

rst antennular segment Dorsal margin with 7-9 small teeth regularly distributedin proximal 45 of its length distal 15 unarmed single seta in each space betweenteeth Ventral margin with one small sharp tooth opposite to rst dorsal tooth( g 5a)

Carapace mdash Shorter than abdomen integument rigid suborbital margin withantennal spine Hepatic spine small below antennal spine reaching beyond marginof carapace ( g 5a)

Abdomen mdash Relative lengths of somites unchanged Ventral margin of pleuronof second somite broadly rounded devoid of setae Pleura of third and fth somiteswith posterior margin subacute bearing a pair of setae pleuron of fourth somitewith posterior margin ovoid pleuron of sixth somite ending in spine on posteriorventral angle ( g 5a)

Telson mdash Approximately rectangular lateral margins parallel posterior marginrounded with 15-19 plumose setae Dorsal surface with two rows of setae alongcentral portion posterior third with two pairs of spines on lateral margins Twopairs of spines on distolateral angles inner pair longer ( g 5j)

Uropods mdash Fully developed both rami similar in shape with internal andposterior margins bearing plumose setae Exopod longer and larger two pairs




of spines on posterolateral angle outer pair small non-articulated inner pairarticulated twice as long Lateral margin of protocerite ending in sharp spine( g 5j)





Stage 5 ( g 6)

Similar to stages 3 and 4 except for the following charactersSize mdash CL 33 mm range 34-33 TL 85 mm range 85-86 mmAntennule mdash Unchanged except for inner agellum 7-segmented external

agellum divided beyond third segment inferior branch 3-segmented superiorbranch 5-segmented ( g 6h)

Stages 6 7 and 8

The few changes that appear in these stages are described belowSize mdash Stage 6 CL 36 mm range 36 mm TL 92 mm range 91-93 mm

Stage 7 CL 38 mm range 37-40 mm TL 952 mm range 91-105 mm Stage 8CL 41 mm range 41-43 mm TL 112 mm range 105-117 mm

Antennule mdash Inner agellum with 7 11 and 12 segments in stages 6 7 and 8respectively Superior branch of external agellum with 9 11 and 13 segments instages 6 7 and 8 respectively

Second maxilliped mdash Epipodite progressively becoming a podobranch throughstages 6-8

DISCUSSION

According to Jalihal et al (1993) the larval development of Macrobrachiumtuxtlaense corresponds to a Type II partially abbreviated development withall pereiopods uniramous well developed pleopods and a fan-shaped telson instage 1 Other characteristics noted by Jalihal et al (1993) for species with thiskind of development (eg medium body size distribution in upper reaches ofrivers and absence of marked sexual dimorphism) t M tuxtlaense well The larvalcharacteristics of M tuxtlaense suggest a closer relationship with the MexicanM vicconi Romaacuten Ortega amp Mejiacutea 2000 and the South American M nattereri(Heller 1862) and M reyesi Pereira amp Garciacutea 1995 rather than with species fromother geographical areas These species have a morphologically similar rst stagewith uniramous pereiopods biramous pleopods and rounded telson

Regarding size the eggs of M tuxtlaense are similar to those of several speciesincluding M vicconi M reyesi M aracamuni Rodriacuteguez 1982 M ferreiraiKensley amp Walker 1982 and M nattereri However the mean and maximum totallengths of the rst stage of M tuxtlaense are larger than in the rest of the Americanspecies The duration of the larval phase is dif cult to compare among speciesdue to the potential in uence of the experimental conditions namely food qualityand temperature (Pereira amp Garciacutea 1995) Macrobrachium tuxtlaense reaches thejuvenile stage after 5 moults that take 25 days


Abbreviated development independent from brackish water has allowedM tuxtlaense to become established in Lake Catemaco at an altitude of 330 mabove sea level Below Lake Catemaco are several waterfalls and a dam that ef-fectively separate the lake fauna from the coastal plain fauna where at least vespecies of Macrobrachium (M acanthurus (Wiegmann 1836) M carcinus (Lin-naeus 1758) M heterochirus (Wiegmann 1836) M olfersii (Wiegmann 1836)and M hobbsi Nates amp Villalobos 1990) with extended larval development occur

ACKNOWLEDGEMENTS

We thank Rolando Mendoza for preparing the gures and the students at theColeccioacuten Nacional de Crustaacuteceos Instituto de Biologiacutea UNAM for their help

REFERENCES

JALIHAL D R K N SANKOLLI amp S SHENOY 1993 Evolution of larval developmental patternsand the process of freshwaterization in the prawn genus Macrobrachium Bate 1868 (DecapodaPalaemonidae) Crustaceana 65 365-376

PEREIRA G amp J V GARCIacuteA 1995 Larval development of Macrobrachium reyesi Pereira(Decapoda Palaemonidae) with a discussion on the origin of abbreviated development inpalaemonids Journ Crust Biol 15 117-133

RODRIGUEZ G 1982 Fresh-water shrimps (Crustacea Decapoda Natantia) of the Orinoco Basinand the Venezuelan Guayana Journ Crust Biol 2 378-391

ROMAacuteN R A L ORTEGA amp L M MEJIacuteA 2000 Macrobrachium vicconi new species afresh-water shrimp from a rain forest in southeast Mexico and comparison with congeners(Decapoda Palaemonidae) Journ Crust Biol 20 186-194

SIGNORET G A L ORTEGA amp D BRAILOVSKY 2000 Partially abbreviated larval developmentin an undescribed freshwater palaemonid prawn of the genus Macrobrachium from ChiapasMexico Crustaceana 73 273-282

VILLALOBOS J L amp F ALVAREZ 1999 A new species of Macrobrachium (Crustacea DecapodaPalaemonidae) with abbreviated development from Veracruz Mexico Proc biol Soc Wash-ington 112 746-753

First received 25 September 2001Final version accepted 4 February 2002


Until recently species of Macrobrachium with abbreviated development on theAmerican continent were known only from South America (Rodriacuteguez 1982Pereira amp Garciacutea 1995) Recent ndings including the description of two newspecies suggest that this type of development is also widespread in southernMexico (Villalobos amp Alvarez 1999 Romaacuten et al 2000 Signoret et al 2000)In this paper we present the larval development of M tuxtlaense Villalobos ampAlvarez 1999 reared in the laboratory

MATERIALS AND METHODS

Ovigerous Macrobrachium tuxtlaense were collected near the type locality ofthe species in a small tributary of Lake Catemaco 1 km south from Las MargaritasVeracruz (Villalobos amp Alvarez 1999) All material studied was collected inMarch 1997 Captured individuals were transported to the laboratory at theInstitute of Biology UNAM in Mexico City in 20 l plastic containers withconstant aeration Once in the laboratory all organisms were placed in 40 l aquariaat 25plusmnC

The process of egg maturation was constantly monitored The eggs graduallychanged from black to transparent over a two-month period When the eyes inthe embryos were noticeable the egg-carrying female was placed in a 500 mljar inside the aquarium closed with nylon mesh and with constant aeration untilthe larvae hatched After hatching the larvae were placed individually in 45 mljars in the same way as the parental female The larvae were fed daily ad libitumwith a ne powder obtained from commercial shrimp pellet food Every jar wasexamined daily all exuvia were removed and xed in 70 alcohol and laterdissected to illustrate each stage Cephalothorax (CL) and total (TL) lengths wereobtained from each exuvium using a dissecting microscope Five to 10 exuvia weremeasured for each stage

Following Pereira amp Garciacutea (1995) we consider that an organism has reachedthe juvenile stage once the telson has adopted its nal shape and has the completeset of spines

RESULTS

The size of ovigerous females was 35 sect 2 mm TL (N D 15) mean sizeof recently spawned eggs was 153 pound 223 mm while that of mature eggs was156 pound 25 mm The hatching process lasted from 1 day for a clutch of 4 to 4days for a clutch of 15 Fecundity was unrelated to female size varying between2 and 20 eggs Three clutches of M tuxtlaense were followed through eight




Stage 1 ( g 2)









indicate 1 mm












Stage 2 ( g 3)






















Stage 3 ( g 4)






















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES











Stage 1 ( g 2)









indicate 1 mm












Stage 2 ( g 3)






















Stage 3 ( g 4)






















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES










indicate 1 mm












Stage 2 ( g 3)






















Stage 3 ( g 4)






















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES



















Stage 2 ( g 3)






















Stage 3 ( g 4)






















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES


























Stage 3 ( g 4)






















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES























Stage 3 ( g 4)






















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES











Stage 3 ( g 4)






















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES


















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES















Stage 4 ( g 5)















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES
















Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES












Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES









Stage 5 ( g 6)



Stages 6 7 and 8





DISCUSSION





ACKNOWLEDGEMENTS


REFERENCES










ACKNOWLEDGEMENTS


REFERENCES








Abbreviated larval development of Macrobrachium hainanense (Parisi, 1919) reared in the laboratory (Decapoda, Caridea, Palaemonidae)

Documents