Some Biological Characteristics of Tadpole Shrimp, Triops Cancriformis, from Seasonal Pools of West Azarbaijan (Iran)

J. Agric. Sci. Technol. (2009) Vol. 11: 81-90

Some Biological Characteristics of Tadpole Shrimp, Triops cancriformis, from Seasonal Pools of West Azarbaijan (Iran)

A. Golzari 1, S. Khodabandeh 1*, and J. Seyfabadi 1

ABSTRACT

The tadpole shrimp of genus Triops is a well-known living f ossil whose f undamental morphology has been unchanged f or 220 million years. W e collected specimens of Triops cancriformis in temporary water bodies near the southern part of Urmia Lake (in the Fall of 2005). Some biological characteristics of this Triops were investigated. The f eeding regime of T. cancriformis was found to be related to the f auna and flora of the temporary pools. Invertebrates and animal detritus were found to constitute major part of the feeding regime. The existence of Triops cysts and particles in the gut also showed certain degree of cannibalism. Morphological and histological investigations showed that the population of T. cancriformis was f emale and there was only one male among 400 samples collected. Observation of sperm among follicle ducts of a few samples indicated some degree of hermaphrodity, but the animal seemed to reproduce mainly through parthenogenesis. Fecundity, varying from 100 to 2500 cysts, was with a f ew exceptions related to the body size. The average cyst diameter was 400 ±±± 85 µµµ m.

Keywords : Crustacean, Feeding Regime, Notostraca, Reproduction, Shrimp, Tadpole Triops cancriformis .

INTRODUCTION Petrov, 1999). Tadpole shrimps are charac- terized by their wide fluctuations in popula- Notostracans are freshwater crustaceans tion density. When their density becomes adapted to temporary water bodies (Su and very high, they can barely find enough food, and they become very active in searching for Mulla, 2001). They are commonly called tadpole shrimps because of their superficial food. Food shortage inhibits their develop- resemblance to frog larvae. They are recog- ment and reduces fecundity, while mortality increases as a result of cannibalistic behav- nizable by their large, horseshoe–shaped dorsal carapace (Martin and Boyce, 2005). ior (Scholnick and Snyder, 1996). Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021 They are benthic branchiopods which swim The tadpole shrimp genus Triops is a well- known living fossil whose fundamental and feed ventral surface down, burrowing into the bottom sediments in search of detri- morphology has remained unchanged for tus and a variety of small organisms (Schol- 220 million years. While the order has wide geographical distribution, many species have nick and Snyder, 1996). Their feeding de- pends on their habitats. Being facultative a restricted local distribution (Suno–Uchi et detritus feeders or scavengers or predators, al., 1997). The ability of Triops to develop and grow they eat algae, bacteria, protozoa, rotifers, earthworms, insects, fairy shrimps, frog eggs quickly is crucial to its success (Davis and Madison, 1999). Fecundity is related to oxy- and also tadpoles (Cvetkovic-Milicic and ______1 Department of Marine Biology, Faculty of Marine Sciences, Tarbiat Modares University, Noor, P.O. Box: 46414-356, Islamic Republic of Iran. * Corresponding author, Surp78@g mail.com

81 ______Golzari et al.

gen tension and temperature and so small tini, 1980; Scanabissi et al ., 2005). changes in temperature or oxygen dramati- Although some scattered networks on this cally change the total number of eggs laid in organism- under its general name “ Apus ”- one season (Scholnick, 1995). have been traced to fish culture ponds in The females possess two ovisacs (brood Iran (Kheirandish, 1975; Kohnehshahri and pouches) on the 11 th appendages. The two Takami, 1974), no specific work on its iden- ovisacs hold an almost equal number of tification and biological characteristics, par- eggs. Eggs are laid randomly and not in ticularly feeding and reproduction, have clusters, although all the eggs in both brood been conducted. In this work, therefore, ef- pouches are released simultaneously (Sea- forts are concentrated towards precise identi- man et al., 1991). The fecundity of T. can- fication, and study of the feeding and repro- criformis is high; individuals may lay more ductive biology of this organisms in sea- than 1000 eggs (Fry and Mulla, 1996). Their sonal ponds of West Azarbaijan Province. eggs lie dormant in the soil and can survive as such for many years. They are resistant to MATERIALS AND METHODS severe drought and extreme temperatures (Takahashi, 1997). It has been suggested that the distribution of many branchiopods Animals results from the transfer of resting eggs, either by birds, people or cattle and sheep. The first adult specimens of T. cancri- Most eggs hatch within a few days of being formis were collected in the Fall of 2003 submerged, but some remain unhatched in from vernal pools of West Azarbaijan Prov- the soil. This is an effective way for the ince in Iran (Figure 1A) (Khodabandeh et shrimps to maintain their populations under al. , 2008). For biological studies 400 speci- unpredictable fluctuations in environmental mens (length = 2-5 cm) were collected dur- conditions (Takahashi, 1997). ing the Spring and Fall of 2005. 95 speci- 15 species of Triops have been reported in mens were fixed for 24 hours in Bouin’s the world. Triops cancriformis inhabits tem- fixative and alcohol 70% separately for porary pools and rice fields in Eurasia and feeding regime and reproduction studies. Africa (Cesari et al ., 2004; Cvetkovic- Milicic and Petrov, 1999). T. cancriformis distribution in Europe has been reported by Feeding Regime Zaffagnini and Trentini (1980). While the order has a wide geographical distribution, Study of the feeding regime was accom- many of the species have a restricted local plished by investigating the gut contents of distribution (Takahashi, 1997). 50 specimens under the microscope, using Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021 T. cancriformis populations reproduce Rose Bengal staining and histological meth- gonochorically, hermaphroditically or even ods. The Triops, gut was separated by scal- parthenogenetically (Zaffagnini and Tren- pel under a stereo microscope, and its con- tini, 1980; Cesari et al ., 2004). They com- tents were preserved in alcohol 70% and prise bisexual populations, with an equal then studied under a light microscope. Rose male: female sex ratio or with a female bias, Bengal (0.5 g in 500 cc distilled water) was and unisexual populations, either hermaph- added to the contents of the gut to distin- roditic or parthenogenetic (Mantovani et al ., guish the vegetal or animal origins of the 2004). The more northerly European popula- food. tions are parthenogenetic, while incidents of For classic histological studies, the animals gonochorism begin appearing towards the were fixed for 24 hours in Bouin’s fixative. southern latitudes, with the northern African The specimens were then fully dehydrated in populations presenting both sexes and am- a graded ethanol series and embedded in phigonic reproduction (Zaffagnini and Tren- paraffin. Sections (5µm) were cut on a mi-

82 Biology of Triops cancrifor mis ______

crotome, collected on albumine-glycerine 1D-F). The anterior dorsal portion of the slides and stained with Haematoxylin, Eosin carapace is equipped with a relatively and Methyl green (Martoja and Martoja- unique optical arrangement (Figure 1E). The Pierson, 1967; Khodabandeh et al ., 2005a trunk consists of a thorax bearing append- and b). ages and an abdomen without them. The oral region of the cephalon possesses specific Reproduction appendages including antennules, antennae, maxillae and mandibles that are covered by a labrum and used for food handling. The Morphological observation and a histo- anterior portion of the thorax consists of 11 logical method were used for sex differentia- segments, each bearing a pair of appendages tion and identification of reproductive sys- (thoracopods) (Figure 1F). The 11 th append- tem structure. Fecundity was determined by ages of females form ovisacs (brood counting the cysts in the ovisacs and gonads pouches) (Figure1F). The many appendages of 45 specimens. Biometry of the cysts was posterior to the 11 th move the spent feeding accomplished by light microscope with a and respiratory current away from the body. micrometer. The diameters of 12 cysts from The proximal portion of the abdomen pos- each specimen were measured, and the aver- sesses 65 pairs of fine, hair-like appendages age, maximum and minimum diameter was that beat in a rhythmic fashion to assist in calculated. movement and food channeling. The distal portion of the abdomen terminates in a RESULTS prominent telson and subsequently branches into two large caudal furca (Figures 1D-F). Stereo microscopic studies revealed that Triops cancriformis is reported from the gut of T. cancriformis was filled with throughout the southern Palaearctic region, microscopic vegetal, animal, and unknown and from vernal pools of West Azarbaijan particles (Figures 2A-H). Invertebrates and Province in Iran (Khodabandeh et al ., 2008). animal detritus were found to constitute a Morphologically the material is a mixture of major part of the T. cancriformis feeding specimens bearing the carapace carinal spine regime. The existence of Triops cysts and arrangements of both T. c. cancriformis and Triops particles in the guts also showed T. c. simplex subspecies, casting doubts as to some degree of cannibalism (Figures 2H and the validity of the subspecific designations. 2D). Application of Rose Bengal showed Specimens appeared with the filling of that the vegetal detritus did not have any ditches and vernal pools by rain and irriga- color and animal detritus were pink color tion. They were also found in almost drained (not shown). Histological studies showed Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021 out ditches (Figure 1B). During the course that in the stained sections the vegetal detri- of sampling the pond had conductivity val- tus was stained green by Methyl Green and -1 ues ranging between 450-860 µS cm animal detritus was stained red by Fuchsin (22°C) and a pH ranging from 7.5 and 8.5. (Figures 3A-C). The gut of Triops in the The highest population of T. cancriformis pool with flora and without flora was more occurred in ponds that had little aquatic green and red, respectively (Figure 3A). vegetation but supported other invertebrates Morphological sex determination showed (cladocerans, hemipterans, coleopterans) and that most of the population of Triops were had relatively high conductivity (860 µS cm- female and there was only one male among 1) (Figure 1C). 400 collected samples. The paired gonads Morphological investigation shows that the extend almost the entire length of the trunk body (total length= 2-5 cm) consists of a on either side of the gut in females (Figure head and a trunk mostly covered by a large 3E). These gonads were filled by numerous (length =1-2 cm) dorsal carapace (Figures oocytes and cysts that occupied the body

83 ______Golzari et al.

* Urmia Lake

1B 1C

Th F-th C FG

Ov Ov A

F 1E 1F

Figure 1. Map of Iran Star showing collection lo cation : ( 1A) S ampling ditches and ve rn al pools; (1B and 1C) Triops cancriformis ; (1D) Dorsal; (1E) Ventral, (1 F) View of Triops

Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021 cancriformis . Abbreviations: A= Abdo men; C= Carapace; CE= Compound eyes; F= Furca; FG= Food groove; F-th= First thoracopod; Ov = Ovisac, Th= Thoracopods. Scale bars: 1 C (15 cm); 1D, 1E and 1 F (1 cm). Arrows (s ampl es).

cavity together with the gut (Figure 3E). cle duct cells on average are 32 µm high and Each gonad consists of a germarium at the 10 µm wide (Figure 3H). They possess tip of several follicles whose ducts join the rather short, scattered microvilli. The ellipti- longitudinal oviduct and, at the end, leading cal nucleus was nearly central. The cyto- to the ovisac (Figure 3D). Each follicle con- plasm of the follicle duct cells seem to se- sists of an oocyte and three nurse cells (Fig- crete eggshell material (Figure 3H). At the ures 3D and 3F). The follicle duct wall is a end of vitellogenesis the oocytes descend to single layer of closely packed cells lying on the follicle duct, then the ripe oocytes pass the basal lamina (Figures 3F and 3G). Folli- along the follicle ducts and enter the longi-

84 Biology of Triops cancrifor mis ______

100 X 100 X 100 X

2A 2B 2C

100X 400X

2D 2E

400 X 300 X 300 X

2F 2G 2H

Figure 2 . (2A-C) So me v eget al particles , ( 2D and 2E) Animal particles and cysts fro m Triops cancriformis gut. Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021

tudinal oviduct to reach the ovisac (Figure present in a tubular and complicated repro- 3D). The number of follicles and oocytes ductive system on either side of the gut, depended on the size and habitat of Triops, which at the end were carried in two ovisacs and were different even in two gonads of the (Figure 3D). Fecundity, that with a few ex- same Triops. Some samples did not have ceptions was found to be related to the body any cysts in their ovisacs, but their gonads size, varied from 100 to 2500; the average were full of cysts. In their ovisacs were cyst diameter was 400 ±85 µm. brownish, spherical and separated cysts. In In the follicle ducts of a few females several their gonads some cysts were milk-white in sperm cells and also degenerated spermato- color and others were brown. The cysts were zoa were observed (Figures not shown).

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3AGo 3B 3C

FD O F G Gu Go

3E LO NC FD

EM EF O Ov O FD

3D 3F

FD O LOE O

EM EM 3G 3H

Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021 Figure 3. Histological stained sections: (3A-C) T ransversal sections of trunk o f Triops cancriformis ; (3B and 3C) Detritus into the gut; (3D) Schematic graph o f a gonad and ovisac, (3E-H) Gonad sections. Abbreviations: EF= Empty follicles; EM= Eggshell materials; F= Follicles; FD Follicle ducts; G= Germariu m; Go= Gonad; Gu= Gut; LO= Longitudinal oviduct; LOE= Longi- tudinal oviduct epithelium; Oo= Oo cyte, Ov = Ovisac. Scale bars: 3A (1mm); 3B and 3 C (400 µ m); 3E (250 µm); 3F and 3G (25 µ m), 3H (3 µm).

DISCUSSION microscopic vegetal and unknown particles. Their feeding regime was found to be related Adult Triops (length 2-5 cm) were used for to the fauna and flora of the temporary biological studies. The gut of T. cancri- pools. The vegetal or animal origin of the formis was filled with small invertebrate and detritus depended on the abundance of floral and faunal forms in the pools. Invertebrate

86 Biology of Triops cancrifor mis ______

and animal detritus were found to constitute tubules and their number varies throughout a major part of T. cancriformis feeding re- the population. In addition, the activity of gime. Some Triops particles in the guts also the testicular vesicles is obviously low when showed a certain degree of cannibalism. compared with those of the bisexual popula- Taylor et al . (1987) have suggested that tions. They have never seen sperm cells in Triops is omnivorous and feeds mainly on their specimens, which leads them to ex- algae, bacteria, protozoa, rotifers and detri- clude any possibility of self-fertilization. tus. Tietze and Mulla (1989) have reported Also, an ultrastructural analysis of the gonad that T. cancriformis progress from juveniles of males of T. cancriformis in Austria has feeding on phytoplankton to mature adults shown a truly functional structure, present- feeding on plant material and that only the ing morphofunctional characteristics typical largest individuals ≥ 10 mm are carnivorous; of normal gametic development and matura- they have also suggested that T. longicauda- tion (Scanabissi et al ., 2005). In T. cancri- tus was polyphagous and found to feed upon formis Engelmann et al . (1997) found signs invertebrates and plant material in both day- of sperm degeneration. light and darkness. They also noted that, in We observed that the female reproductive the laboratory, early instar mosquito larvae apparatus consisted of two gonads lying were consumed. In a more detailed study in ventrolaterally in the haemocoel and filling the laboratory, this species was found to be a the body cavity along with the gut. Each fe- predator of mosquito larvae ( Culex quinque- male gonad consists of a germarium at the fasciatus ) and is currently being considered tip of several follicles whose ducts join the as a biological control against mosquitoes longitudinal oviduct. These results are in (Tietze and Mulla, 1989). agreement with the results of previous stud- Our investigation showed that the popula- ies (Trentini and Scanabiss, 1978 and 1982). tion of Triops were almost exclusively fe- The ovarian follicles consisted of an oo- male and there was only one male among cyte and three nurse cells. These oocyte fol- 400 collected samples. Observation of sperm licles were surrounded by a thin layer of fol- among the follicle ducts of a few females licle cells. The oocytes were initially smaller indicated some degree of hermaphrodity, but than nurse cells. The cytoplasm of oocytes the demonstration of a high amount of de- was filled with differently sized yolk glob- generated spermatozoa in the gonads of ules that were surrounded by membranes of hermaphroditic specimens is contrary to the endoplasmic reticulum. The nurse cells hermaphroditic reproduction and the animals had yolk globules too and they were charac- seemed to reproduce mainly through parthe- terized by slightly darker cytoplasm. This nogenesis. The presence of male T. cancri- was also in agreement with the results of

Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021 formis had already been reported in Hungary previous studies (Trentini and Scanabiss, (Abonyi, 1926), in Yugoslavia (Petrov and 1978 and 1982; Engelmann et al ., 1997). Cvetkovic, 1996 and 1999) and in Austria The follicle duct wall is a single layer of (Scanabissi et al ., 2005). Trentini and Sca- closely packed cells lying on the basal lam- nabissi (1982) have suggested that the sex ina. The oocyte has no coating at the begin- ratio of T. cancriformis in Italy is heavily ning of its descent but pushes the shell mate- biased in favor of the females, which display rial towards the end of the tubules into the testicular lobes scattered along their follicle oviduct where it accumulates. The egg cov- ducts. Such populations have been consid- ering begins to form as the egg passes ered to be hermaphroditic by some authors, through this material before entering the while other authors consider them to be longitudinal oviduct or uterus. The egg coat- parthenogenetic with rudimentary hermaph- ing is uniform and becomes the definitive roditism. Whenever testicular lobes are pre- eggshell only after a vacuolization takes sent, they lie along some female gonadal place within it. It is already reported that, when the eggs are finally deposited in the

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ovigerous pockets, they display the charac- REFERENCES teristic shell structure (Trentini and Scana- biss, 1978 and1982; Tommasini et al ., 1. Abonyi, A. 1926. The Mmales of Apus cac- 1989). riformis Schaeffer Described on the Basis of In our study fecundity was, with a few ex- Specimens Collected in the Region of Lake ceptions, related to the body size and means Balaton Biological Station of Reofulop. that the larger specimens have more cysts; Arch . Balatonicum ., 1: 71-90. fecundity varided from 100 to 2500 cysts. 2. Cesari, M., Mularoni, L., Sabelli-Scanabissi, Fry and Mulla (1996) reported that the fe- F. and Mantovani, B. 2004. Characterization cundity of T. cancriformis was high: indi- of Dinucleotide Microsatellite Loci in the Living Fossil Tadpole Shrimp Triops cancri- viduals may lay more than 1000 eggs. formis (Crustacea Branchiopoda Notostraca). Scholnick (1995) found a positive linear re- Molec. Ecol. Notes ., 4: 733-735. lationship between the number of eggs and 3. Cvetkovic-Milicic, D. and Petrov, B. 1999. wet mass for T. longicaudatus ; Seaman et al . Life Histories of Triops cancriformis (Bosc, (1991) have also reported a similar relation- 1801) and Lepidurus apus (Linnaeus, 1758) ship between carapace length and fecundity (Crustacea, Notostraca) in a Group of Rain- for T. granarius. Su and Mulla (2001) have pool in the Banat Province in Yugoslavia. reported a positive correlation between any Crustacean Issues , 12 : The Biodiversity Cri- two parameters of growth (CL at Death), sis and Crustacea , Vol 2, Rotterdam, PP. longevity and egg production. Longer sur- 411-417, 4. Damgaard, J. and Olesen, J. 1998. Distribu- viving T. newbrryi with a larger body size at tion, Phenology and Status for the Larger death produced more eggs during their life- Branchiopoda (Crustacea: Anostraca, Noto- time. This relationship is also true for other straca, Spinicaudata and Laevicaudata) in branchiopod species. We observed that the Denmark. Hydrobiologia , 377 : 9-13. average cyst diameter in the ovisacs of T. 5. Davis, J. M. and Madison, D. 1999. The cancriformis was not significantly different. Ontogeny of Light-dark Response in Triops Cyst diameter was not found to be related longicaudatus as a Response to Changing Selective Pressures. Crustaceana , 73(3): body size. Some of the cysts in the gonads were milky-white and the other were brown. 283-288. 6. Engelman, M., Hahn, M. and Hoheisel, G. Previous investigations have shown that T. 1997. Ultrastructural Characterization of the granarius has two types of eggs, one Gonads of Triops cancriformis (Crustacea, drought resistant, the other not (Seaman et Notostraca) from Populations Containing al ., 1991). Trentini and Scanabissi (1982) Both Females and Males: No Evidence have found that the cyst shell of T. cancrifor Hermaphroditic Reproduction. Zoomor- formis is subdivided into two layers: an phology , 117 : 175-180. 7. Fry, L. L. and Mulla, M. S. 1996. Optimal

Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021 outer uniform and compact one about 3 µm thick (cortex) and an inner alveolar layer Conditions for Rearing the Tadpole Shrimp, Triops cancriformis (Notostraca: which is about 25 µm thick. The chitinous Triopsidae), a Biological Control Agent embryonic cuticle lies deep to it. against Mosquitoes . J. Am. Mosq. Control. Assoc ., 12(3):446-453.

ACKNOWLEDGEMENTS 8. Khodabandeh, S., Charmantier, G., Blasco, C., Grousset, E. and Charmantier-Daures, M. 2005a. Ontogeny of the Antennal Gland in This study was supported by Tarbiat Mo- the Crayfish Astacus leptodactylus (Crusta- dares University, Iran. Dr. Christopher cea, Decapoda): Anatomical and Cell Differ- Rogers from Laboratory of Invertebrate entiation. Cell Tissue Res ., 319 : 153-165 Ecologist/Taxonomist (Woodland, CA, 9. Khodabandeh, S., Charmantier, G. and Charmantier-Daures, M. 2005b. Ultrastruc- USA) is thanked for the specimen’s identifi- tural Studies and Na +,K +-ATPase Immu- cation. nolocalization in the Antennal Urinary

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Glands of the Lobster Homarus gammarus longicaudatus to Hypoxia. Crustaceana , (Crustacea, Decapoda). J. Histochem. Cyto- 69(8): 937-948. chem ., 53(10): 1203-1214. 21. Seaman, M. T., Kok, D. J., Schlichting, B. J. 10. Khodabandeh, S., Golzari, A. and Seyfabadi, and Kruger, A. J. 1991. Natural Growth and J. 2008. Occurrence of the Tadpole Shrimp, Reproduction in Triops granarius (Lucas) Triops cancriformis (Bosc, 1801) (Crusta- (Crustacea: Notostraca). Hydrobiologia , 212 : cea: Notostraca) in Iran. Zoology in the 87-94. Middle East , (In Press). 22. Suno-Uchi, N., Sasaki, F., Chiba, S. and 11. Kheirandish. M. B., 1975. Investigation on Kawata, M. 1997. Morphological Stasis and Adverse Circumstances on the Growth of Phylogenetic Relationships in Tadpole Apus and Leptestheria in Sturgeon Fish Shrimps, Triops (Crustacea: Notostraca). Ponds. PhD. Thesis, Veterinary College, Te- Biochemi J. Linnean Soc ., 61(4): 439-457. hran University. 23. Su, T. and Mulla, M. S. 2001. Effects of Nu- 12. Kohnehshahri, M. and Takami, Gh. A. 1974. tritional Factors and Soil Addition on Artificial Propagation and Culture of Stur- Growth Longevity and Fecundity of the geons . Tehran University Press. PP. 181- Tadpole Shrimp Triops newberryi (Noto- 186. straca: Triopsidae), a Potential Biological 13. Mantovani, B., Cesari, M. and Sabelli- Control Agent of Immature Mosquitoes . J. Scanabissi, F. 2004. Molecular Taxonomy Vector Ecol ., 26(1): 43-50. and Phylogeny of the 'Living Fossil' Line- 24. Takahashi, F. 1997. Use of the Tadpole ages Triops and Lepidurus (Branchiopoda: Shrimp (Triops spp.) as a Biological Agent Notostraca). Zoologica Scripta ., 33 : 367- to Control Paddy Weeds in Japan . PP.128- 374. 137. 14. Martin, J. W. and Boyce, S. L. 2005. 25. Taylor, C. M., Bryant, M. and Hartman, R. Crustacea: Non Cladoceran Branchiopoda , E. 1987. Eastward Range Extension of the PP. 284-297. Tadpole Shrimp, Triops longicaudatus (Le- 15. Martoja, R. and Martoja-Pierson, R. 1967. conte), in Oklahoma. Proc. Okla. Acad. Sci ., Initiation aux Techniques de L’histologie 67 : 75-76. Animale . Masson et Cie, Paris. 345 PP. 26. Tietze, N. S. and Mulla, M. S. 1989. Prey- 16. Petrov, B. and Cvetkovic, M. D. 1996. A size Selection by Triops longicaudatus (No- First Record of Males of Triops Cancri- tostraca: Triopsidae) Feeding on Immature formis (Bosc, 1801) (Crustacea, Branchio- Stages of Culex quinquefasciatus . J. Am. poda) from Yugoslavia. Belgrade, Aech. Mosquito Cont. Asson ., 5(3): 392-396. Biol. Sci ., 48 (1-2): l 59-62, 27. Trentini, M. and Scanabissi, F. S. 1978. Ul- 17. Petrov, B. and Cvetkovic, M. D. 1999. trastructural Observations on the Oogenesis Morphological Variability of Triops of Triops cancriformis (Crustacea, Noto- cancriformis (Bosc, 1801) (Crustacea, straca), I. Origin and Differentiation of Notostraca) in Yugoslavia. Contributions to Nurse Cells. Cell Tiss. Res., 194 : 71-77. the Zoogeography and Ecology of the East- 28. Trentini, M. and Scanabissi, F. S. 1982. Fol-

Downloaded from jast.modares.ac.ir at 0:11 IRST on Tuesday September 28th 2021 ern Mediterranean Region , Vol. 1, Athens licle Duct Cell Ultrastructure and Eggshell PP. 379-385. Formation in Triops cancriformis (Crusta- 18. Scanabissi, F. S., Eder, E. and Cesari, M. cea, Notostraca). J. Morphol., 172 : 113-121 2005. Male Occurrence in Austrian Popula- 29. Tommasini, S., Scanabissi, F. S. and tions of Triops cancriformis (Branchiopoda, Trentini, M. 1989. Scanning Electron Micro- Notostraca) and Ultrastructural Observations scope Study of Eggshell Development in of the Male Gonad. Invertebr. Biol ., 124(1): Triops cancriformis (Bosc) (Crstacea, Noto- 57-65. straca). Vieet Milieu , 39 (1): 29-32. 19. Scholnick, D. A . 1995. Sensitivity of Meta- 30. Zaffagnini, F. and Trentini, M. 1980. The bolic Rate, Growth and Fecundity of Tad- Distribution and Reproduction of Triops pole Shrimp Triops longicaudatus to Envi- cancriformis (Bosc) in Europe (Crustacea, ronmental Variation. Biol. Bull ., 189(1): 22- Notostraca). Monitore Zool. Ital., (N. S.) , 28. 14 :1-8. 20. Scholnick, D. A. and Snyder, G. K. 1996. Response of the Tadpole Shrimp Triops

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ﻣﻄﺎﻟﻌﻪ ﺑﺮﺧﻲ ﺧﺼﻮﺻﻴﺎت زﻳﺴﺘﻲ ﻣﻴﮕﻮي ﺑﭽﻪ وزﻏﻲ، Triops cancriformis ، در آﺑﮕﻴﺮﻫﺎي ﻓﺼﻠﻲ اﺳﺘﺎن آذرﺑﺎﻳﺠﺎن ﻏﺮﺑﻲ

ا . ﮔﻠﺰاري، ص . ﺧﺪاﺑﻨﺪه و ج . ﺳﻴﻒ آﺑﺎدي

ﭼﻜﻴﺪه

ﻣﻴﮕﻮي ﺑﭽﻪ وزﻏﻲ ﺟﻨ ﺲ Triops ﻓﺴﻴﻞ زﻧﺪه ﺷﻨﺎﺧﺘﻪ ﺷﺪه اي اﺳﺖ ﻛﻪ در ﻃﻮل 220 ﻣﻴﻠﻴﻮن ﺳﺎل ﮔﺬﺷﺘ ﻪ ﺗﻐﻴﻴﺮات اﺳﺎﺳﻲ رﻳﺨﺖ ﺷﻨﺎﺳﻲ اﻧﺠﺎم ﻧﺪاده اﺳﺖ . در ﭘﺎﻳﻴﺰ ﺳﺎل 2005 ، ﻧﻤﻮﻧﻪ ﻫﺎي ازﮔﻮﻧﻪ اروﭘﺎﻳﻲ اﻳﻦ ﺟﻨﺲ ، ، ﻳﻌﻨﻲ Triops cancriformis در آﺑﻬﺎي اﺳﺘﺨﺮﻫﺎي ﻓﺼﻠﻲ واﻗﻊ در ﺑﺨﺶ ﺟﻨﻮﺑﻲ درﻳﺎﭼﻪ اروﻣﻴﻪ ﺟﻤﻊ آوري ﻛﺮده و ﺑﺮﺧﻲ ﺧﺼﻮﺻﻴﺎت زﻳﺴﺘﻲ آﻧﻬﺎ ﻣﻮرد ﻣﻄﺎﻟﻌﻪ ﻗﺮار ﮔﺮﻓﺖ. رژﻳﻢ ﺗﻐﺬﻳﻪ اي اﻳﻦ ﺳﺨﺖ ﭘﻮﺳﺘﺎن واﺑﺴﺘﻪ ﺑﻪ ﻣﺠﻤﻮﻋﻪ زﻳﺴﺘﻲ ﻣﻮﺟﻮد در اﺳﺘﺨﺮﻫﺎي ﻓﺼﻠﻲ ﺑﻮده و ﺑﻲ ﻣﻬﺮﮔﺎن ﻛﻮﭼﻚ و ﭘﻮده ﻫﺎي ﺟﺎﻧﻮري ﺑﻴﺸﺘﺮﻳﻦ ﻣﺤﺘﻮﻳﺎت روده را ﺗﺸﻜﻴﻞ ﻣﻲ دادﻧﺪ . وﺟﻮد ﺳﻴﺴﺖ ﻫﺎ و ﺧﺮده ﻫﺎي ﺗﻨﻪ ﻣﺮﺑﻮط ﺑﻪ ﻫﻤﻨﻮﻋﺎن در دﺳﺘﮕﺎه ﮔﻮارش ﻧﻤﻮﻧﻪ ﻫﺎي ﺑﺮرﺳﻲ ﺷﺪه ﻧﺸﺎﻧﮕﺮ ﺣﺎﻟﺖ ﻫﻤﺠﻨﺲ ﺧﻮاري در آﻧﻬﺎ ﺑﻮ د . ﺑﺮرﺳﻴﻬﺎي ﺑﺎﻓﺖ ﺷﻨﺎﺳﻲ و رﻳﺨﺖ ﺷﻨﺎﺳﻲ ﻧﺸﺎن داد ﻛﻪ ﻏﺎﻟﺐ اﻓﺮاد ﺟﻤﻌﻴﺖ ﻣﺎده ﺑﻮده و ﺗﻨﻬﺎ ﻳﻚ ﻣﻮرد از 400 ﻧﻤﻮﻧﻪ ﮔﺮﻓﺘﻪ ﺷﺪه ﻧﺮ ﻣﻲ ﺑﺎﺷﺪ . ﻣﺸﺎﻫﺪه اﺳﭙﺮم در ﻟﻮﻟﻪ ﻫﺎي ﻓﻮﻟﻴﻜﻮﻟﻲ ﺑﺮﺧﻲ از ﻧﻤﻮﻧﻪ ﻫﺎ ﻧﺸﺎﻧﮕﺮ دوﺟﻨﺴﻲ ﺑﻮدن آﻧﻬﺎ ﺑﻮد، اﻣﺎ ﺑﻪ ﻧﻈﺮ ﻣﻲ رﺳﺪ ﻛﻪ روش ﻋﻤﺪه ﺗﻮﻟﻴﺪ ﻣﺜﻞ آﻧﻬﺎ ﺑﻜﺮزاﻳ ﻲ ﺑﺎﺷﺪ. ﺑﺎروري ﺑﺴﺘﻪ ﺑﻪ ﺳﺎﻳﺰ ﺟﺎﻧﻮر، ﺑﻴﻦ 100 - 2500 ﺳﻴﺴﺖ ﻣﺘﻔﺎوت ﺑﻮد . ﻣﺘﻮﺳﻂ ﻗﻄﺮ ﺳﻴﺴﺖ ﻫﺎ 85 ± 400 ﻣﻴﻜﺮون اﻧﺪازه ﮔﻴﺮي ﺷﺪ.

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