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Preliminary Data on Larval Development of Caucasian Cave-Dwelling Shrimp Troglocaris (Xiphocaridinella) Kumistavi Marin, 2017 (Crustacea: Decapoda: Atyidae)

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Preliminary Data on Larval Development of Caucasian Cave-Dwelling Shrimp Troglocaris (Xiphocaridinella) Kumistavi Marin, 2017 (Crustacea: Decapoda: Atyidae) Arthropoda Selecta 26(4): 297–302 © ARTHROPODA SELECTA, 2017 Preliminary data on larval development of Caucasian cave-dwelling shrimp Troglocaris (Xiphocaridinella) kumistavi Marin, 2017 (Crustacea: Decapoda: Atyidae) Ïðåäâàðèòåëüíûå äàííûå ïî ëè÷èíî÷íîìó ðàçâèòèþ êàâêàçñêîé ïåùåðíîé êðåâåòêè Troglocaris (Xiphocaridinella) kumistavi Marin, 2017 (Crustacea: Decapoda: Atyidae) Ivan N. Marin1,2, Sergey Yu. Sinelnikov3 Èâàí Í. Ìàðèí1,2, Ñåðãåé Þ. Ñèíåëüíèêîâ3 1 A.N. Severtzov Institute of Ecology and Evolution of RAS, Moscow, 119071 Russia. E-mail: [email protected],[email protected] 1 Институт проблем экологии и эволюции им. А.Н. Северцова РАН, Москва, 119071 Россия. 2 Biological Department, Altai State University, Barnaul, 656049 Russia. 2 Биологический факультет, Алтайский государственный университет, Барнаул, 656049 Russia. 3 I.D. Papanin Institute of Biology of Inland Waters of RAS, Yaroslavl Province, 152742 Russia. E-mail: [email protected] 3 Институт биологии внутренних вод им. И.Д. Папанина, Ярославская область, 152742 Россия. KEY WORDS: Crustacea, Decapoda, Atyidae, Troglocaris, Xiphocaridinella, lipid reserves, stygobiotic, development, larvae, Caucasus. КЛЮЧЕВЫЕ СЛОВА: Crustacea, Decapoda, Atyidae, Troglocaris, Xiphocaridinella, липидные запасы, стигобиоты, развитие, личинка, Кавказ. ABSTRACT. Up to date, any characteristics of the представлено описание первой находки яйценос- development of stygobiotic Troglocaris shrimp larvae ных самок, личиночного развития и использование remained unknown. The article briefly describes the желтка в развитии личинок пещерной креветки first case of finding of ovigerous females, larval devel- Troglocaris (Xiphocaridinella) kumistavi Marin, 2017 opment and the use of yolk lipid in larval nutrition of (Crustacea: Decapoda: Atyidae). Показано, что ли- Caucasian cave-dwelling shrimp Troglocaris (Xipho- чинки Troglocaris являются планктонными и леци- caridinella) kumistavi Marin, 2017 (Crustacea: Deca- тотрофными, в течение всего периода развития спо- poda: Atyidae). Troglocaris larvae are planktonic and собны развиваться исключительно за счет запасов lecithotrophic, able to develop using the expense of яичного желтка, не питаясь. Данная особенность egg yolk exclusively, without feeding. This trait allows позволяет креветкам рода Troglocaris размножать- shrimps of the genus Troglocaris completing metamor- ся в олиготрофных, лишенных планктона, пещер- phosis in oligotrophic planktonless cave water reser- ных водоемах в период межени, когда уровень воды voirs during the low water period, thus shrimp larvae максимально низкий и личинки не вымываются; are not washed away; however, alochtonic organics do однако и аллохтонная органика в этот период в not enter into caves during this period as well. Charac- водоемы также не поступает. Особенности личи- teristics of larval development of T. kumistavi also ночного развития T. kumistavi также позволяют по- make it possible to understand that these cave shrimps нять, что эти пещерные креветки могут размно- can reproduce only in large stable water reservoirs жаться лишь в крупных водоемах со стоячей водой (underground lakes), and therefore spend much of the (подземных озерах), а значит и проводят там боль- life cycle there, than in rivers or streams. шую часть жизненного цикла, нежели в подземных How to cite this article: Marin I.N., Sinelnikov реках или ручьях, куда их, скорее всего, вымывает. S.Yu. 2017. Preliminary data on larval development of Caucasian cave-dwelling shrimp Troglocaris (Xipho- caridinella) kumistavi Marin, 2017 (Crustacea: Deca- Introduction poda: Atyidae) // Arthropoda Selecta. Vol.26. No.4. P.297–302. doi: 10.15298/arthsel.26.4.03 Limitation and irregular supply of food is a major driving force for the evolution of cave animals [Van- РЕЗЮМЕ. Особенности развития личинок рода del, 1965; Barr, 1968]. Subterranean animals, includ- пещерных креветок Troglocaris оставались до пос- ing shrimps, fishes and salamanders, are able to live леднего времени неизвестными В статье кратко without food for long periods, up to several years [Van- 298 I.N. Marin, S.Yu. Sinelnikov del, 1965; Hüppop, 1985]. According to the energy- Material and methods economy hypothesis, the adaptation of subterranean animals to low energy includes a reduction in pigments Shrimps were collected in subterranean cave streams and eyes, infrequent reproduction, small brood size, and lakes in Kumistavi Cave, Tskaltubo-Kumistavi reduced metabolism and growth rates, and increased (42°22′35.8″N 42°36′03.2″E) by hand net in February longevity [Poulson, White, 1969; Howarth, 1983]. 2016 that represent the period of larva hatching clearly Most of specialized animals possess specific me- correlated with the low water in cave reservoirs oc- tabolism and organs storing food and resources. The curred in winter season, from December to March. crustacean hepatopancreas is the primary site for lipid This period in the winter season differs from the sum- digestion and metabolism [Chang, O’Connor, 1983; mer season (June–September) by a lower level of karst Gibson, Barker, 1979]. The lipid reserves in cave- water in the cave. Collected ovigerous females were dwelling shrimps of the genus Troglocaris are stored transported to the lab in the Moscow State University extracellularly in specialized compartments of hepato- where 4 planktonic larvae successfully hatched. After pancreas, named oleospheres [Vogt, Strus, 1992, 1999], hatching, all larvae were removed to à separate clean a fascinating adaptive mechanism to survive in the water 1-liter tank, and were photographed alive each 2 nutrient-poor hypogean environment. Well-developed days and then fixed in 90% ethanol for further morpho- oleospheres in T. anophthalmus allowing survival of logical analysis. Fixed larvae are deposited in the col- shrimps without food for more than 2 years [Vogt, lection of A.N. Severtzov Institute of Ecology and Strus, 1999]. At the same time, nothing is known about Evolution of Russian Academy of Sciences, Moscow, Troglocaris larval development and larva nutrition as Russia (LEMMI). Species names and modern taxo- only few females with eggs has been sampled in Bal- nomic position are given according to the international kans [Matjasic, 1958; Juberthie-Jupeau, 1969; Jugovic database WoRMS (World Register of Marine Species) et al., 2015]. Moreover, the release of decapod larvae and Marine Species Identification Portal. The study in the non-hypogean environment has been correlated does not include endangered or protected species. to different environmental cycles [Morgan, Christy, 1994] that remain poorly studies in cave biotopes. Results and discussion The stygobiotic atyid shrimp genus Troglocaris Dormitzer, 1853 currently includes six Caucasian spe- Only 4 larvae were successfully reared and hatched cies referring to the subgenus Xiphocaridinella Sa- under the temperature about 20–21 °C in the water dowsky, 1930 (according to WoRMS, 2017). Three tank. It was impossible to assess the number of zoeal species, Troglocaris (Xiphocaridinella) kutaissiana (Sa- stages and examine zoeal morphology in details be- dowsky, 1930) (the type species of the subgenus), Tro- cause we tried to keep all larvae alive. Detailed mor- glocaris (Xiphocaridinella) fagei Birštein, 1939 and phological description of larval stages will be the aim Troglocaris (Xiphocaridinella) jusbaschjani Birštein, of our further study. However, it is possible to describe 1948, were currently re-described based on topotypic the development of T. kumistavi larvae as complete, material [Marin, Sokolova, 2014; Marin, 2017] as well but abbreviated and accelerated. Preliminary, we con- as a new species, Troglocaris (Xiphocaridinella) clude that T. kumistavi possess 4 or 5 zoeal stage in kumistavi Marin, 2017 was recently described from their development showed in Figs 1, 2. At the same underground stream and lakes of Kumistavi (Prome- time, no exuviae were found in the tank where larvae of theus) Cave, Tskaltubo, Imereti region, Western Geor- Troglocaris stayed during the study as well as it was gia [Marin, 2017] based on morphological investiga- impossible to select exact molting stages in their devel- tion and DNA analysis. Among the specimens of T. opment. It is possible that the remains of the exuviae kumistavi collected in Kumistavi Cave several oviger- were eaten by larvae very quickly, before they were ous females with eggs were sampled, representing the seen. Studied Troglocaris larvae possessed with thin first described locality in Georgia where Caucasian soft (membranous) cuticle and, possibly, were able to Troglocaris shrimps were found breeding (“breeding grow appendages without molting. Non-feeding larvae pool”) [Marin, 2017]. Previously, only non-ovigerous are deprived of the opportunity to replenish the food females of Troglocaris shrimps were found in Cauca- resources and the absence of molting as well as the sian underground lakes and cave streams; probably discharge of the exuvium seems to be a very valuable they were washed out there from their underground feature of their development and an excellent adapta- “breeding pools” by flood water [Marin, Sokolova, tion to the underground way of life. Freshly hatched 2014]. Herewith, we present a preliminary description larvae possessed with partly developed pereiopods and of larvae of T. kumistavi and indications of the yolk developed pleopods (Fig. 2a, b) used for swimming lipid nutrition in larval development of Troglocaris as and skipping. They actively swam with tail
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