Cytologicalrelationshipsof Some Pacificgastropodsi・2 IJSCV ?'I
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The malacologicalsocietymalacological society of Japan Cytological Relationships of some Pacific Gastropodsi・2 J. B. BuRcH3 (The 406th Medical Laberatory, U.S. Army Medical Command, Japan and The University of Michigan, Ann Arbor, Michigan, U,S.A.) IJSC V ?'i':- Jdi Hti J(l! ts{ cD fiEM ECtl t):-i ]'4 ts pm tw ma en J. B. A---.e (I'ptLIa Text-fig, 1 ; fi Tables 1-17) Introduction Information on the chromosomes of snails of the Pacific area is scarce, and it is only recently that any reliable data have been gathered. Nevertheless, it is of in- terest and value to compare what cytologica! information is available for Pacific gastropod mollusks with similar information on snails from other geographical regions, and to discuss the cytotaxonomic value of current data and the prospective usefulness of cytological studies as anticipated in the future. The existing reliable information on chromosome numbers and the taxonomic use- fulness of this information was reviewed by BuRcH (1965a) for the gastropod Euthy- neura and by PA'rrF.RsoN (1967a) for the gastropod Streptoneura. From these reviews it was observed or concluded that in the Euthyneura 1) haploid chromosome numbers (including polyploid numbers) range from 5 to 72; 2) although different chromosome numbers characterize certain taxa within this subclass, the chromosome numbers of the various species comprised by these taxa remain remarkably constant ; 3) therefore, mechanisms responsible for addition or deletion of chromosomes must operate at a very low frequency or efficiency in these taxa;4) changes in chromo- some numbers among the higher systematic categories are brought about through aneuploidy ; 5) polyploidy has been found only in the Basommatophora, but even there it is notcommon and, when present, is evident mainly at the species level, Although polyploidy, where it exists, may produce new species, it does not appear to be of significance in the formation of higher taxa; 6) in most cases, the more advanced 1) Contribution No. 11, Asian Mollusks Program, Institute of Malacology. This paper was pre- "Evolution, sented to Symposium No. 4, Distribution and Migration of Plants and Animals in the Pacific Area" at the E'eventh Pacific Science Congress, Tokyo, Japan, 1966. Gratefu] acknowledgement is made te the National Research Council, National Academy of Sciences, Washington, D. C., fer travel funds to attend th{s Congress. 2) This investigation wa$ sponsored by the Commission on Parasitic Diseases of the Armed Forces Epidemiological Board, and was supported (in part) by the U. S. Army Medical Research and Development Comrnand and (in part) by research grants GB-3006 and GB-787 from the National Science Foundation, Washington, D. C, 3) Supported by a Research Career Program Award (Ne. 5-K3-AI-19, 451) from the National of and Infectious Diseases, U. S. Public Health Institute Allergy Service. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan BuRcH: Cytological Relationships of Some Pacific Gastropods 119 groups among the Euthyneura, or those often considered more specialized, also have the greater number of chromosomes (Text-fig. 1). In the Streptoneura, it was observed or concluded that 1) haploid chromosome numbers range from 7 to 47 ; 2) like the Euthyneura, the Streptoneura exhibit a con- servativeness in chromosome numbers, with variation in chromosome numbers seldom more than ± 2 bivalents in the lower taxa; 3) within the Streptoneura, the Archaeo- gastropoda and Mesogastropoda have relatively low numbers, whereas the mere advanced order Neogastropoda has higher numbers ; 4) however, with the fragmentary information available, there appears to be no general clear-cut correlation between "primitiveness" !ow chromosome numbers and among the various groups. However, within the Viviparacea, such a correlation may exist ; 5) polyploidy, suspected in the Hydrobiidae, and reported in the Thiaridae, needs to be confirmed. Chromosomes have been studied of snail species from the following Pacific areas : Japan, Taiwan (Forrnosa), New Zea!and, New Ca!edonia, the Solomon Islands, the Marshall Islands. Hawaii and the Galapagos Islands. In order to provide a framework of reference for chromosome numbers of Pacific snails, I would like to briefly review eurrent information on chromosome numbers in gastropods. The Opisthobranchiate Orders In the Notaspidea only one species has had its chromosome number determined, Pleurobranchaea novae2ealanditze from Japan, with 12 pairs of chromosomes (Table 1). In the Nudibranchia, chromosomes have been studied in 17 species of 14 genera and 10 families, all from the Pacific area. It is of considerable interest to note that all of these had the same chromosome number, h =13. Table 1. Chromoseme Numbers in Notaspidea and Nudibranchia' Chromosome 1 Family I.ecality No, Genera No. Species Nurnber (n) Notaspidea Pleurobranchidae 12 Japan i 1 / 1 Nudibranchia Dorididae 13131313131313131313 Japan / 4121111111 52211I1121 Marshall Is, Dendrodorididae i Japan, Triophidae japan Goniodorididae Japan E :ilt1 Fimbriidae 1, Japan Dotonidae i Japan Arminidae l Japan Cuthonidae , Japan ': Facelinidae Japan i1! Favorinidae Marshall Is. l1i " From INABA & Hmoi'A, 1954, 1958 ; INABA, 1959b,c, l961a; Bulusu & NATARA,JAN, 1967. NII-Electronic Library Service The malacologicalsocietymalacological society ofJapanof Japan 120 VENUS: 25<3.4)1967 In other opisthobranch 17 species have the groups, been fpund to have 17 pairs of chromosomes, which again demonstrates the great conservativeness of chromosome numbers in the lower Euthyneura (Table 2). Only in the Smaragdinellidae of the Cephalaspidea, the Polybranchiidae of the Sacoglossa, and the Soleolifera (Table 3) have haploid numbers other than 17 been found, and, except for the sacoglossan, these have not varied from 17 more than ± 1 bivalent Table 2.ChromesomeNumbers inOpisthobranchs" Chromosome Family Locality No. Genera No. Species Number (n)1213 Notaspidea JapanJapan, i 115 118 Nudibranchia Marshall Is, . Entomutaemata Pyramidellidae 17 '' Japan 1 1 Anaspidea /F 4 Aplysiidae 17 Japan,Marshall Is. 4 Cephalaspidea / Acteonidae 1717171718 JapanJapanJapanMar$hall 11112 11122 Philinidae l' Aglajidae 1 1 Atyidae Is. Smaragdinellidae Marshall Is. Saceglossa i1i11 Elysiidae 17 Japan, Italy 1411 2511 Stiligeridae 17 Japan, Italy Juliidae 17 JapanItalyMarshall 11,' Polybranchiidae 716-18 Soleolifera Is,, Japan, !I India, U. S, A. 4 5 1 1BvRcH, * From 1959c, MANeiNo &SOIMI, 1964i NATARAJAN, 1959, 1960; 1965a; BuRcHINABA.,& NATARAJAN,1961a; 1967. Table 3. Chromosome Numbers in Soleolifera' Species Chromosome LocaHty /1/I Number (n)i1 Veronicellidae Veronicelta floridana , 1617 U, S, A. i Laevicaulis alte l India Onchidiidae Onchidella kurodai i 1718 JapanMarshall / 1 Onchidella evelinae Is. Onchidium / verracutatum 18 India * From 1959, 1960; INABA, 1961a; BuRcH, 1965a; BvzRcHNATARAJAN,& NAaTARAJA)i, 1967. NII-Electronic Library Service The malacologicalsocietymalacological society ofJapanof Japan Buku]it :CytologicalRelationships of SomePacificGastropods 121 The Basommatophora In the primitive marine Basommatophora, the haploid chromosome number of the 4 species of SiPhonaria studied is 16 (Table 4). Additionally, I recently studied (un- published) another Japanese species, S. sirius, which also had 16 pairs of chromosomes. The haploid chromosome number of Salinator taleii of the operculated Amphibolidae is 18, the number found in most of the other Basommatophora studied. In the EIIobiidae, haploid chromosome numbers range from 17 to 19, but most species have 18 pairs of chromosomes (MEyER, 1955; NATARA.TA/N, 1958a; BuRcii, 1960al NATARAJAN & BuRcH, 1966). In the primitive freshwater Basommatophora, the 3 species studied all had 18 pairs of chromosomes (Table 5). Acroloxus lacustris is cytologically quite distinct from all other Euthyneura, and quite different from the New Zealand Latiafluviatilis, where some think its atfinities lie Csee HuBENDIcK, 1962). In the higher limnic Basommatophora, 7 species of the Physidae have been studied, all with the haploid number 18 (LE CALvEz & CERTAiN, 1950; BuRcH, 1960a; BuRcH & BusH, 1960; INABA, 1965). In the Lymnaeidae, 34 species have been studied, 23 of which had 18 pairs of chromosomes (Table 6). But the 9 species of the genus Radix Table 4. Chromosome Numbers in Marine Basommatophora* - ... tt tttt .ttttttttttmttt.t.. .. tt-- tttt tt .==.. CNhurmO ii・ species I8rSO(Mt)e Locaiity Siphonariidae 16 Siphenaria l'aPonica Japan 16 Marshalt ls. SiPhonaria g"aneensis, / laciniosa l・ 16 New Caledonia SiPhonaria ' , 1 / SiPhonaria Pectinata 16 / Florida, U. S. A. Amphibolidae ! 18 ,. Salinator takii Japan . ! - * From INABA, 1953 ; NA'rARAJAN & BuRcH, 1966. Table 5. Chromosome Nurnbers of Primitive Aquatic Basommatophora* Chromosome Spec.ies Locality Number (n) Latiidae Latia neritoides 18 Zealand / New Chilinidae / Chilina nttviatitis 18 Uruguay Acroloxidae Acroloxus lacustris 18 Eng]and * From BuRcJt, BuRcH & PAT7・EKsON,1963; NATARAJAN & BvRcH, 1966,1962; NII-Electronic Library Service The malacologicalsocietymalacological society ofJapanof Japan 122 VENUS: 25 (3.4) 1967 Table6. Chromosome Numbers in Lymnaeidae* tt tt ' 1 Chromosome I Genera No, Species Numbers Cn) , ' '' / Bakerilymnaea-- l' 16 1' Radix 17 9 i L,ymnaea 18 1 Stagnicola 18 11 Acella 18 ・ 1 / Pseudosuccinea 18 1 Butimnea 18 1 Fossaria 18-19 4 'i Lanx 18 3 Fishereta 18 2 ' * From PERRoT, lg3o, lg34; J.-L., PERRoT & pl]Rl[oT,'lggs; LE CAnvEz & CERTAiN, 1950; BuRcH, 1960a, 1963a, 1965a,