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Title Morphometric Variation Analyses and Revision of the Japanese Toads

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Title Morphometric Variation Analyses and Revision of the Japanese Toads Morphometric Variation Analyses and Revision of the Title Japanese Toads (Genus Bufo, Bufonidae) Author(s) MATSUI, Masafumi Contributions from the Biological Laboratory, Kyoto Citation University (1984), 26(3-4): 209-428 Issue Date 1984-07-31 URL http://hdl.handle.net/2433/156031 Right Type Departmental Bulletin Paper Textversion publisher Kyoto University Contr. biot. Lab. K),oto Univ., Vol. 26, pp. 209-428 Issued 31 July 1984 Morphometric Variation Analyses and Revision of the Japanese Toads (Genus Bufo, Bu fonidae)i) Masafumi MATsui Abstract Two problems havc been involved in the classification of the Japanese toads : (1) relation- ships with the continental forms, and (2) relationships among forms hitherto described within the islands. The present study was undertaken with the purpose ot- resolving the latter problem, and to achieve this, morphometric variation analyses were made ofa large number of preserved specimens. The results were synthesized with the hitherto accumulated non- metrical and non-morphological information so as to taxonomfcally revise the Japanese toads. A historical review of the classificat{on of theJapanese toads revealed that the most complex problems were involved in the still-water breeding forms from the main islands (= the Japanese common toad), and that thc taxonomic confusion was caused mainly by previous authors' classification through examination ofa small number ofspecimens notwithstanding the great variability ofexternal morphometric characters ofthe toads. The critical points for measurc- ments fbr each character, and problcms with measurement crrors duc to variant fixation and preservation are discussed. Allomorphic relations of 30 charactcrs to SVL were analyzed fbr four age/sex groups o{' 147 Japanese common toads from rvlomoyama, Kyoto, with thc result that many characters should be trcated separately tbr three groups, i.c., young, adult males, and adult females. Besides, it was determined that simple ratios to SVL should not be calculated for many characters. For theJapanese common toad, SVL and relativc value (ACV:=theoretical value calculated by the allomorphic relationship of each character to SVL) of 10 selected characters for each of 2,525 specimens belonging to 96 populations were analyzed, and the presence or absence of intrapopulational age and sexual variations and of interpopulational variations was noted. From these analyses, the Japanese common tbad was divided into northeastern (A) and southwestern (B) types chiefiy by the relative size of thc tympanum diameter. For the two types of the .Japanese common toad, presence of morphometric clines in re!ation to geographical and/or climatological parameteis was in- vestigated. Clear clinal tendencies were found mostly in type A, whereas few clear trends were detected in type B. Taxonomical meaning of the clines was discussed, and it was suggested that the Japanese common toad should be split into taxa beiow the species level. Similar morphometric analyses were made on the still-water breeding Miyako toad from thc Ryukyu Archipelago. The results for 176 specimens from 4 populations ind{cated that this form was very uniform in. external morphometry, that it greatly differed from the Japanesc common toad in several characters, and that the differences were greater bctween this form and geograph{cally adjacent type B Japanese common toads in some characters. Similar analyscs were also performed on 265 specimens of 13 populations of thc stream-breeding ,Japanese stream toad. Interpopulation variation in this form was very small. Comparisons with sympatric and allopatric populations of thc Japanese common toad wcre made with the, result that the two fbrms were found to be morphometrically almost complctcly separatcd, showing grcatest differentiation in thc zQne. of sympatry. Taxonomic ('onclusions wcre drawn with thesc rcsults in mind. For tlie.Japa. ncsÅë c,ommon toad, thc typc scries or topc)- types of each of the hitherto described fbrms wcre compared with thc othcr populations and it was determined that neither type A nor typc B can bc split further. Thc syntypes of Bqfo l) Part of a thesis submitted to the Faculty of Scien(:e of Kyoto University in partial fu1filment ot' the requirein(,".nts for the D. Sc. degree. 210 MMATsul vul•garis .iaPonicus, which is the oldest of all the described forms from Japan and is now kept in Leiden, were examined and a lectotype was designated and described. This form clearly belonged to type B. 'rhc nature of the intergradation of morphometric. variations between type A and type B is discussed, and special attention is paid to the sudden change in the morphocline in the tympanum diameter. From this discussion, the relation of thc two typcs is regarded as subspecific, and they are treated as B. j'aponicus fbrmosus (type A) and B. i j'aponicus (type B). For the Miyako toad, the morphometric difference f'rom the Japanese, common toad in the adjacent distribution range was studied, and the two fbrms are regarded as belonging to the different lineages. In addition, the known results ofartificial hybridization support this idea, and therefbre, the form is regarded as specifically different from J'aPonicus and is treated as a subspecies (mi akonis) of the northeastern Chinese species B. gargarizans. The Japanese stream toad has been showii to have strong genetic compatibility with the Japanese common toad. Therefbre, from the viewpoint of evolutionary taxonomy, the taxonomic value of the evidence from genetical sources is discussed. The two forms are distributed sympatrically, but are judged to seldom interbreed or show morphological inter- gradation in nature. Thus, notwithstanding the strong genetic compatibility of the artificial hybrids, theJapanese stream toad is regarded as a full species, B. torrenticota, as was originally described. In conclusion, the Japanese toads are taxonomically divided into three species, Bufo j'aPonicus (.i. J'aPonicus Schlegel, 1838 and i formosus Boulenger, 1883), B. gargarizans miyakonis Okada, l931, and B. torrenticola M. Matsui, 1976. Contents Introduction .,............,...............,..........,.........................,...... 214 I History and problems of the taxonomy of the Japanese toads . 216 1. A historical review ofclassification oftheJapanese toads ........,.. 216 a) 1826-1907 ••••J•••'''''''''''''''''''''''''''''''''''''''''''''''''''''''-'''' t--t 216 b) 1908-1942 .......................,.....•-•-••••••••J••••••••••••-•••-i•••-•-•• .220 c) 1943-1966 ........................,......,..,,,.,......,..,............,...... .221 - .---+- . d) 1967- .""".."",".".H"H.,.,..,.-."..".,.,",..",.",H",...,.,. 22l 2. Taxonomic problems and characters to be chosen ,.••J.J•.••J•JJ-•••• 222 a) Taxonomicproblem.......,....,.,.,...,...,.,.,..,.,......,...,....,,..,., .222 b) Characters to be chosen .....,.........,.....,.,.............,...,......., 224 +- --..---- ------ II General description of materials and methods of observation - -- -t -- -J- -i- -- -- - . 225 l. "daterials .,.,H,.""."..,.,".,.".."..,,",,H,.,H.,". ---J-.-J----- - -. - --- -- l .-- - -J -- - 225 a) Population ..............................,.,........,.,..,. 22Jr --------------- -----J------------ d b) Determination ofsex and maturity ......,......,. ---t------l-- . ----4---------- . 230 2. 0bservation methods ......,....,......••-•••-•••••i•••- ---------+----- ------------------ 234 a) Methodsofmeasurements ....,...,.,.,...,.,........ ---J-J--------- -t -- -. -- -- --- --- -J 234 b) Dimensions measured ............,.,................., --------------- -- -- -- -J -- --- -- E-. 236 i) Standardcharacterdimension .......,......,.., -i -- -- -- -- --- -------------..-.. 236 ii) Characterdimensionsofhcadregion ..,...... -. -.. - --t --- --t -- 237 iii) Characterdimensionsofforelimb .,,,.,.......,. 238 iv) Character dimensions ofhindlimb ,..,.........,. 238 c) Shrinkag.ein fixedspecimens .....,....,.........,,., 239 1) Shrinkag. e caused by forrnalin fixation ......,., ..239 ------t------ -t -- - --. --- -- --- -J 2) Shrinkagcin toads long preservcd in cthanol ----t--t----4 -tt--tttttttt-- . 240 llI Intrapopulation allomorphosis in the Japanese commontoad from Momeyama) Kyoto...................,...........................,....,....,., 242 -----.------------ . Materials and methods ...........................,..,...,.... 243 ------------- . .-------------+-.- . Results .....".......................................,............, , 244 -- -- -'- i' -- -- -l---t--l--------- . a) Regression to allomorphicformulae ,.............. 244 --Jl--t-----l-- -----------------l . ' b) 'I"ypes ofallomorphosis in each agc/sex group ••, 249 ----t---,--------t MorPhometr7 and Revision ofJaPanese Toads 211 c) Comparisons ofallomorphic relations among agelsex groups ....... .... ..250 . + . + . ...254 d) CharacteristicsofallomorphiccoeMcientsinthecombinedgroups . Discussion .................................................................................. ..257 . + Regression formula and correlation coefficients ............................... .257 + Slope and position of the regression lines .....................................,.. .258 . , Intrapopulationvariation,.......................,..........................,......... ,....259 . ' Growth gradient and function .................................................... ......260 . IV Morphometric variation in the Japanese cornmon toad ............. ...261 . , Materials and methods ..,..............,...........,.....,•......,............,.•••••• . .261 , . a) Materials ..........................,...........,.................................
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