Introduction

Museum specimens provide valuable insights into the evolutionary and ecological history of living and extinct populations, their taxonomy and knowledge for conservation biology (Suarez and Tsutsui 2004). For example, morphological and molecular data on historical specimens assist in the clarification of the classification and various other biological issues (Wandeler et al. 2007). In this study, we focused on specimens of endangered freshwater molluscs in a collection of Tomotaro Iwakawa from the National Museum of Nature and Science, Tokyo. Tomotaro Iwakawa (1855-1933) was an early contributor to zoology in Japan and he researched freshwater molluscs in Japan with particular attention (Taki 1933). Furthermore, he established one of the earliest systematic malacological lists in Japan (Taki 1933) and specimens of the list were deposited at the National Museum.

Viviparidae Gray, 1847, belonging to Caenogastropoda, is a family of freshwater gastropods with 125–150 valid described species globally and have a wide geographic range in Asia (Hirano et al. 2019b, Van Bocxlaer and Strong 2019, Stelbrink et al. 2020). Historically, Japanese viviparid gastropods have been classified as four species, based primarily on conchological features (Habe 1990, Masuda and Uchiyama 2004): Cipangopaludina chinensis laeta (Martens, 1860), Cipangopaludina japonica (Martens, 1861), Heterogen longispira (Smith, 1886) and Sinotaia quadrata histrica (Gould, 1859). Recently, some attributions were modified by Hirano et al. (2019b), namely, Cipangopaludina japonica was modified to Heterogen japonica, based on molecular phylogeny. Furthermore, molecular analyses have led to the recognition of two subspecies of C. chinensis in Japan: C. c. laeta and C. c. chinensis and the existence of an undescribed species of Heterogen sp was also suggested (Hirano et al. 2019a, Hirano et al. 2019b). Thus, based on all the above-mentioned work on Japanese Viviparidae, we consider six viviparid species/subspecies to exist in Japan. Of these six species/subspecies, H. longispira is listed on the IUCN Red List as EN (Köhler and Rintelen 2011). Furthermore, C. chinensis is listed on the IUCN Red List as LC, although the subspecies is not designated (Köhler et al. 2012). These two species are also listed on the Red List issued by the Government of Japan as NT and VU, respectively (Ministry of the Environment, Government of Japan 2019). Furthermore, although H. japonica is invasive on some Continents (Van Bocxlaer and Strong 2016), it is also listed on the Japanese Red List as NT (Ministry of the Environment, Government of Japan 2019).

The highly-endangered H. longispira (Köhler and Rintelen 2011, Nakai 2016, Ministry of the Environment, Government of Japan 2019) is endemic to Lake Biwa drainage (Hirano et al. 2015, Hirano et al. 2019b) which consists of Lake Biwa, a river flowing from Lake Biwa, Lake Yogo (artificially drained from Lake Biwa) and Lake Biwa Canal (Fig. 1). Besides, Lake Biwa has an ancient origin with a remarkable biodiversity (Horie 1971, Yokoyama 1984, Kawanabe 1996, Rossiter 2000, Okuda et al. 2013). Prior to its description, H. longispira was included under Paludina ingallsiana Lea, 1856 (Kobelt 1879, Iwakawa 1895, Iwakawa 1897a, Iwakawa 1897b). Then, Pilsbry (1902) indicated that P. ingallsiana does not occur in Japan and H. longispira was included in Vivipara sclateri Frauenfeld, 1865. After this, Vivipara sclateri (or Viviparus sclateri) was adopted as the species name for specimens from Lake Biwa (Hirase 1909, Hirase 1910, Kobelt 1909, Lake Biwa fisheries experimental station 1915, Annandale 1916, Kawamura 1918, Iwakawa 1919). In addition, the name Vivipara sclateri was used for viviparid specimens from Japanese localities other than Lake Biwa (Kobelt 1909, Annandale 1916, Iwakawa 1919). Furthermore, Annandale (1921) pointed out the difference between Vivipara sclateri by Frauenfeld and the distinctive viviparid gastropods from Lake Biwa (= H. longispira), based on comparison with the type illustration of Vivipara sclateri and he described specimens from Lake Biwa as Heterogen turris Annandale, 1921. However, Smith (1886) had already described the endemic species in Lake Biwa as Viviparus longispira Smith, 1886, although this paper was not referred to by other malacologists at that time. Finally, Kuroda (1929) reclassified Viviparus longispira as H. turris and considered Viviparus sclateri to represent a regional subspecies of H. japonica, based on the type illustration of V. sclateri. Later, Vivipara sclateri was generally considered a junior synonym of H. japonica (Yagura 1935, Kuroda 1947b, Kuroda 1955, Kuroda 1963). As Annandale (1921) indicated, H. longispira is clearly different from typical Vivipara sclateri; however, the early malacologists have considered the species to occur in other regions beyond Lake Biwa (Kobelt 1879, Iwakawa 1895, Iwakawa 1897a, Iwakawa 1897b, Iwakawa 1919, Pilsbry 1902, Annandale 1916), although this conclusion remains uncertain. Iwakawa (1919) listed 12 museum lots from 10 localities labelled with 'Naga-tanishi', the current Japanese name for H. longispira, which was at the time equivalent to Viviparus sclateri and three of the 10 localities are outside Lake Biwa drainage. Although Kuroda (1929) said "these may be elongated H. japonica" without any examination, these “H. longispira” from areas outside Lake Biwa drainage in Iwakawa (1919) have not been sufficiently examined and have not been illustrated to date. In addition, there were two records of "H. longispira" from the southern part of Lake Biwa and the Seta River flowing from Lake Biwa; however, H. longispira is now very rare in this area (Nishino 1991, Kihira et al. 2003, Kihira et al. 2009, Nakai 2016). Furthermore, one locality of “H. longispira” was drained and is now terrestrial (Ozawa 2012) and the viviparid gastropods from these localities have not been examined. In this study, to clarify the actual status of “H. longispira” (=Viviparus sclateri) from areas outside the current distribution and to classify extinct populations, we examined the collection of Tomotaro Iwakawa from the National Museum of Nature and Science, Tokyo.

Figure 1.  

A map of collected localities of Viviparus scalateri in Iwakawa's collection . Yellow parts show the estimated ranges, based on the label of the museum lots. Numbers indicate the sample numbers in this study (see also Table 1). Sample numbers 2964J and 2964L were collected from somewhere in Lake Biwa, but the entire Lake is not coloured to make it easier to see. Green letters and downstream of the Seta River are Lake Biwa drainage as defined in this paper. A map is created from digital national land information (Ministry of Land, Infrastructure and Transport of Japan: https://nlftp.mlit.go.jp/ksj/index.html).

Table 1.

Download as

CSV

XLSX

List of examined viviparid lots deposited in the National Museum of Nature and Science, Tokyo, which were treated as Viviparus sclateri (Frauenfeld, 1865) in Iwakawa 1919. The number in each species/subspecies indicates examined individuals in each sample.

Sample No. in this study	Lot Acc. No. in the museum	Locality	Heterogen japonica	Heterogen longispira	Sinotaia quadrata histrica	Remarks
2959J	NSMT-Mo 2959	Japan, Ibaraki Pref., Lake Kasumigaura	3			Probably one of No. 2950 in Iwakawa (1919).
2960J	NSMT-Mo 2960	Japan, Nagano Pref., Lake Suwa	1
2960H	NSMT-Mo 2960	Japan, Nagano Pref., Lake Suwa			1
2961J	NSMT-Mo 2961	Japan, Nagano Pref., Lake Suwa	27
2961H	NSMT-Mo 2961	Japan, Nagano Pref., Lake Suwa			5
2962J	NSMT-Mo 2962	Japan, Nagano Pref., Lake Suwa	6
2963J	NSMT-Mo 2963	Japan, The western part of Aichi Pref.	1
2964J	NSMT-Mo 2964	Japan, Shiga Pref., Lake Biwa	1
2964L	NSMT-Mo 2964	Japan, Shiga Pref., Lake Biwa		1
2966J	NSMT-Mo 2966	Japan, Shiga Pref., Hikone City, Hikone	1
2966L	NSMT-Mo 2966	Japan, Shiga Pref., Hikone City, Hikone		2
2967J	NSMT-Mo 2967	Japan, Shiga Pref., Maibara City, Irie	7
2968J	NSMT-Mo 2968	Japan, Shiga Pref., Tsukuma Lagoon	15			This locality was drained and converted into terrestrial areas
2969J	NSMT-Mo 2969	Japan, Shiga Pref., Otsu City, Zeze	4
2969L	NSMT-Mo 2969	Japan, Shiga Pref., Otsu City, Zeze		6		H. longispira are almost extinct in this locality
2970J	NSMT-Mo 2970	Japan, Shiga Pref., the Seta River	2
2970L	NSMT-Mo 2970	Japan, Shiga Pref., the Seta River		15		H. longispira are almost extinct in this locality

Materials and methods

Data resources

Taxon treatments

Heterogen japonica (Martens, 1861)

Nomenclature

Synonyms:

Materials   Download as CSV 

1. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Ibaraki
   ; locality:
   Lake Kasumigaura
   ; verbatimLocality:
   Kasumigaura (in Japanese) [Lake Kasumigaura]
   ; year:
   before 1919
   ; individualCount:
   3
   ; lifeStage:
   adults
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2959J ex.NSMT-Mo 2959
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965303
   ; occurrenceRemarks:
   NMST-Mo 2959 from Kasumigaura was presumed as one record of 2950, Viviparus sclateri from Kasumigaura in Iwakawa’s catalogue (1919) (see data resources).
   ; recordNumber:
   Probably 2950 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2959J
2. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Nagano
   ; locality:
   Lake Suwa
   ; verbatimLocality:
   Suwako (in Japanese) [Lake Suwa]
   ; year:
   before 1919
   ; individualCount:
   1
   ; lifeStage:
   adult
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2960J ex.NSMT-Mo 2960
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965315
   ; recordNumber:
   2960 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2960J
3. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Nagano
   ; locality:
   Lake Suwa
   ; verbatimLocality:
   Suwako (in Japanese) [Lake Suwa]
   ; year:
   before 1919
   ; individualCount:
   27
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2961J ex.NSMT-Mo 2961
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965316
   ; recordNumber:
   2961 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2961J
4. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Nagano
   ; locality:
   Lake Suwa
   ; verbatimLocality:
   Tenryu-gawa-no-suigen (in Japanese) [The water source of Tenryu River = Lake Suwa]
   ; year:
   before 1919
   ; individualCount:
   6
   ; lifeStage:
   adult
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2962J ex.NSMT-Mo 2962
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965304
   ; recordNumber:
   2962 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2962J
5. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Aichi
   ; locality:
   western part of Aichi Prefectue
   ; verbatimLocality:
   Owari (in Japanese) [Old name of western part of Aichi Prefectue]
   ; year:
   before 1919
   ; individualCount:
   1
   ; lifeStage:
   adult
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2963J ex.NSMT-Mo 2963
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965307
   ; recordNumber:
   2963 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2963J
6. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Shiga
   ; locality:
   Lakw Biwa
   ; verbatimLocality:
   Biwa-ko (in Japanese) [Lake Biwa]
   ; year:
   before 1919
   ; individualCount:
   1
   ; lifeStage:
   adult
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2964J ex.NSMT-Mo 2964
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965302
   ; recordNumber:
   2964 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2964J
7. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Shiga
   ; locality:
   Hikone, Hikone City
   ; verbatimLocality:
   Hikone (in Japanese) [Hikone, Hikone City]
   ; year:
   before 1919
   ; individualCount:
   1
   ; lifeStage:
   adult
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2966J ex.NSMT-Mo 2966
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965311
   ; recordNumber:
   2966 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2966J
8. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Shiga
   ; locality:
   Maibara City, Irie
   ; verbatimLocality:
   Irie-mura (in Japanese) [Irie Village, old name of Irie, Maibara City]
   ; year:
   before 1919
   ; individualCount:
   7
   ; lifeStage:
   adults
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2967J ex.NSMT-Mo 2967
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965312
   ; recordNumber:
   2967 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2967J
9. scientificName:
   Heterogen japonica (Martens, 1860)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Oh-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Shiga
   ; locality:
   Tsukuma Ragoon
   ; verbatimLocality:
   Tsukuma-uchiko (in Japanese) [Tsukuma Ragoon]
   ; year:
   before 1919
   ; individualCount:
   15
   ; lifeStage:
   13 adults 2 young
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2968J ex.NSMT-Mo 2968
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965309
   ; recordNumber:
   2968 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2968J
10. scientificName:
    Heterogen japonica (Martens, 1860)
    ; kingdom:
    Animalia
    ; phylum:
    Mollusca
    ; family:
    Viviparidae
    ; vernacularName:
    Oh-tanishi (in Japanese name)
    ; country:
    Japan
    ; stateProvince:
    Shiga
    ; locality:
    Ohtsu City, Zeze
    ; verbatimLocality:
    Zeze (in Japanese) [Zeze, Ohtsu City]
    ; year:
    before 1919
    ; individualCount:
    4
    ; lifeStage:
    3 adults 1 young
    ; preparations:
    shell (dried)
    ; catalogNumber:
    2969J ex.NSMT-Mo 2969
    ; occurrenceDetails:
    https://www.gbif.org/occurrence/2864965310
    ; recordNumber:
    2969 in Iwakawa 1919
    ; previousIdentifications:
    Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
    ; identifiedBy:
    Takumi Saito
    ; dateIdentified:
    2020-02-03/04
    ; identificationRemarks:
    Some specimens seemingly have the intermediate morphology between H. japonica and H. longispira (see taxon discussion section).
    ; language:
    japanese
    ; bibliographicCitation:
    Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
    ; institutionCode:
    NSMT
    ; collectionCode:
    Mo
    ; basisOfRecord:
    PreservedSpecimen
    ; occurrenceID:
    paper:SK2020:2969J
11. scientificName:
    Heterogen japonica (Martens, 1860)
    ; kingdom:
    Animalia
    ; phylum:
    Mollusca
    ; family:
    Viviparidae
    ; vernacularName:
    Oh-tanishi (in Japanese name)
    ; country:
    Japan
    ; stateProvince:
    Shiga
    ; locality:
    theSeta River
    ; verbatimLocality:
    Seta-gawa (in Japanese) [the Seta River]
    ; year:
    before 1919
    ; individualCount:
    2
    ; lifeStage:
    2 juveniles
    ; preparations:
    shell (dried)
    ; catalogNumber:
    2970J ex.NSMT-Mo 2970
    ; occurrenceDetails:
    https://www.gbif.org/occurrence/2864965317
    ; recordNumber:
    2970 in Iwakawa 1919
    ; previousIdentifications:
    Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
    ; identifiedBy:
    Takumi Saito
    ; dateIdentified:
    2020-02-03/04
    ; identificationRemarks:
    Some specimens seemingly have the intermediate morphology between H. japonica and H. longispira (see taxon discussion section).
    ; language:
    Japanese
    ; bibliographicCitation:
    Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the Tokyo Imperial Museum, Tokyo, Japan, p. 72.
    ; institutionCode:
    NSMT
    ; collectionCode:
    Mo
    ; basisOfRecord:
    PreservedSpecimen
    ; occurrenceID:
    paper:SK2020:2970J

Diagnosis

Adult shell large and thick; shell shape subconical to pyramidal. Adult shell colour dark brown or brown or dark olive, often covered with many environmental attachments, such as alga; shell surface glossy, sometimes having weak hollows, growth lines and spiral striae. Suture moderately deep. Young shell small, relatively thin and fragile; shell shape pyramidal. Young shell colour yellowish-olive; shell surface quite glossy with no spiral ridges on the upper part of the spire.

Taxon discussion

Heterogen japonica can be distinguished from other Japanese Viviparidae by the following criteria which are based on the illustrations and information from literature (Suppl. material 1).

H. japonica can basically be distinguished from Cipangopaludina chinensis laeta by its subconical to pyramidal shell shape (Kihira et al. 2009; Fig. 2). In addition, the spire is higher, the spire angle is narrower and the shell apex is more pointed than those of C. c. laeta. Furthermore, C. c. laeta often has a lipped aperture, but the aperture of H. japonica does not have a pronounced lip (Masuda and Uchiyama 2004). Next, the differences with C. chinensis chinensis are not fully revealed, as the shell morphology of C. c. chinensis in Japan was not examined in depth. Nevertheless, based on the analysis and illustration of Hirano et al. (2019a), the only study that explicitly examined Japanese C. c. chinensis, the shell shape of C. c. chinensis is similar to that of C. c. laeta except for the pointed shell apex of C. c. chinensis. In any case, C. c. chinensis is distributed only around Kyushu Island, the south-western part of Japan (Hirano et al. 2019a) and no specimens of this subspecies were included in our study.

Figure 2.  

Representative shells of Heterogen japonica (Martens, 1861) from Japan, deposited in the collection of the National Museum of Nature and Science, Tokyo, which were treated as Viviparus sclateri (Frauenfeld, 1865) in Iwakawa (1919). a: 2959J from Lake Kasumigaura, Ibaraki Pref. b-g: specimens from Lake Suwa, Nagano Pref. (b: 2960J, c-e: 2961J, f-g: 2962J). h: 2963J from the western part of Aichi Pref. i: 2964J from Lake Biwa, Shiga Pref. j: 2966J from Hikone City, Shiga Pref. k: 2967J from Maibara City, Shiga Pref. l-m: 2968J from Tsukuma Lagoon, Shiga Pref. (l: adult, m: young).

Adult Sinotaia quadrata histrica is generally smaller than H. japonica. There is a high morphological diversity in the shell shape of S. q. histrica (Kihira et al. 2009); however, the body whorl and the spire whorl are more rounded and arched than that of H. japonica. In addition, the shell apex of S. q. histrica is more rounded and the aperture is relatively smaller than those of specimens of H. japonica having the similar shell shape, as the spire is dense. Moreover, the shell colour of S. q. histrica seems to be brighter than that in H. japonica and often has a yellowish colour.

H. japonica differs in shell shape from H. longispira (Hirano et al. 2019b). Quantitatively, H. japonica has a lower spire and a broader spiral angle than H. longispira (Kihira et al. 2009). In addition, H. longispira has a strong basal angulation even on the adult shell (Okada and Kurasawa 1950). Furthermore, the body whorl and the spire whorl of H. longispira are linear (like vertical) in lateral view and the upper periphery of the whorls turns sharply to nearly horizontal. Accordingly, the suture is quite deep and whorls have a strong shoulder just below the suture. This feature is extremely pronounced in juveniles and young shells and this morphological difference is diagnostic between two species (Okada and Kurasawa 1950, Hirano et al. 2019b). In addition, a difference that rarely appears in the shell shape is the distinct spiral ridges on the upper whorl of adult H. longspira (Hirano et al. 2019b). The early whorl of adult H. japonica does not have such spiral ridges. Moreover, Heterogen sp. has weak spiral ridges on the upper whorls and so this is a distinguishing feature from Heterogen sp., which is not distinguished by the shell shape (Hirano et al. 2019b). The spiral ridges of H. longispira are also pronounced on the body whorl of the adult shell. Furthermore, some H. japonica usually has greenish shells without reddish colour (e.g. Fig. 2h), whereas the shell colour of most H. longispira contains reddish colour and accordingly are often brown or dark brown in colour.

Besides, H. japonica has one morphotype, ver. iwakawa, which had been described and synonymised. This morphotype has a pyramidal shell shape, a broad spire angle and a strong basal angulation (e.g. Fig. 2d). Typical specimens of Viviparus japonicus var. iwakawa (Pilsbry 1902) is very easy to distinguish from any other Japanese Viviparidae; however, the morphology is continuous with H. japonica (Okada and Kurasawa 1950).

Notes

Heterogen japonica was identified in all 11 studied lots (Figs 2, 3). Some specimens (Fig. 2b-c, h and l) have the high spire and the narrow spire angle and the shell shapes of these specimens are relatively similar to H. longispira. However, specimens assigned to H. japonica lack a lot of the distinctive features of H. longispira. Lots Mo2969 and Mo2970 contained specimens with an intermediate morphology between H. japonica and H. longispira (e.g. Fig. 3c and g-h). Therefore, in this study, we treated only specimens that were clearly distinguished from H. longispira (e.g. Fig. 3a-b and l) as H. japonica.

Figure 3.  

Shells of Heterogen japonica (Martens, 1861), H. longispira (Smith, 1886), and the intermediate morphology between the two species stored at the National Museum of Nature and Science, Tokyo, which were treated as Viviparus sclateri (Frauenfeld, 1865) in Iwakawa's catalogue (1919). a-b: H. japonica in 2969J from Otsu City, Shiga Pref. c-d: non-typical H. longispira in 2969L from Otsu City, Shiga Pref. e: typical H. longispira in 2969L from Otsu City, Shiga Pref. f: subadult H. japonica in 2969J from Otsu City, Shiga Pref. g-j: non-typical and intermediate H. longispira in 2970L from the Seta River, Shiga Pref. K: typical H. longispira in 2970L from the Seta River, Shiga Pref. l: a juvenile shell of H. japonica in 2970J from the Seta River, Shiga Pref. m: a typical juvenile shell of H. longispira in 2970L from the Seta River, Shiga Pref.

Based on the previous taxonomy (Habe 1973), synonymy included species-group name, iwakawa (and its mandatory changes in spelling and incorrect subsequent spellings) in addition to species-group name, japonica (and its mandatory changes in spelling and incorrect subsequent spellings). Furthermore, since Paludina oxytropis in Kobelt (1879) and Vivipara oxytropis in Pilsbry (1895) were later synonymised by the authors themselves (Kobelt 1909, Pilsbry 1902), these were also listed in synonymy. As to the species-group names ingallsiana and sclateri, we refer to the discussion.

Heterogen longispira (Smith, 1886)

Nomenclature

Synonyms:

Materials   Download as CSV 

1. scientificName:
   Heterogen longispira (Smith, 1886)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Naga-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Shiga
   ; locality:
   Lake Biwa
   ; verbatimLocality:
   Biwa-ko (in Japanese) [Lake Biwa]
   ; year:
   before 1919
   ; individualCount:
   1
   ; lifeStage:
   adult
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2964L ex.NSMT-Mo 2964
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965305
   ; recordNumber:
   2964 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2964L
2. scientificName:
   Heterogen longispira (Smith, 1886)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Naga-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Shiga
   ; locality:
   Hikone, Hikone City
   ; verbatimLocality:
   Hikone (in Japanese) [Hikone, Hikone City]
   ; year:
   before 1919
   ; individualCount:
   2
   ; lifeStage:
   1adult 1 young
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2966L ex.NSMT-Mo 2966
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965303
   ; recordNumber:
   2966 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2966L
3. scientificName:
   Heterogen longispira (Smith, 1886)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Naga-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Shiga
   ; locality:
   Zeze, Ohtsu City
   ; verbatimLocality:
   Zeze (in Japanese) [Zeze, Ohtsu City]
   ; year:
   before 1919
   ; individualCount:
   6
   ; lifeStage:
   adults
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2969L ex.NSMT-Mo 2969
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965313
   ; recordNumber:
   2969 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; identificationRemarks:
   Some specimens are presumed to be hybrid species between H. longispira and H. japonica (see taxon discussion section).
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2969L
4. scientificName:
   Heterogen longispira (Smith, 1886)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Naga-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Shiga
   ; locality:
   Seta River
   ; verbatimLocality:
   Seta-gawa (in Japanese) [the Seta River]
   ; year:
   before 1919
   ; individualCount:
   15
   ; lifeStage:
   11 adults 4 juveniles
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2970L ex.NSMT-Mo 2970
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965314
   ; recordNumber:
   2970 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; identificationRemarks:
   Some specimens are presumed to be hybrid species between H. longispira and H. japonica (see taxon discussion section).
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2970L

Diagnosis

Adult shell moderately large and very thick; shell shape subconical. Adult shell dark brown or greenish-brown sometimes covered with attachments; shell surface often having growth lines and several spiral ridges. Suture deeper than in any other viviparid from Japan; whorls strongly shouldered. Young shell small and relatively thin, but not fragile; shell shape pyramidal to subconical. Young shell bright olive; shell surface quite glossy having strong spiral ridges on upper part of spire. The shell size upon birth is larger than that of any other viviparid gastropod in Japan.

Taxon discussion

Heterogen longspira can be distinguished from other Japanese Viviparidae by criteria based on the illustrations and information from literature (Suppl. material 1). In particular, the juvenile and young shell have strong spiral ridges on the upper part of each whorl, this feature having crucial diagnostic value (Okada and Kurasawa 1950, Hirano et al. 2019b). Furthermore, the shell size of the juvenile upon birth is larger than that of any other viviparid gastropods in Japan.

Firstly, H. longispira is easily distinguished from Cipangopaludina species/subspecies in Japan, based on the pyramidal shell shape, the higher spire, the narrower spire angle and the linear body whorl (Kihira et al. 2009; Fig. 4).

Figure 4.  

Representative shells of Heterogen longispira (Smith, 1886) from Japan, deposited in the collection of the National Museum of Nature and Science, Tokyo, which were treated as Viviparus sclateri (Frauenfeld, 1865) in Iwakawa (1919). a: 2964L from Lake Biwa, Shiga Pref. b: 2966L from Hikone City, Shiga Pref.

Besides, adult Sinotaia quadrata histrica is generally smaller than H. longispira. There is a high morphological diversity in the shell shape of S. q. histrica (Kihira et al. 2009); however, the body whorl and the spire whorl are much more rounded and arched than those of H. longispira. In addition, the suture of S. q. histrica is shallower than that of H. longispira. The upper whorl of adult S. q. histrica does not have the pronounced spiral ridges. Moreover, the shell colour of S. q. histrica is brighter than that in H. longispira and often has a yellowish colour.

Both adult H. longispira and Heterogen sp. have the spiral ridges on the upper whorl (Hirano et al. 2019b), but the ridges of H. longispira are stronger than Heterogen sp. In addition, the shell shape of Heterogen sp. is indistinguishable from that of H. japonica and then there are some differences in the shell shape between H. longispira and Heterogen sp. (Hirano et al. 2019b; refer to taxon discussion on the section of H. japonica for the difference).

Notes

Heterogen longispira was identified in four of the 11 lots examined, which were all from Lake Biwa drainage only (Figs 3, 4). Lots Mo2969 and Mo2970 contained non-typical H. longispira with an intermediate morphology to H. japonica and H. longispira (e.g. Fig. 3c and g-k). The shell shapes of these specimens tend to be slightly more similar to H. japonica; however, they have several distinctive morphological features of H. longispira, namely, the pronounced spiral ridges, the shouldered whorl, the deep suture, the strong basal angle and the high spire . Therefore these specimens were tentatively identified as H. longispira here (see also discussion).

Based on the previous taxonomy (Kuroda 1929), synonymy included the species-group name turris (and incorrect subsequent spellings) in addition to the species-group name longispira. For species-group names ingallsiana and sclateri, refer to the discussion.

Sinotaia quadrata histrica (Gould, 1859)

Nomenclature

Synonyms:

Materials   Download as CSV 

1. scientificName:
   Sinotaia quadrata histrica (Gould, 1859)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Hime-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Nagano
   ; locality:
   Lake Suwa
   ; verbatimLocality:
   Suwako (in Japanese) [Lake Suwa]
   ; year:
   before 1919
   ; individualCount:
   1
   ; lifeStage:
   adult
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2960H ex.NSMT-Mo 2960
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965306
   ; recordNumber:
   2960 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2960H
2. scientificName:
   Sinotaia quadrata histrica (Gould, 1859)
   ; kingdom:
   Animalia
   ; phylum:
   Mollusca
   ; family:
   Viviparidae
   ; vernacularName:
   Hime-tanishi (in Japanese name)
   ; country:
   Japan
   ; stateProvince:
   Nagano
   ; locality:
   Lake Suwa
   ; verbatimLocality:
   Suwako (in Japanese) [Lake Suwa]
   ; year:
   before 1919
   ; individualCount:
   5
   ; lifeStage:
   adults
   ; preparations:
   shell (dried)
   ; catalogNumber:
   2961H ex.NSMT-Mo 2961
   ; occurrenceDetails:
   https://www.gbif.org/occurrence/2864965308
   ; recordNumber:
   2961 in Iwakawa 1919
   ; previousIdentifications:
   Viviparus sclateri (Frauenfeld, 1865) in Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; identifiedBy:
   Takumi Saito
   ; dateIdentified:
   2020-02-03/04
   ; language:
   Japanese
   ; bibliographicCitation:
   Tomotaro Iwakakawa. 1919. Catalogue of Japanese Mollusca in the Natural history department, Tokyo Imperial Museum, the tokyo Imperial Museum, Tokyo, Japan, p. 72.
   ; institutionCode:
   NSMT
   ; collectionCode:
   Mo
   ; basisOfRecord:
   PreservedSpecimen
   ; occurrenceID:
   paper:SK2020:2961H

Diagnosis

Adult shell small and thin, but not fragile; shell shape subconical to pyramidal. Adult shell bright brown or yellowish- or reddish-brown often covered with many attachments such as alga; shell surface slightly glossy, weak growth lines and spiral lines are usually present. Suture shallow; spire rounded. Young shell small, very thin and fragile; shell shape pyramidal with rounded angle. Young shell bright olive; shell surface quite glossy with no spiral ridges on upper part of the spire.

Taxon discussion

Sinotaia quadara histrica can be distinguished from other Japanese Viviparidae by the following features, based on the illustrations and information from literature (Suppl. material 1). In particular, the adult shell width tends to be relatively smaller than that of other Japanese Viviparidae (Masuda and Uchiyama 2004; for example, under 24 mm except for one exception in Kagawa et al. (2019)).

Firstly, S. q. histrica is easily distinguished from Cipangopaludina species/subspecies in Japan, based on the small shell width, pyramidal shell shape and the higher spire (Fig. 5).

Figure 5.  

Representative shells of Sinotaia quadrata histrica (Gould, 1859) from Japan, deposited in the collection of the National Museum of Nature and Science, Tokyo, which were treated as Viviparus sclateri (Frauenfeld, 1865) in Iwakawa (1919). a-c: Lake Suwa, Nagano Pref. (a: 2960H, b-c: 2961H).

The size of Heterogen sp. is larger and the apical whorls are also larger than those of S. q. histrica. In addition, Heterogen sp. has the weak spiral ridges on even the early whorls, whereas S. q. histrica does not have them. Diagnostic differences between S. q. histrica vs. H. japonica and H. longispira have been provided above, so we refer to the taxon discussions of these latter two species.

Notes

Sinotaia quadara histrica was identified in two of the 11 lots examined, which were from Lake Suwa (Fig. 5). Some specimens of H. japonica from Lake Suwa have a similar shell shape to S. q. histrica, but S. q. histrica has a more rounded shell apex, a broader spire angle, shallower sutures and a brighter colour than those of H. japonica.

We treated only the Japanese Sinotaia species for synonymy, as the taxonomic relationship between the continental Sinotaia and the Japanese Sinotaia species is not clear and not the point of this study. The species-group name, nitens was synonymised with a question mark by Pilsbry (1902) and we followed this treatment. Vivipara lacustris in Kawamura (1918) was documented as a small species that is distributed in the Kyushu Region. In the past, S. q. histrica was considered to be distributed in Kyushu Region (Pilsbry 1902) and so V. lacustris in Kawamura 1918 may be a junior synonym of S. q. histrica.

Analysis

The principal results of the PCA of EF analysis are shown in Fig. 6 and Fig. 7 (for full results of the PCA and SW, see Suppl. material 2). PC1 and PC2 explained 74.6% of the variance and the first five components explained 86.7% of the variance (PC1: 59.5%; PC2: 15.1%; PC3: 6.3%; PC4: 3.0%; PC5: 2.7%). Heterogen japonica and H. longispira from Hirano et al. (2019b) had little overlap in each region and Heterogen sp. was fully included in the morphological range of H. japonica. Many of the H. japonica in Iwakawa's collection identified qualitatively by diagnostic features were within the morphological range of H. japonica from Hirano et al. (2019b), but some were located within the range of H. longisira from Hirano et al. (2019b) or in parts of the morphospace not covered by either. Some of H. longispira in Iwakawa's collection, which were identified qualitatively, were within the morphological range of H. longispira from Hirano et al. (2019b), but some were located within the morphological range of H. japonica from Hirano et al. (2019b) or in parts of the morphospace not covered by either.

Figure 6.  

a: Plots of principal component analysis (PCA) of all analysed specimens. The top left graph shows the proportion of variance of each component. Figures on the background indicate reconstructed shell morphology, based on principal components. b: Plots of PCA of Iwakawa's specimens and old literature specimens. The morphospace occupation by specimens from Hirano et al. (2019b) is shown as coloured polygons. The top left graph shows the proportion of variance of each component. Background outlines indicate the reconstructed shell morphology, based on principal components.

Figure 7.  

Plots of principal component axis one versus shell width for specimens from Iwakawa and Hirano et al. (2019b). The morphospace occupation by specimens from Hirano et al. (2019b) is also shown as coloured polygons.

Discussion

Acknowledgements

Author contributions

References

Anma K, Nagaoka M, Niwa S, Sekimoto K, Yoshikawa M, Fujine H (1990)
A geological survey for the study on Neotectonic movement and geoenvironment of Lake Suwa, Nagano Prefecture
.
Memoirs of the Geological Society of Japan
36
:
179
‑
194
. [In
Japanese with English abstract
].
Annandale N (1916)
Zoological results of a tour in the Far East. The Mollusca of Lake Biwa, Japan.
Memoirs of the Asiatic Society of Bengal
6
(
1
):
39
‑
74
. URL: https://biodiversitylibrary.org/page/47681217
Annandale N (1921)
Zoological results of a tour in the Far East. The viviparous water-snail of Lake Biwa, Japan
.
Memoirs of the Asiatic Society of Bengal
6
(
7
):
397
‑
401
. URL: https://biodiversitylibrary.org/page/47681678
Annandale N (1922)
The macroscopic fauna of Lake Biwa
.
Annotationes Zoologicae Japonenses
10
:
127
‑
153
.
Bonhomme V, Picq S, Gaucherel C, Claude J (2014)
Momocs: Outline analysis using R
.
Journal of Statistical Software
56
(
13
):
1
‑
24
. https://doi.org/10.18637/jss.v056.i13
Frauenfeld GRV (1865)
Zoologische Miscellen. IV
.
Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien
15
:
259
‑
266
. URL: https://www.biodiversitylibrary.org/page/16400700
Gould AA (1859)
Dr. A. A. Gould read the following descriptions of new species of shells
.
Proceedings of the Boston Society of Natural History
7
:
40
‑
45
. URL: https://www.biodiversitylibrary.org/page/9249577
Habe T, Kosuge S (1967)
A standard illustrated handbook in natural color. III. Mollusca.
Hoiku-sya
,
Osaka
,
223
pp. [In
Japanese
].
Habe T (1973)
Mollusca
. In: Ueno M (Ed.)
The late Tamiji Kawamura freshwater biology of Japan, enlarged and revised edition
.
Hokuryu-kan
,
Tokyo
,
309-341
pp. [In
Japanese with English title
].
Habe T (1990)
The list of Japanese freshwater mollusks: 1
.
Hitachiobi
54
:
3
‑
6
. [In
Japanese
].
Hannibal H (1912)
A synopsis of the recent and tertiary freshwater Mollusca of the Californian province, based upon an ontogenetic classification
.
Proceedings of the Malacological Society of London
10
:
112
‑
211
. URL: https://www.biodiversitylibrary.org/page/15237075
Hirano T, Saito T, Chiba S (2015)
Phylogeny of freshwater viviparid snails in Japan
.
Journal of Molluscan Studies
81
(
4
):
435
‑
441
. https://doi.org/10.1093/mollus/eyv019
Hirano T, Saito T, Tsunamoto Y, Koseki J, Ye B, Do VT, Miura O, Suyama Y, Chiba S (2019a)
Enigmatic incongruence between mtDNA and nDNA revealed by multi-locus phylogenomic analyses in freshwater snails
.
Scientific Reports
9
(
1
). https://doi.org/10.1038/s41598-019-42682-0
Hirano T, Saito T, Tsunamoto Y, Koseki J, Prozorova L, Do VT, Matsuoka K, Nakai K, Suyama Y, Chiba S (2019b)
Role of ancient lakes in genetic and phenotypic diversification of freshwater snails
.
Molecular Ecology
28
(
23
):
5032
‑
5051
. https://doi.org/10.1111/mec.15272
Hirase S (1927)
Mollusca
. In: Uchida S (Ed.)
An illustrated encyclopedia of the fauna of Japan
.
Hokuryu-kan
,
Tokyo
. [In
Japanese
].
Hirase S, Kuroda T (1947a)
Cipangopaludina japonica iwakawa (Pilsbry)
. In: Uchida S (Ed.)
An illustrated encyclopedia of the fauna of Japan, revised ver.
Hokuryu-kan
,
Tokyo
,
1162-1162
pp. [In
Japanese
].
Hirase S, Kuroda T (1947b)
Viviparus (Sinotaia) histricus (Gould)
. In: Uchida S (Ed.)
An illustrated encyclopedia of the fauna of Japan, revised ver.
Hokuryu-kan
,
Tokyo
,
1160-1160
pp. [In
Japanese
].
Hirase S, Kuroda T (1947c)
Cipangopaludina japonica (Martens)
. In: Uchida S (Ed.)
An illustrated encyclopedia of the fauna of Japan, revised ver.
Hokuryu-kan
,
Tokyo
,
1161-1161
pp. [In
Japanese
].
Hirase S, Taki Is (1951)
A handbook of illustrated shells in natural colors from Japanese Islands and their adjacent territories
.
Bunkyo-kaku
,
Tokyo
,
134 pls and 43
pp. [In
Japanese with English title
].
Hirase Y (1909)
Introduction book of Mollusks
.
Hirase-kaikan
,
Kyoto
,
112
pp. [In
Japanese
]. URL: https://dl.ndl.go.jp/info:ndljp/pid/993608
Hirase Y (1910)
One thousand kinds of shells existing in Japan
.
Hirase-kaikan
,
Kyoto
,
48
pp. [In
Japanese
]. URL: https://dl.ndl.go.jp/info:ndljp/pid/994074
Horie S (1971)
Ancient inland waters in the world and Lake biwa as the natural heritage
. In: The scientific investigation committee of Lake Biwa Quasi-national park (Ed.)
The scientific report of Lake Biwa Quasi-national park
.
The scientific investigation committee of Lake Biwa Quasi-national Park
,
Otsu
,
491-493
pp. [In
Japanese
].
Horikoshi M, Itabashi Y (1994)
Republication of "List of molluscs collected from Yokohama and its environs" compiled by the late Dr. Isao Taki in 1933 part 1
.
Natural History Report of Kanagawa
(16)
:
17
‑
28
. URL: http://nh.kanagawa-museum.jp/files/data/pdf/nhr/16/nhr16_017_028horikoshi.pdf
Iwakawa T (1895)
Freshwater mollusks in Japan (2)
.
Zoological Magazine
7
(
86
):
411
‑
414
. [In
Japanese
]. URL: https://dl.ndl.go.jp/view/download/digidepo_10825080_po_ART0003837109.pdf?contentNo=1&alternativeNo=
Iwakawa T (1897a)
Freshwater mollusks in Japan (2), with plate 2
.
Zoological Magazine
9
(
99
):
5
‑
10
. [In
Japanese
]. URL: https://dl.ndl.go.jp/view/download/digidepo_10825396_po_ART0003840054.pdf?contentNo=1&alternativeNo=
Iwakawa T (1897b)
Notes on the Paludina-species of Japan
.
Annotationes Zoologicae Japonenses
1
:
83
‑
92
. URL: https://dl.ndl.go.jp/view/download/digidepo_10852805_po_ART0003842255.pdf?contentNo=1&alternativeNo=
Iwakawa T (1919)
Catalogue of Japanese Mollusca in the natural history department, Tokyo Imperial Museum
.
Tokyo Imperial Museum
,
Tokyo
,
375
pp. [In
Japanese
]. https://doi.org/10.5962/bhl.title.13147
Kagawa O, Saito T, Uchida S, Chiba S (2019)
Phenotypic divergence in viviparid snails in a recently converted freshwater lagoon
.
Plankton and Benthos Research
14
(
3
):
189
‑
196
. https://doi.org/10.3800/pbr.14.189
Kanamaru T (1920)
An investigation report of Mollusca in Mie Prefecture
.
Mie Prefecture
,
Tsu
,
17
pp. [In
Japanese
].
Kawamura T (1918)
Japanese freshwater biology
.
I
.
Shoka-bo
,
Tokyo
,
362
pp. [In
Japanese
]. https://doi.org/10.11501/956620
Kawanabe H (1996)
Asian great lakes, especially Lake Biwa
.
Environmental Biology of Fishes
47
(
3
):
219
‑
234
. https://doi.org/10.1007/bf00000495
Kihira H, Matsuda M (1990)
Freshwater mollusks in Lake Biwa and the Yodo River
.
Tatara-syobo
,
Hirakata
,
131
pp. [In
Japanese
].
Kihira H, Matsuda M, Uchiyama R (2003)
Freshwater Molluscs of Japan 1: Freshwater Molluscs of Lake Biwa and the Yodo River
.
Pisces
,
Tokyo
,
159
pp. [In
Japanese
].
Kihira H, Matsuda M, Uchiyama R (2009)
Freshwater Molluscs of Japan 1: Freshwater Molluscs of Lake Biwa and the Yodo River, revised ver.
Pisces
,
Tokyo
,
159
pp. [In
Japanese
].
Kitahara T (1895)
Animals in Lake Kasumigaura
.
Zoological Magazine
7
(
77
):
87
‑
90
. [In
Japanese
]. URL: https://dl.ndl.go.jp/view/download/digidepo_10824909_po_ART0003801509.pdf?contentNo=1&alternativeNo=
Kobelt W (1879)
Fauna molluscorum extramarinorum Japoniae, nach den von Professor Rein gemachten Sammlungen.
Christian Winter
,
Frankfurt a. M.
,
171
pp. https://doi.org/10.5962/bhl.title.12881
Kobelt W (1909)
Die Gattung Paludina Lam. (Vivipara Montfort) : neue Folge
. In: Martini FHW, Chemnitz JH (Eds)
Sytematisches Conchylien-Cabinet
.
Bd. 1 Abt. 21a
.
Verlag von Bauer und Raspe
,
Nürnberg
,
98-430
pp. https://doi.org/10.5962/bhl.title.119758
Köhler F, Rintelen T (2011)
Heterogen longispira
.
IUCN Red List of Threatened Species
https://doi.org/10.2305/iucn.uk.2011-2.rlts.t189576a8750326.en
Köhler F, Do V, Jinghua F (2012)
Cipangopaludina chinensis.
IUCN Red List of Threatened Species
https://doi.org/10.2305/iucn.uk.2012-1.rlts.t166265a1124988.en
Kuhl FP, Giardina CR (1982)
Elliptic Fourier features of a closed contour
.
Computer Graphics and Image Processing
18
:
236
‑
258
. https://doi.org/10.1016/0146-664X(82)90034-X
Kurasawa H (1988)
Pass down the Tenryu River, 9: A eutrophication in Lake Suwa and the biological community response
.
The Ministry of Construction of Japan, Construction Bureau in the Chubu region, the work office of the upper Tenryu River
,
Komagane
,
67
pp. [In
Japanese
].
Kuroda T (1928)
A catalogue of mollusks in Amami-Oshima
.
Kagoshima Prefecture
,
Kagoshima
,
120
pp. [In
Japanese
].
Kuroda T (1929)
Notes on the nomenclature of the Japanese species of the genus Viviparus
.
Venus
1
(
3
):
98
‑
102
. [In
Japanese
]. https://doi.org/10.18941/venusomsj.1.3_98
Kuroda T (1947a)
Freshwater mollusks of Lake Biwa (2)
.
Yume-hamaguri
(20)
:
2
‑
15
. [In
Japanese
].
Kuroda T (1947b)
Classified list of freshwater shells from the Japanese Islands
.
Mokuhachi-tengu-sya
,
Kyoto
,
22
pp. [In
Japanese with English title
].
Kuroda T (1948)
A list of mollusks in Lake Biwa
.
Yume-hamaguri
(22)
:
26
‑
27
. [In
Japanese
].
Kuroda T (1955)
A list of freshwater molluscan shells of Japan
.
Yume-hamaguri
(80)
(
Supplement
):
1
‑
12
. [In
Japanese with English title
].
Kuroda T (1963)
A catalogue of the non-marine mollusks of Japan, including the Okinawa and Ogasawara Islands.
The Malacological Society of Japan
,
Tokyo
,
70
pp. [In
Japanese with English title
].
Kuroda T, Habe T (1965a)
Cipangopaludina (Heterogen) longispira (SMITH)
. In: Okada Y, Uchida S, Uchida T (Eds)
New illustrated encyclopedia of the fauna of Japan (vol. 2)
.
Hokuryu-kan
,
Tokyo
,
48
pp. [In
Japanese with English title
].
Kuroda T, Habe T (1965b)
Sinotaia quadratus histrica (GOULD)
. In: Okada Y, Uchida S, Uchida T (Eds)
New illustrated encyclopedia of the fauna of Japan (vol. 2)
.
Hokuryu-kan
,
Tokyo
,
48
pp. [In
Japanese with English title
].
Kurozumi T (2001)
Japanese crested ibis and Viviparid gastropods —Animals showing the transition of rice field
.
Birder
15
(
4
):
30
‑
33
. [In
Japanese
].
Kurozumi T (2019)
Seeking unspoilt landscape in Japan using shells
. In: Toyokawa K, Tomoda A, Yura H (Eds)
30th anniversary book of the Natural History Museum and Institute, Chiba
.
Chiba
,
45-46
pp. [In
Japanese
]. URL: http://www2.chiba-muse.or.jp/www/NATURAL/contents/1549010275781/simple/shell.pdf
Lake Biwa fisheries experimental station (1915)
Fisheries research report of Lake Biwa (2): Mollusks of Lake Biwa
.
Lake Biwa fisheries experimental station
,
Fukuman [Hikone]
,
36
pp. [In
Japanese
]. https://doi.org/10.11501/928661
Masuda O, Uchiyama R (2004)
Freshwater mollusks of Japan 2: freshwater mollusks of Japan, including brackish water species.
Pisces
,
Tokyo
,
240
pp. [In
Japanese
].
Matsuoka K (2001)
Freshwater mollusks in irrigation ponds
.
Nature of irrigation pond
34
:
9
‑
11
. [In
Japanese
].
Ministry of the Environment, Government of Japan (2019)
Red List of Ministry of the Environment, 2019
.
Ministry of the Environment, Government of Japan
,
Tokyo
,
129
pp. [In
Japanese
]. URL: https://www.env.go.jp/press/files/jp/110615.pdf
Nakai K (2016)
Heterogen longispira
. In: Scientific Committee for Research into the Wildlife in Shiga Prefecture (Ed.)
Red Data Book of Shiga Prefecture 2015
.
Sunrise Publishing
,
Hikone
,
583-583
pp. [In
Japanese
].
Nishino M (1991)
Benthos in Lake Biwa —Wildlife of the shores. Mollusca.
Lake Biwa Research Institute, Shiga Prefecture
,
Otsu
,
46
pp. [In
Japanese
].
Nishiwaki S, Masu H, Hanawa T (2008)
Seasonal and regional variations in the number of shelled embryos of Sinotaia quadrata histrica in Lake Kasumigaura, Japan
.
Venus
67
(
1-2
):
73
‑
80
. [In
Japanese with English abstract
]. https://doi.org/10.18941/venus.67.1-2_73
Okada Y, Kurasawa H (1950)
A study of Viviparidae in Japan
. In: Okada Y (Ed.)
Studies of fishery animals (1)
.
Japan Publishing Cooperation
,
Tokyo
,
141-165
pp. [In
Japanese
]. https://doi.org/10.11501/2470294
Okuda N, Watanabe K, Fukumori K, Nakano S, Nakazawa T (2013)
Origin and diversification of freshwater fishes in Lake Biwa
.
Springer Briefs in Biology
1
‑
19
. https://doi.org/10.1007/978-4-431-54150-9_1
Oyama K, Kajiyama H (1959)
The relationship between Heterogen turris and Cipangopaludina japonica (a preliminary note)
.
Yume-hamaguri
(98)
:
1
‑
4
. [In
Japanese
].
Ozawa S (2012)
A study of the relationship between the designation of Lake Biwa as a Quasi-national Park and concurrent reclamation work on lagoons around the Lake
.
Landscape Research Japan Online
5
:
5
‑
16
. [In
Japanese with English title and abstract
]. https://doi.org/10.5632/jilaonline.5.5
Pilsbry HA (1895)
Appendix I. List of land and freshwater mollusks collected in Japan by Frederick Stearns
. In: Pilsbry HA (Ed.)
Catalogue of the marine mollusks of Japan, with descriptions of new species and notes on others collected by Frederick Stearns
.
Frederick Stearn
,
Detroit
,
155-159
pp. https://doi.org/10.5962/bhl.title.32672
Pilsbry HA (1902)
Revision of Japanese Viviparidae, with notes on Melania and Bithynia
.
Proceedings of the Academy of Natural Sciences of Philadelphia
54
(
1
):
115
‑
121
. URL: https://www.jstor.org/stable/4062827
Prashad B (1928)
Recent and fossil Viviparidae. A study in distribution, evolution, paleogeography
.
Memoirs of the Indian Museum
8
(
4
):
153
‑
253
.
R Core Team (2018)
R: A language and environment for statistical computing
.
3.5.1
.
R Foundation for Statistical Computing, Vienna, Austria
. URL: https://www.R-project.org/
Reeve LA (1863)
Monograph of the genus Paludina. Pl. X.
In: Reeve LA (Ed.)
Conchologia Iconica: or, Illustrations of the Shells of Molluscous Animals
.
14
.
Lovell Reeve
,
London
. URL: https://www.biodiversitylibrary.org/page/10971864
Rossiter A (2000)
Lake Biwa as a topical ancient lake
.
Advances in Ecological Research
31
:
571
‑
598
. https://doi.org/10.1016/s0065-2504(00)31031-5
Smith EA (1886)
Descriptions of three new species of freshwater shells from Japan
.
Journal of Conchology
5
:
57
‑
59
. URL: https://biodiversitylibrary.org/page/31653661
Stelbrink B, Richter R, Köhler F, Riedel F, Strong EE, Van Bocxlaer B, Albrecht C, Hauffe T, Page TJ, Aldridge DC, Bogan AE, Du L, Manuel-Santos MR, Marwoto RM, Shirokaya AA, von Rintelen T (2020)
Global diversification dynamics since the Jurassic: low dispersal and habitat-dependent evolution explain hotspots of diversity and shell disparity in river nails (Viviparidae)
.
Systematic Biology, in press.
https://doi.org/10.1093/sysbio/syaa011
Suarez A, Tsutsui N (2004)
The value of museum collections for research and society
.
BioScience
54
(
1
). https://doi.org/10.1641/0006-3568(2004)054[0066:tvomcf]2.0.co;2
Taki Is (1933)
The life of Prof. Iwakawa Tomotaro and a list of his papers on conchology (with a portrait)
.
Venus
4
(
2
):
63
‑
69
. [In
Japanese
]. https://doi.org/10.18941/venusomsj.4.2_63
Taki Is (1946)
The freshwater mollusks in Musashino
.
Collecting and breeding
8
(
11
):
202
. [In
Japanese
].
Tanba K, Shibata S, Takamatsu K (1883)
General Zoology
.
Shimamura Toshisuke, Maruya-zenshiti, and Tyu-kin-do
,
Tokyo
,
397
pp. [In
Japanese
]. https://doi.org/10.11501/832886
Tanba K, Shibata S, Takamatsu K (1891)
General Zoology, revised ver.
Shimamura Toshisuke, Maruzen, Nanko-do, and Umehara Kameshichi
,
Tokyo
,
336
pp. [In
Japanese
]. https://doi.org/10.11501/832887
Van Bocxlaer B, Strong E (2016)
Anatomy, functional morphology, evolutionary ecology and systematics of the invasive gastropod Cipangopaludina japonica (Viviparidae: Bellamyinae)
.
Contributions to Zoology
85
(
2
):
235
‑
263
. https://doi.org/10.1163/18759866-08502005
Van Bocxlaer B, Strong EE (2019)
Viviparidae, Gray 1847
. In: Lydeard C, Cummings KS (Eds)
Freshwater mollusks of the world: a distribution atlas
.
Johns Hopkins University Press
,
Baltimore
,
43-50
pp.
von Martens E (1861)
Die Japanesischen Binnenschnecken im Leidner Museum
.
Malakozoologische Blätter
7
:
32
‑
61
. URL: https://biodiversitylibrary.org/page/15919572
Wandeler P, Hoeck PA, Keller L (2007)
Back to the future: museum specimens in population genetics
.
Trends in Ecology & Evolution
22
(
12
):
634
‑
642
. https://doi.org/10.1016/j.tree.2007.08.017
Wickham H (2016)
ggplot2: Elegant graphics for data analysis
.
Springer-Verlag New York
. URL: https://ggplot2.tidyverse.org
Yagura H (1935)
A research of viviparid gastropods and land snails
.
Bulletin of the Society for Hyogo Natural History
(9)
:
7
‑
13
. [In
Japanese
].
Yagura W (1932)
A catalogue of Mollusca in Hyogo, new and revised ver.
Konan-Kairui-so
,
Nishinomiya
. [In
Japanese
]. https://doi.org/10.11501/1903884
Ye B, Saito T, Hirano T, Dong Z, Do VT, Chiba S (2020)
Human‐geographic effects on variations in the population genetics of Sinotaia quadrata (Gastropoda: Viviparidae) that historically migrated from continental East Asia to Japan
.
Ecology and Evolution
10
(
15
):
8055
‑
8072
. https://doi.org/10.1002/ece3.6456
Yokoyama T (1984)
Stratigraphy of the Quaternary system around Lake Biwa and geohistory of the ancient Lake Biwa
. In: Horie S (Ed.)
Lake Biwa, Monographiae Biologicae
.
54
.
Dr. W Junk Publishers
,
Dordrecht
,
43-128
pp.

Supplementary materials