Flammulinini Crosse, 1895

Mahlfeld, Karin, Climo, Frank & Roscoe, David, 2025, Systematics, conservation status, and biogeography of 16 new species of Sinployea Solem, 1983 (Gastropoda: Charopidae) from New Zealand, Zoological Journal of the Linnean Society 204 (1) : -

Tribe Flammulinini Crosse, 1895

Remarks: The subfamily Flammoconchinae Schileyko, 2001 was proposed for two NZ genera, Flammoconcha and Flammocharopa Climo, 1970 (now Calymna ), on the basis of a single character, a large glandular appendage on the terminal male reproductive tract. Schileyko (2001: 1024) wrote: ‘Two New Zealand genera [ Flammocharopa and Flammoconcha ] differ from any other charopid genera by the conspicuous (glandular?) penial appendage’. Our studies show that the use of the penial appendage to define a subfamily is problematic. Some undescribed species of Flammoconcha (F.C. obs.), in addition to Calymna costulata Hutton, 1883 and Calymna montana ( Suter, 1891) , have a basal penial appendage, but none of the Flammoconcha species dissected and illustrated by Climo (1969) does. Powell (1941) and Dell (1952) did not illustrate the male reproductive systems of Flammoconcha cumberi ( Powell, 1941) or Flammoconcha stewartensis Dell, 1952 . Schileyko (2001) did not give any other source of information for his statement that a

Specimen First teleoconch Second teleoconch Rib count D/U Maximal shell Shell Shell whorl width whorl width first teleoconch ratio diameter height whorls whorl

NZ Sinployea

S. kaipara Okahukura 325 μm 563 μm Peninsula

(M.082966)

S. kaipara Thomson 349 μm

Kauri Grove ScR

(M.075729)

S. solemi? Ohinau Island 347 μm

S. accelerata (Climo) 301 μm 556 μm Motukaramarama

S. accelerata (Climo) ? Gut 308 μm 534 μm Rock

S. accelerata (Climo) ? 366 μm 656 μm Ruamahuaiti

S. ohinepoutea Puketi 449 μm

(M.165180)

S. solemi ?

Ngunguru (M.058119)

S. mangamuka 404 μm 669 μm Mangamuka (M.062484)

S. karangahake Hotwater 333 μm 538 μm Beach (M.076163)

S. karangahake 417 μm 787 μm Karangahake (M.103006)

S. fiordlandica Mount 280 μm 506 μm Troup (M.081649)

S. australis Invercargill 316 μm 612 μm (M.078966)

S. waipoua Waipoua 411 μm

(M.098308)

S. waipoua Parakao 393 μm

S. capensis Cape Sanc- 269 μm 539 μm tuary (M.151454)

S. capensis 258 μm 579 μm Cape Sanctuary

(MA73638)

S. capensis 270 μm 569 μm Cape Sanctuary

(MA73639)

S. ohinepoutea 287 μm 580 μm Ohinepoutea

(M.078239)

S. solemi Unuwhao 337 μm

(MA96903)

S. solemi Unuwhao 343 μm

(MA96903)

S. solemi Waterfall Gully 311 μm 590 μm (MA96904)

S. solemi Waterfall Gully 281 μm 536 μm (MA96904)

S. solemi Pandora Road 367 μm

(MA152583)

67 4.45 2.2 mm 1.22 mm 3

75 3.86 1.83 mm 1.1 mm ~3

56 ~4.75 ~ 1.7 mm ~905 μm ~2.8 65 3.99 3.0 mm ~ 1.7 mm ~3.75

66 4.11 2.5 mm ~ 1.3 mm ~3.5

42 3.93 2.7 mm ~ 1.4 mm ~3.4

65 4.36 1.5 mm 830 μm 2.5

56 ~4.06 ~ 1.3 mm 2.25

49 4.99 2.3 mm 1.28 mm 3

66 4.57 2.7 mm 1.58 mm 3.25

56 3.96 2.4 mm 1.28 mm 3.1

102 4.17 2.1 mm 950 μm ~3

78 3.92 2 mm 1.09 mm ~3

85 ~4.90 ~2.0 mm ~2.75

92 ~4.72 1.5 mm ~2.5 32 5.02 2.3 mm 1.21 3.25

33 5.24 1.87 mm ~937 μm ~3

33 5.35 1.9 mm 935 μm 3

59 4.15 1.95 mm ~933 μm ~3

42 5.53 1.4 mm 849 μm 2.6

45 6.84 1.6 mm 960 μm 2.7

67 5.47 1.7 mm 965 μm 3

65 6.05 1.7 mm 974 μm 3

50 5.60 1.5 mm 856 μm 2.5 large penial appendage is unique to Calymna and Flammoconcha . Our dissections show that Flammulina jacquenetta ( Hutton, 1883) , Charopidae , also has a basal glandular appendage, as has Sinployea capensis ( Figs 3, 4). Two further dissections of undescribed Calymna species, Charopidae sp. 37 (NMNZ M.075855) and Charopidae sp. 38 (NMNZ M.029851), have pocket pilasters without a basal penial appendage like the Pacific Sinployea species ( Fig. 5). The basal glandular appendage on the male terminal reproductive tract of Sinployea accelerata ( Climo, 1970) comb. nov. is depicted in Figure 6 for further comparisons. The reproductive system of Calymna costulata is shown in Figure 4 and that of Calymna montana ( Suter, 1891) in the paper by Climo (1970: fig. 16). The dissections suggest two possibilities: homology of external appendage and internal pocket pilaster, and a more complex evolutionary history than that implied by Schileyko’s subfamily Flammoconchinae . For example, the penial pilasters of Charopidae sp. 37 depicted in Figure 5B appear to be at an intermediate stage of internalization of an appendage. Pustules occur in the upper chamber of the penis of Flammulina jacquenetta ( Fig. 3), as they do in Charopa coma (Gray, 1843) ( Fig. 7) and Flammulina zebra ( Le Guillou, 1842) as illustrated by Solem (1983: fig. 29e).

The male genitalia of the more tightly coiled Charopinae with higher whorl count like Charopa coma are different. Solem (1983: 73 and 82) considered the male genitalia of Sinployea to be of ‘ Charopa type’ but also comparable to those of Flammulina , but we find that the reproductive anatomy of Charopa Albers, 1860 is more different from Sinployea (currently classified in Charopinae ) than the latter from Calymna ( Flammoconchinae sensu Schileyko ). The male genitalia of Polynesian and Micronesian Sinployea species and Charopa coma are illustrated in Figures 7–9.

The male genitalia of Pacific islands Sinployea species have a standard pattern of apical or near-apical insertion of the vas deferens into the epiphallus, with the epiphallus inserting apically into the penial lumen through a verge or papilla. The retractor muscle attaches apically near the junction of the epiphallus and phallus, and the outline of the penis often appears to show a constriction, marked internally by a circular pilaster. A pocket pilaster occurs in the internal lumen of most of the dissected species ( Figs 7, 9). The internal male lumen of Charopa coma also shows a circular pilaster or sphincter, a pocket pilaster, and an epiphallic papilla, but the pilasters are predominantly pustulose, and the pocket pilaster is situated above the sphincter. The cluster of pilasters adjacent to the entrance of the epiphallus below the retractor muscle attachment in Charopa coma appears to be a separate structure ( Fig. 7). These small pilasters possibly represent a reduced caecum similar to that seen in the charopid species Corinomala gressitti ( Solem, 1970) from New Guinea ( Solem 1970: 250, fig. 2) and charopid Cavellia colensoi ( Suter, 1890) from NZ (dissected specimen from Pongaroa, Tararua District; Mahlfeld 2005 and auth. obs.). No such structure nor pustulose pilaster structures are known from Sinployea or Calymna dissections so far.

The dissections of Calymna ( Figs 4–6) species illustrate the different degrees of reduction and internalization of an external appendage to an internalized remnant caecum structure or pocket pilaster or to even further simplified lumen pilasters. The trend to internalization of structures and vertical compaction of reproductive morphologies is particularly clear in the very loosely coiled Flammulina E. von Martens, 1873 taxa and Ranfurlya Suter, 1903 (or Climo 1969: figs 11–13; and our pers. obs.). Solem (1983: 61) discussed this trend in reference to a dissection of Flammulina zebra : ‘In Flammulina zebra , the genotype, the epiphallus has been compacted back into the penis … and is thus completely undetectable from external view’. Similar trends have been found in other NZ charopine snails, which suggests a general pattern of structurally defined and ordered evolutionary pathways from external appendages to internalized, reduced structures involving internalization and reduction of radially branched structures through telescopic contraction and compaction (tubes within tubes), and reduction of structures to simple pilasters or, eventually, no pilasters at all in the penial lumen ( Fig. 5B; Solem 1970). Dissections of Allodiscus Pilsbry, 1892 and Phenacohelix Suter, 1892 species have likewise shown complex arrangements and fusions of appendages (auth. obs.), sometimes involving several appendages. Different stages of fusion of the epiphallic papilla with the internal phallic lumen wall and various stages of reduced pocket pilasters and caeca have also been observed in dissections of species in the charopid genus Cavellia Iredale, 1915 (Mahlfeld 2005). Moreover, pilaster structures seem to be interchangeable between male and female systems, for instance ‘valve-like’ structures or pores.

To summarize, Schileyko (2001) included two NZ genera, Calymna and Flammoconcha , in the new subfamily Flammoconchinae , while retaining other taxa with similar morphological characters in Charopinae or Flammulininae . However, the glandular penial appendage supposedly defining Flammoconchinae does not occur in all Flammoconcha , but it does occur in other Charopidae species that Schileyko (2001) did not include in Flammoconchinae . Thus the subfamily Flammoconchinae sensu Schileyko (2001) is not accepted here. Instead, we suggest treating Flammoconchinae as a junior synonym of Flammulinini and assigning Sinployea , Calymna , Flammoconcha , Ranfurlya , and Flammulina to this tribe in the subfamily Charopinae . This is based on the trends observed in the reproductive morphologies of different NZ charopid genera ( Figs 3–9). A recently published punctoid phylogeny ( Salvador et al. 2020) proposed a clade including Flammulina , Ranfurlya , Charopa , and Sinployea based on 26 of ≥300 NZ charopid species. Flammulinini as a tribal group in Charopinae would be in agreement with results of Salvador et al. (2020).