Evolution of Janthina and Recluzia (Mollusca: Gastropoda: Epitoniidae)
Author
Beu, Alan G.
text
Records of the Australian Museum
2017
Rec. Aust. Mus.
2017-08-23
69
3
119
222
http://dx.doi.org/10.3853/j.2201-4349.69.2017.1666
journal article
10.3853/j.2201-4349.69.2017.1666
d63960a1-e0c5-4b97-8e7d-f8be80d376d6
2201-4349
4677010
Recluzia johnii
(
Holten, 1802
)
Figs 2P, R
,
36B, D, I–J
Helix johnii
Holten, 1802: 76
(refers to Chemnitz, 1795: 284, pl. 210, fig. 2076).
Recluzia hargravesi
Cox, 1870: 172
, pl. 16, fig. 8;
Angas, 1877: 190
;
Tryon, 1887: 38
, pl. 10, fig. 23;
Brazier, 1894: 536
;
Hedley, 1907: 481
;
Iredale, 1910: 71
;
Iredale, 1929: 279
; Iredale & McMichael, 1962: 49.
Recluzia johnii
(Chemnitz)
.–
Jousseaume, 1872: 205
;
Tryon, 1887: 38
, pl. 10, fig. 24;
Hedley, 1910: 358
;
Winckworth, 1943: 148
.
Recluzia erythraea
Jickeli, 1882: 367
;
Tryon, 1887: 39
;
Thiele, 1928: 83
, fig. 6.
Type material
.
Holten (1802)
provided a list of binominal names from Martini & Chemnitz’s (1769–1795) famous but non-binominal iconography in a catalogue prepared for the sale of Chemnitz’s collection (
Winckworth, 1943
). Part of the collection was purchased for Peter the Great and is now in the Zoological Institute, Russian Academy of Sciences, St Petersburg (
Martynov, 2002
). Chemnitz’s illustrated shell is not in ZMUC (J. Knudsen ZMUC pers. comm.
07 Feb 1978
; AGB personal obs.). The possibility that Chemnitz’s (1795: 284, pl. 210, figs 1076–1077) illustrated specimen of
Helix johnii
remains in Chemnitz’s collection in the Zoological Institute, St Petersburg, was investigated at the writer’s request by B. Sirenko (pers. comm.
07 Sep 2012
); the illustrated specimen is not present. Chemnitz’s drawing shows a cream and pale grey shell with a tall spire, closely similar in shape and general appearance to the
holotype
of
Recluzia hargravesi
(
Figs 36I–J
), although with a still slightly taller spire, a shallower suture, and one more teleoconch whorl.
Jousseaume (1872: 203–206)
discussed the identity of the specimen, translated Chemnitz’s description and discussion into French, referred the specimen to
Recluzia
, and redescribed it carefully. Chemnitz received his specimen from “M. John, a missionary … living in
Tranquebar
, whom I must thank for many conchological riches. Whether it inhabits Coromandel,
Ceylon
, the
Nicobar Islands
or other parts of the East Indies, and whether it is a land, freshwater or marine shell he gave not the slightest information” (translation from
Jousseaume, 1872: 204
). The strongly and evenly inflated whorl shape, essentially smooth surface, circular aperture, faint spiral colour band at the continuation of the suture around the last whorl, and narrowly reflected inner lip agree with characters of
Recluzia johnii
of the present report. It seems feasible that this is the earliest name for a species of
Recluzia
, despite some early authors referring it to terrestrial gastropod groups such as
Pupa
Draparnaud, 1801
(i.e.,
Pupilla
Fleming, 1828
,
Pupillidae
; not
Pupa
Röding, 1798
,
Acteonidae
) or
Cochlogena
Férussac, 1821
,
Helicidae
.
Férussac (1821: 57)
cited the name as
Helix
(
Cochlogena
)
johnii
Chemnitz
, but added “Habit. Les grandes Indes. Espèce incertain pour le genre”, whereas
Gray (1825: 412)
simply listed it as
Pupa johnii
without comment.
Smith (1910: 203)
also doubted that this name referred to a
Recluzia
species because of its description by Chemnitz (1795) as “subkeeled”, but this appears to refer only to the colour band around the last whorl in Chemnitz’s (1795: pl. 210, fig. 2076–2077) drawings; the drawings show no obvious sculpture. Similar narrow, irregular colour bands or faint spiral grooves are present on many specimens of
Recluzia
(e.g., Habe & Tokioka, 1953: fig. 12). Jousseaume’s note seems to have been inspired by a specimen in his collection, in
MNHN
(
Figs 36B, D
). The name has not been applied to any other mollusc during the last century, to the writer’s knowledge, and
Jousseaume (1872)
and
Winckworth (1943: 148)
are followed in accepting it as the earliest name for a species of
Recluzia
.
Recluzia hargravesi
,
holotype
NHMUK
197432, stated to be from Miall River, Port Stephens,
New South Wales
,
Australia
, but corrected by the collector, W. Glover (cited by
Brazier, 1894: 536
) to the beach inside the North Head of Port Stephens. It is necessary to remove the name
Helix johnii
from uncertain status and associate it unambiguously with this species rather than
R. lutea
. The name
H. johnii
originally was not provided with a type locality. The
holotype
of
Recluzia hargravesi
, NHMUK
197432 (
Figs 36I–J
) is here designated the
neotype
of
Helix johnii
Holten, 1802
. Therefore, the type locality of
Helix johnii
becomes Port Stephens,
New South Wales
.
Thiele (1928
: fig. 6) illustrated the presumed
holotype
of
Recluzia erythraea
, ZMB
/Moll-112603 (E. Strong,
USNM
, pers. comm.
20 Sep 2012
; not seen), from Dahlak Island, Red Sea, showing that this name also refers to a tall, narrow specimen of
Recluzia johnii
.
Other material examined
.
Australia
:
New South Wales
:
Collaroy Beach, Sydney (
AMS
); Maroubra Beach, Sydney (
AMS
); Middle Harbour, Port Jackson (
NMV
); Port Stephens (two lots,
AMS
); Port Macquarie (
NMV
).
Queensland
:
Green I. (
AMS
C9824); Palm Islands (
AMS
C9823);
31–37 m
, Masthead I. (
AMS
C19351
a); trawled,
150–200 m
, Swain Reefs (
GNS
WM17360, 1); King’s Beach, Caloundra (
NMV
); Wreck I., off Yeppoon (
AMS
); Fairfax I., Bunker Group (
AMS
C69053
).
The only other specimens observed in world museums are the
holotype
of
Recluzia hargravesi
, listed above, and Jousseaume’s specimen in
MNHN
. A specimen in
MNHN
labelled “Rade de Batavia”, i.e.,
Jakarta
,
Indonesia
(E. Strong,
USNM
, photograph sent pers. comm. 20 Sep 20012) has a tall, narrow spire and possibly belongs in
R. johnii
, but is too incomplete for certain identification.
Distribution
. The tall-spired species
Recluzia johnii
is much rarer than
R. lutea
. In this study, almost all specimens observed are from
Queensland
and northern
New South Wales
, eastern
Australia
; none have been observed from
New Zealand
. Living specimens have also been collected on northern New South Wales beaches by
Riek (2017
;
Fig. 7
). However, the
type
specimens of the synonyms demonstrate that
R. johnii
also occurs in the northern Indian Ocean and the Red Sea. It probably occurs throughout the tropical Indo-West Pacific province. The writer is not aware of records from the Atlantic Ocean or the eastern Pacific, but this probably results merely from its rarity.
Dimensions
. See
Table 12
.
Diagnosis
. Moderately large (H to 39, D to
24 mm
), larger than
Janthina
species other than
J. janthina
and
J. globosa
; taller and narrower than all other neustonic
Epitoniidae
including
Recluzia lutea
; mean height: diameter 1.56, mean diameter: spire height 1.20. Teleoconch of 6–7 whorls; whorls strongly and evenly inflated, of almost circular crosssection, narrower than in
R. lutea
; suture deeply impressed; without obvious sculpture. Aperture almost circular, almost all specimens without anterior spout-like expansion. Lays smooth, narrow, cylindrical to weakly club-shaped, yellow egg capsules on underside of float, as in
R. lutea
.
Figure 36. Putative fossil
Recluzia
, and specimens of
Recluzia johnii
(Holten)
and
Recluzia lutea
(Bennett)
.
(A
,
C)
unidentified
Epitoniidae
formerly identified as
Recluzia
sp., NMB locality 17516, Casa Cantaure, Paraguana Peninsula,
Venezuela
; Cantaure Fm, Burdigalian (late early Miocene); SEM.
(B
,
D)
MNHN, Jousseaume collection, unlocalized specimen referred to
Recluzia johnii
(photos by E. Strong).
(E
,
H)
Recluzia lutea
, USNM
56411,
holotype
of
Lymnaea palmeri
Dall, Taqui River
delta, head of Gulf of Mexico, eastern Pacific (photos by E. Strong).
(F–G)
Recluzia lutea,
MNHN IM.
25667,
syntype
of
Recluzia jehennei
Petit de la Saussaye
and
neotype
designated here of
Recluzia lutea
Bennett
, of
Janthina turrita
Philippi
and of
Recluzia rollandiana
Petit de la Saussaye
; Gulf of Arabia (MNHN photos).
(I–J)
Recluzia johnii,
NHMUK
197432,
holotype
of
Recluzia hargravesi
Cox
and
neotype
designated here of
Helix johnii
Holten
; Port Stephens,
New South Wales
,
Australia
(photos by E. Strong).
(K–L)
Recluzia lutea,
MNHN IM.
25668, one of two
syntypes
of
Recluzia annamitica
Wattebled
; Thuan-an, Hué,
Vietnam
(MNHN photos).
(M–P)
Recluzia lutea
, range of shape of large specimens in one collection; Great Exhibition Bay, northern North Island,
New Zealand
,
Mar 1998
; in collection of P. Poortman (
Auckland
) (specimen in
Fig. 2S–T
is from the same sample). Scale bars:A,
1 mm
; C,
300 µm
; uppermost bar,
10 mm
, applies to all other figures
.
Table 12
. Dimensions of
Recluzia johnii
(*dimensions from original publications).
H,
height;
D,
diameter;
SH,
spiral height.
| species/locality |
H |
D |
SH |
H/D D/SH |
|
R. hargravesi
holotype, neotype of
R. johnii
|
38.8 |
22.9 |
22.7 |
1.69 |
1.00 |
|
R. erythraea
, from Jickeli*
|
13.0 |
8.5 |
— |
1.53 |
— |
|
R. turrita
, from Philippi*
|
14.5 |
10.0 |
— |
1.45 |
— |
|
R. johnii
, Jousseaume’s specimen, MNHN
|
23.3 |
12.6 |
9.3 |
1.85 |
1.35 |
|
R. hargravesi
, AMS
, Queensland
|
16.9 |
10.9 |
— |
1.55 |
— |
|
R. hargravesi
, AMS
, Queensland
|
22.7 |
14.8 |
— |
1.53 |
— |
|
R. hargravesi
, AMS
, Queensland
|
22.5 |
13.9 |
— |
1.62 |
— |
|
R. hargravesi
, AMS
, Queensland
|
7.5 |
4.5 |
— |
1.67 |
— |
| AMS C94285, Queensland |
22.2 |
14.8 |
14.0 |
1.50 |
1.06 |
| AMS C94286, Queensland |
24.5 |
14.8 |
16.0 |
1.66 |
0.93 |
| AMS C94287, Queensland |
17.9 |
12.2 |
10.3 |
1.47 |
1.18 |
| AMS C94287, Queensland |
20.7 |
14.7 |
13.5 |
1.41 |
1.09 |
| NMNZ M242099, Pearl Bay, Queensland |
35.7 |
24.04 18.3 |
1.49 |
1.31 |
| GNS WM17360, Swain Reefs, Queensland |
28.8 |
20.5 |
15.1 |
1.40 |
1.36 |
Remarks
.
Recluzia johnii
is so rare that little can be stated about it. It is concluded here that it is a second species of
Recluzia
with a taller, narrower spire with straighter outlines than in
R. lutea
, produced by its greater translation rate and slightly more numerous whorls of narrower cross-section. Most specimens have faint, irregular sculpture of weak spiral and axial grooves and ridges, varying individually, as in
R. lutea
. The teleoconch also tends to be a slightly warmer shade of yellow-brown than in
R. lutea
. Further material is required before the status and distribution of
R. johnii
is certain.
Riek (2017)
published on his web page several excellent photographs of specimens collected alive on northern
New South Wales
beaches, showing two specimens of
Recluzia hargravesi
(i.e.,
R. johnii
) feeding on the floating anemone
Actinecta
sp. The
Recluzia
specimens have extremely long, narrow, undivided cephalic tentacles, apparently much more extended than those in the photographs of
R. lutea
. However, the bright yellow animal, cylindrical egg capsules, and other characters are identical to those of
R. lutea
. It is notable that the floats of all specimens on this web page are as irregularly arranged as those of
R. lutea
in all other photographs observed, and it appears that the float of
Recluzia
species is less regular than and formed slightly differently from that of
Janthina
. Denis Riek (Brunswick Heads, NSW, pers. comm.
11 Nov 2015
) confirmed that the cephalic tentacles of all
Recluzia
specimens he has seen are undivided; the double appearance in some of his photographs is due to reflection on the undersurface of the water in the aquarium. He also stated that the photographed specimens ate the
Actinecta
anemones voraciously; they had consumed one anemone before he reached home and they could be transferred to his aquarium. Thereafter he was able to photograph them consuming the remaining anemone.
An unlocalized specimen from Jousseaume’s collection in
MNHN
(
Figs 36B, D
; point identified by “JJ” in
Fig. 35
; photographs provided by E. Strong, USNM, pers. comm.
20 Sep 2012
) is similar in spire height to Chemnitz’s (1795: 284, pl. 210, fig. 2076) drawing of the lost
holotype
of
Recluzia johnii
. It is unusually narrow, and has a still narrower appearance than other
Recluzia
specimens because its suture is markedly shallower and the spire outline consequently less strongly stepped than in any other
Recluzia
specimen the writer has examined (H 23.3, D 12.6, SH
9.3 mm
; H/D = 1.85, D/SH = 1.35; i.e., intermediate between the means of
R. lutea
and
R. johnii
; dimensions included in
Table 12
and in calculations of dimensions and means for
R. johnii
, including in
Table 11
). The anterior end of the aperture is flared narrowly. It is an unusually pale yellowish grey, similar to the colour of Chemnitz’s (1795: 284, pl. 210, fig. 2076) drawing, and paler than all other specimens of
R. johnii
and
R. lutea
observed by the writer. It also has seven teleoconch whorls, indicating that the whorls are still narrower than those of other specimens assigned to
R. johnii
. This specimen presumably inspired
Jousseaume’s (1872)
note on this species. It possibly represents a third living species of
Recluzia
. However, as only one specimen has been observed it must be assumed to be part of the variation of
R. johnii
until more specimens allow it to be evaluated more fully. The available material suggests that there are two species of
Recluzia
.