Shells from Alfred Russel Wallace in
Alfred Russel Wallace
(1823-1913) is widely
l<nown as the 'father of
biogeography' and for
the j oint announcement,
witlr Charles Darwin, of
the theory of evolution
by natr-rral selection
(figure I, left, in a
photograph taken in
Singapore in 1862
(public domain photo)
and figure 4). The recent
death has seen a revival
in interest in Wallace
and brought his work to
wider audiences. In
Wales he has become
something of a local
hero (whether he was
'oflicially' Welsh or not is another question). The range of
frelds to which he contributed can be seen from his two-
volume 'The Malay Archipelago...', which records his
eight-year voyage ofdiscovery in Southeast Asia as a
professional collector-naturalist. One of these fields was
conchology. Wallace himself records collecting nearly 7500
shells during the Malay Archipelago years, tal<ing any
opporlunity, br-rt favoLrring land snails in particular (Wallace,
1 869a,b). Perhaps it was already clear to him that terrestrial
molluscs had great potential in biogeography. Much of the
biological material Wallace collected was intended for
immediate sale to museums and collectors in Britain and
beyond. He shipped it in consignments to his London agent,
Samuel Stevens. Samuel was the brother of the auctioneer
John Crace Stevens ofCovent Garden (Baker,200l) whose
auction house was the venue ofthe great shell sales ofthe
1860s onwards (Dance, 1986). Although it is commonly
believed Wallace travelled and collected mainly to earn
money through sales, he himself regarded this as a
secondary objective and kept his best specimens for his own
collection and study (C. Beccaloni, pers. con.rm.). Those
specimens that Wailace instructed Samuel Stevens not to
sell included material lor Wallace's own collection and that
put aside tbr the entomologist William Wilson Saunders
Soon after his return fiom Asia, Wallace (1865) published
his own list of Malay land snails. He fbund that his
expeditions had collected 125 species, ofwhich over 50
were new, and had provided the first accurate localities fbr
rnany of the others. At the end of the paper, Henry Adams -
the less well travelled of the two Adams brothers. but no
less an authority on Asian land snails wrote the
descriptions for Wallace's last f-ew new snail taxa. Although
the types and other shells were said to be in Saunders'
collection, their whereabouts are now unclear. The
Moluccas work of Woutera S. S. van Benthem Jutting (e.g.
1941, 1953, 1958, 1959) cites several of the Adams/Wallace
species. She examined collections in London, Frankfurt,
Leiden and Amsterdam (1959), and mentior.rs at least one
'original lot described by Adams after Wallace's
the National Museum of Wales
Ben Rowson* and Harriet Wood
collections' in London (1958: p.322), although its
dimensions do not match those given by Adams in Wallace
(1865). Saunders' collections were sold at two auctions in
1874, and the majority of his insects are now at the Oxford
University Museum of Natural History. However, Saunders'
shells have not yet been located at Oxford or at the Natural
History Mnseum, London (D. J. Mann & J. Ablett, pers.
Wallace (1865) says that all his shells were identified with
reference to the incomparable collection of Hugh Cuming,
which is now at the Natural History Museum, London. As
Cuming's letters to others reveal, he had been a customer of
Wallace (or Stevens) during the Malay Archipelago years
(Dance, 1980), including obtaining several Moluccas
species described by Louis Pf-eilier (e.g. Pfeiffer, 1861). As
Cuming's collection remained in private hands until
Cuming's death in 1866 (Dance, 19t36), Wallace must have
made use of it befbre then and one wonders whether the two
men l<new one another personally. Both knew Darwin, of
course. and had much else in common as both field
collectors and dealers (Dance, 1980). Alternatively,
Sar.Lnders or Adams cor-Lld have done the comDarisons for
Wallace without hin.r meeting Cuming himself.
At this time, James Cosmo Melvill (1845 1929) was
beginning to build up his own collection. Eventr-rally, when
combined with that of John Read le Brockton Tomlin. this
was to become the biggest ever in private hands in Britain
(Dance, 1 986). MelvilI later wrote an admiring obituary of
Cuming, whom he encountered at one of J. C. Stevens' shell
sales when Cuming was an old man (Melvill, 1895). He
does not say whether they corresponded, but does point out
one further similarity between Cuming and Wallace. Melvill
notes that in SoLrtheast Asia, both collectors obtained many
land snails by paying local children to look in woods and
forests, and also carried quinine and other medicines, each
'practising as a nredicine man' (Melvill, 1895). This
apparently aided relations with locals as well as at least in
Wallace's case treating people directly and possibly
saving his own life (Wallace, l869a,b). Melvill says the
effect was to raise superstitions about the travellers. Wallace
(1869b) wrote that the people of the Aru Islands, New
Cuinea were 'imputing hidden medical virtue' to the land
snails he collected. This, he thought, was their explanation
of why he kept snails while rejecting the common, nacreoLrs
marine specimens they brought him.
The Melvill-Tomlin collection continued to be built up by
purchase and exchange until the 1940s, before being
bequeathed to the National Museum of Wales in 1955. Lists
of secondary sources, with whom Melvill or Tomlin
corresponded or obtained material from (Trew, 1987, 1990)
suggest shells from Wallace could easily have made their
way into the collections. Finding and investigating the
relevant shells is now immeasurably easier, thanks to the
careful curation and item-documentation of the Melvill-
Tomlin collection during the late 1990s and early 2000s.
The publication of many of Wallace's notes in recent years
is also an aid to establishing what he collected (Baker,200l;
van Wyhe & Rookmaaker,2013; see also .,
*ben.rowson (rf; nr useumwales.ac.uk
The Wallacean land snails
Our investigation shows the Melvill-Tomlin collection
contains at least eight lots of tropical Asian land snails with
labels reading either'Wallace', 'Wallace coll.' or ,A. R.
Wallace coll.' in a range of handrvritings, including
Melvill's and Tomlin's. Some labels include orices in
shillings. The lots are labelled, and databased, as belonging
to eight different species, most of which are relatively large
and 'showy' (figures 2 and 3). Five of the species were
mentioned in Wallace's snail paper (1365). Six of the
species belong to the Helicoidea and are from various
islands of the Moluccas (Maluk-u Islands), Indonesia. The
Moluccas lie between continental shelves in the regron now
known to biologists as Wallacea (see van Benthem Jutting,
e.g. 1941,1953,1958, 1959 for the snail fauna). Wallace
spent several years visiting almost all the major Moluccas
islands, collecting and exporling land snails from many of
them (Wallace, 1865; Baker, 2001). Famously, ir was in the
Moluccas in 1858 that Wallace wrote up his ideas on
evolution by natural selection the 'Ternate Essay' - and
sent them to Darwin. This led to the jointly credited
announcement ofthe theory at the Linnean Society of
London later that year, although the sequence ofevents is
not without its controversies (van Whye & Rookmaaker,
Five of the helicoid lots belong to Camaenidae, and one to
Bradybaenidae. The bradybaenid is a shell labelled ,Hel.
pyrostoma Ind Islands' to which Tomlin has added ,1512
Wallace Gilolo' and'v. nigrescens Kobelt.' The shell seems
corectly identified with Phania pyrostoma (Fdrussac, I 821)
from Gilolo (Halmahera) in Wallace's list. Wallace even
singled this species out in the Malay Archipelctgo as ,very
fine and handsome' among those found by a local on Gilolo
(Wallace, I869b); Pilsbry (1891) also considered it .a
magnificent species'. The variety nigrescens Kobelt was
acknowledged (in Additions and Corrections) by pilsbry
(1894). The genus (the name Phania was later replaced with
Pyrochilus) is endemic to Gilolo, Batchian (Bacan) and
Ternate in the Moluccas (Pilsbry, 1891; 1894; van Benthem
Jutting, 1 959; Schileyko, 2004).
The camaenids all belong to the genus Plani.spira, which
ranges from Sulawesi and the Moluccas to New Guinea
(Pilsbry, 1891, 1894; van Benthem Jutting, 1959; Schileyko,
2003). One lot of two shells has an old label reading .p.
kuni Pfr. Batchian Wallace Coll.' (handwriting unknown;
see Leptopoma below); Batchian was a locality given for p.
kurriby Wallace (1865). Two other lots, each containing
one shell, are from localities not recorded by Wallace for
their respective species. One has an old label in a different
unknown handwriting reading 'Planispira zonalis Var [?]
Wallace Batchian 1 at 4l- [i.e., 4 shillings]'. Tomlin has
added 'Fer.' in pencil. Wallace records P. zonalis Fdrussac
only from Gilolo. The other has a label in Torrlin's
handwriting reading 'Planispira expansa pfr. Bouru
(Wallace)'. Wallace (1865) records P. expansa pfeiffer only
frorn Batchian, although lists several other snails from
Bouru (Buru), where he collected widely (Baker,2001). lt is
possible that there has been some confusion over
identification, but both are a good match for illustrations in
Pilsbry (1891), who also extended the known distributions
of both species. So neither their names nor localities rule out
their having been collected by Wallace, especially if they
were despatched to Europe before being identified.
The remainingtwo Planispira lots, both from Ceram
(Seram), bear species names introduced by pilsbry ( I 991 ),
long after Wallace's return to Britain and disposal of what
he had retained as his personal collections (Wallace, lg65;
Baker,2001). Pilsbry described both species from the
Moluccas without giving a further locality or collector. One
lot, containing one shell, is labelled 'Planispira iaddae
Pilsbry Ceram 5/-'. lt seems correctly identified. It also
contains a pale blue label reading 'Wallace' in unknown
handwriting. The other lot, containing two shells, is labelled
'H nitidiuscula 4 at3l: Tomlin has added .Bttg.' and a
second label in his handwriting reads 'Ceram Wallace 2g,.
The identification with P. chariessa recorded on the label by
A. Trew (c. 1997) seems less certain. According to pilsbry
(1891) both species are similar to the widespread and
variable P. zonaria (Linnaeus, 1767); indeed, the name
nitidiuscula Boettger, I 891, described from Amboina
(Ambon) is considered a form of P. zonaria by pilsbry
( 1 8e4).
Finally there are the two non-helicoid 'Wallace' lots in the
Melvill-Tomlin collection. One contains three shells of
ariophantoids, with three labels. Apparently the oldest label,
in unknown handwriting, reads 'Borneo Wallace Xestia
citrina var. albocincta 3 at3l: . The reverse of this label
reads'large! Citrina [;] 2 small fillegible] sp 1,. There are
two labels in Tomlin's handwriting. One reads .X. citrina L.
v. albocincta Borneo Wallace'; the other reads .v. coagulata
Pfr. f. Fulton'. If the 'f.' is for 'f,rde' it would appear that the
dealer Hugh Fulton reidentified at least one of the shells at
sonre point; Helix coagulata Pfeiffer, 1856 was described
from Amboina. Wallace (1365) was aware of the well-
known species 'Nanina' li.e. |tlaninia] citrina (Linnaeus,
1758), at least when identifying his shells in London. He
listed it from several islands of the Moluccas and noted its
variation in colour and pattern. Wallace had also, of course,
collected widely in Borneo and recorded shipping land-
snails home (Baker, 2001). The problem is that 1y'. citrina
and other Naninia are species of the Moluccas and New
Guinea (van Benthem Jutting, 1959; Schileyk o,2002).
Borneo is on the other side of the famous biogeographical
divide established by Wallace that became known as the
Wallace Line. Wallace himself would have been unlikely to
have made any mistake in this regard, noting that snails
generally followed the same pattern (Wallace, 1865).
Although some of the genera he records on both sides of the
line include superficially similar species, we found it
difficult to refute the identification of the Bomreo shells as
N. citrina, so the locality and identification seem to conflict.
The eighth lot comprises one shell of a caenogastropod with
the operculum in place. It is labelled 'Leptopoma Batchian
Wallace Coll.', in the same handwriting as appears on the
lot of Plctnispira kuwi. Another label, in the capitalised
handwriting of Hugh Fulton. reads 'Leptopo-u flurn,o.u,,
Pf. Ceylon Batchian ld'. On the reverse, Fulton has written
'Batchian Id is probably correct'. Wallace's Malay paper
(l865) lists eight species of Leptopoma from Borneo, the
Moluccas and New Guinea, but of course none from Ceylon
(Sri Lanka). He is known to have visited Sri Lanka only en
route to and from Europe - to change ships at Galle in 1g54,
and to call briefly in 1862 (van Wyhe & Rookmaaker,
2013). However, the shell does resemble the Sri Lankan
speci es L ep t o
p om o ides fl a mm eus' (Pfei ffer, 1 8 5 4), as fi gured
by Reeve (1862) and Naggs & Raheem (2000). lt does not
look like any Moluccas Leptopoma we have been able to
flgLrre 2: Latrd snails rrith \\rallace associatiors in the McLvill-Tonrlrn colLectiott. Natiorral MLtscttttt of \\'ales (.r)
21 ivu,ri,.couchsoc. org
figure 3Land snails with Wallace associations in the Melvill-Tomlin coliection. National Museurn of Wales (b),
examine. As with the Nanini.q shells, the Wallace locality
and specimens seell to conflict. Interestingly, Fulton gave
his opinion on both lots and may have had a similar
problem; van Benthem Jutting (1959) gives other examples,
sonre relating to Wallace.
It seems highly likely that the Melvill-Tomlin shells were
indeed collected by Wallace from the Moluccas. The two
non-helicoid lots are more dubious, given the mismatch
between their identities and localities, so some mir-up may
have occurred. The diversity of labels among the shells,
some clearly from dealers, suggest they entered the
collection by various routes, some perhaps bought from
Samuel Stevens while Wallace was still overseas. and even
from the period in which he was forrrr-rlating his ideas on
As Wallace's 1865 paper suggests, snails like these were
among those that helped lead to his famous conclusions on
species and biogeography. His name was evidently familiar
to conchologists ofhis day, his conchological career
overlapping that of Cuming and Melvill, and may well have
known some collectors personally. Furlher conclusions on
Wallace's molluscs would demand a more detailed study of
his material in London and elsewhere.
www. conch soc. org
Ac k nowledge ments
We thank George Beccaloni (Natural History Museurn, London)
lbr helpfirl commeuts, and Jon Ablett (Natural H istory Muser-rm)
and Darren Mann (Oxtbrd University Museum ol'Natural History)
for their help with information on the Saunders collection.
Baker, D.B. (2001). Alfied Russel Wallace's recold of his consignnrents ro
Sanruel Stevcns, I 854 1 861 Zoohgische l.lededelingen Leiden 75 (16):
van Benthenr Jutting, W.S S. (l9.tl). On a collection of Non-Marine
Mollusca from the Talaud lslands and From Molotai (Moluccas). Treubia
18: I 27.
van Benthcm Jutting, W.S.S (1953). Annotated ltst o1'the non-nrarine
Mollusca o1'the Moluccan Jslands Atnbon, Harulku, Saparua and Nusa LaLrt
van Benthenr JLrtting, W.S.S (1958) Non-marirre Mollusca ol'the lsland o1'
Misool. Nova Cttineu 9:293 338
van Benthen .lutting, W.S S. (1959). Non-rrarine rnollusca olthe North
Moluccarr lslands Haln-rahera, Ternate, Batjan and O6i Treuhh 25:25 87.
Dance, S.P. (1980). Hugh Curling prince ofshell collectors Journal of
the Society for the Bibliograph.v o/ Naturtrl Histott 9:477 501 .
Dance, S.P. (1986) I History o./ Shell Collecting. E. J Brill W
Davics, R. (2012). How Charles Darwil received Wallace's Ternate paper
l5 days earlier than he clainred: a col.).1'ner1L orr van Whye & Rookmaaker
(2012) Biologitul Jottrnal of the Linnean SocieD: 705 412111
Melvill, J C (1S95) An epltome of the life of the late Hugh Cuming' FLS'
CMZS etc Journal of Conchoktg 8: 59-70
Society oJ London 1861:2029'
Pilsbry, H.A. (.1891) Manuttt oJ Concholog: st:yc.tu:1l ond $)stemdtic'
wilh iiluslrtttions o[ the species' Ser' 2, Vol VI: Helicidae' Vol l Acadenty
of Natural Sciences, Phi ladelph ia'
Pilsb 894). Manual of Con and tystematic'
tvith s of the speciei. Ser. e' [ol 7: guide lo
the s lces. AcademY of Na adelPhia'
Reeve, L (1862). Monograph ofthe genus Leptopoma Conchologica
lconica 13l 8 Pls
Schileyko, A.A. (2002). Treatise on Recent terestrial pulmonale molluscs'
Part 9: Heiicarionidae' Gymnarionidae, Rhysotinidae' Ariopharrtidae'
Ruthenica, SuPPlement 2 Moscow'
Schileyko, eat rrestrial pulmonate molluscs'
Part I l: 1'r ae, e, Vitrinidae' Limacidae'
Bielziidae, ,B amaenidae Ruthenica'
Supplemenl 2 Moscow. molluscs,
Trew, A. ( 1987). .lames Cosmo Melvill's new molluscan names National
Museum of Wales, Cardiff.
Trew, A. (1990). John R le B. Tornljn's new molluscan narnes National
Museum of Wales, Cardifi
Wallace, A.R. (1865). L d shells collected by Mr' Wallace in
the Malay Archipelago, tions of the new species by Mr' Henry
Adams. -Proceetiings tf cil Society d l'ondon ( 1865): 405 41 6'
Wallace, A.R. (1869a). The Malay Archipelago: the land oflhe oranF;-utan'
anr! the brcl o1'paratJise' with slttdies of man and nature Yttl' L
MacMillan & Co, London.
Wallace, A.R. (tS69b). The Malay Archipelago: the land.of lhe orong-Ltnn'
antl the iird ttf paradi.se' with studies of man and naltLre Y ol ' 2'
MacMillan & Co, London.
aaker, K (201 2)' A new theory 1o erplain the
nate Essay by Darwin in l858 Blological 'Iournal
van Whye, J., & Rookmaaker, K' (eds ) (2013) Alfred Russel Wallace:
letters fiom the Malay Archipelago Oxford University Press' Oxford'
figure 4: poftrajt of Alfred Russel Wallace and his signature on the
fr-ontispiece of Darwinism (1839) (public domain image)
Errata,Mollusc World issue 36 (Nov' 2014)
Readers will also have noticed that the page numbering was
omitted from issue 36. My apologies for this, which only
came to light after the issue was printed' Despite the
Society's iery limited resources every attempt is made to
ensure accuracy, but inevitably errors will occasionally
continue to creeP in!
Another excellent issue of Mollusc World' The new he ad
teacher t the
speed r where he
now pr me to
keep up the good work,
A11 the best,
Chris du Feu