An Updated Classification of the Recent Crustacea

have been used inconsistently in the past and that
there are no rules governing the name given to a
taxon above the family-group level. Thus, according
to McLaughlin and Holthuis, the Rule of Priority
need not be applied (Anomala is, strictly
speaking, the older of the two names). Furthermore,
they argued that, for stability, the name Anomura
MacLeay, 1838, should be used for the taxa
traditionally considered to belong to this group
(lomisoids, galatheoids, paguroids, and hippoids),
and we have followed their suggestion. Phylogenetic
relationships within the Anomura remain largely
unsettled; studies addressing this question include
McLaughlin (1983b), Martin and Abele (1986),
Cunningham et al. (1992), Tudge (1997b), Mc-
Laughlin and Lemaitre (1997, 2000), and Morrison
and Cunningham (1999).
McLaughlin (1983a) recognized the unusual nature
of Lomis hirta and placed it in its own family
(Lomidae) and superfamily (Lomoidea) (corrected
herein to Lomisidae and Lomisoidea, respectively).
McLaughlin (1983b) concluded that the hermit
crab families were monophyletic, and she therefore
treated all six families as members of the superfamily
Paguroidea. This arrangement has been adopted
by a variety of workers (e.g., Tudge, 1991; Richter
and Scholtz, 1994; Scholtz and Richter, 1995; Tudge,
pers. comm.) and seems to us both logical and simple,
and we have used it here. In his treatment of
the Pylochelidae (treated as Pomatochelidae in
Bowman and Abele, 1982), Forest (1987) indicated
that the family is more closely allied with the Diogenidae
than with other anomuran families, but
we have not indicated this alliance pending formal
recognition of that relationship.
The family name Lomisidae and the superfamily
name Lomisoidea, containing only the monotypic
genus Lomis, occasionally have been spelled, beginning
with Glaessner (1969), as Lomidae and Lomoidea
(see especially McLaughlin, 1983a). However,
the genus Lomis is not a Greek or Latin word,
and thus it has no Greek or Latin stem (such as
Lom-) to which the -idae ending can be added; the
original author of Lomis, Bouvier, coined the
French common name ‘‘Lomisinés’’ for these crabs
(G. Poore, pers. comm.). Thus, the preferred spelling
for the family is Lomisidae and for the superfamily
is Lomisoidea.
A recent analysis of anomuran phylogeny based
on mitochondrial DNA gene rearrangements (Morrison
and Cunningham, 1999; C. Morrison and C.
Cunningham, pers. comm.) largely supports Mc-
Laughlin’s (1983b) recognition of the major anomuran
groups and their phylogeny. According to
the findings of Morrison and Cunningham (1999),
lithodids are strongly associated with pagurids and
together these groups constitute a monophyletic
clade (confirming the earlier report by Cunningham
et al., 1992). The Hippoidea is also strongly supported
as a monophyletic clade, and the Galatheoidea
(including both Aegla and Lomis) is depicted
as basal to the remaining Anomura. Thus, a classification
based on these data would differ from
McLaughlin’s (1983a, b) in that the superfamily
Lomisoidea would be removed, with the monotypic
Lomisidae being placed within the Galatheoidea
(which also contains the Aeglidae, Porcellanidae,
Galatheidae, and Chirostylidae; see Baba (1988)
for a thorough review of the latter family). However,
support for this particular node (placement of
Lomis) was not as strong in the Morrison and Cunningham
tree, and indeed C. Morrison (pers.
comm.) has suggested that we might be better off
depicting a separate lineage for Aegla and Lomis
from the remaining galatheoids. We have for now
retained Lomis in its own family and superfamily,
the Lomisoidea, which we have placed adjacent to
the Galatheoidea as a concession to the new data.
Similarly, we have moved the Paguroidea closer to
the Hippoidea, also reflecting the findings of Morrison
and Cunningham (1999). Several workers
have discussed the fact that the lithodids (at least
some of them) appear to have stemmed from within
the Paguridae (Cunningham et al., 1992; Richter
and Scholtz, 1994; Tudge, 1991, Tudge et al., 1998;
C. Morrison, pers. comm.). Additionally, Cunningham
(pers. comm.) suggested a rather close tie between
the Aeglidae (restricted to freshwater streams
and lakes in temperate South America) and the
Lomisidae (a monotypic and exclusively marine
family known only from Australia). According to
Scholtz and Richter (1995), two groups of the Anomura,
hippoids and galatheoids, share the apomorphic
character of a telson stretch receptor not
found in any other malacostracan group (Scholtz
and Richter, 1995, citing Paul, 1989).
In contrast with the phylogenetic hypotheses of
McLaughlin (1983b) and Morrison and Cunningham
(1999), evidence from sperm ultrastructure
(reviewed in Tudge, 1997b) would suggest that the
Anomura is not monophyletic, that Lomis does not
belong to the Anomura sensu stricta, that at least
some of the thalassinoids are within the Anomura,
and that the superfamilies Thalassinoidea, Paguroidea,
and Galatheoidea are not monophyletic. Because
at this time the bulk of the evidence (i.e.,
adult morphology combined with molecular sequence
and gene arrangement data) seems to support
the more conservative approach of Mc-
Laughlin (1983b), we have modified our arrangement
of anomuran taxa only slightly. Our classification
is therefore more in agreement with the
findings of Morrison and Cunningham (1999) than
with the sperm ultrastructural findings presented by
Tudge (1997b).
In the Bowman and Abele (1982) classification,
the hermit crab families were divided among two
superfamilies, Coenobitoidea and Paguroidea. The
Coenobitoidea was removed following the suggestion
of McLaughlin (1983b), and the family Coenobitidae
is now treated within the superfamily Paguroidea.
Thus, our infraorder Anomura contains
four superfamilies: Lomisoidea (the distinctness of
which is questionable in light of the Morrison and
48 Contributions in Science, Number 39 Rationale