Salticidae: Diagnostic Drawings Library
by Jerzy Proszynski 2006
Phylogeny of Lapsiines and Hisponines Maddison & Needham 2006
No 0000 Ecuador
Source: MADDISON1 & NEEDHAM 2006. Lapsiines and hisponines as phylogenetically basal salticid spiders (Araneae: Salticidae). Zootaxa, 1255: 37, 45, f 1. (2006 - online edition) [http://salticidae.org/wpm/reprints/Maddison2006NewLapsiines.pdf]
p. 37
Morphological and molecular data have begun to resolve the basal
phylogenetic structure
of salticid spiders (Wanless, 1980, 1982, 1984, Rodrigo & Jackson, 1992,
Maddison, 1988,
1996, Wijesinghe, 1992, 1997, Maddison & Hedin, 2003). One of the best
corroborated
clades is the Salticoida (Maddison & Hedin, 2003), within which falls
the vast majority of
salticids, about 95% of the approximately 5000 described species (Platnick,
2005).
Excluded from the Salticoida are three much smaller groups: the lyssomanines,
the
spartaeines, and the Cocalodes group. Six extant Old World and 2 New World
genera are
placed in the Lyssomaninae (Wanless, 1980, Logunov, 2004); 15 genera, entirely
from the
Old World, are placed in the Spartaeinae (Wanless, 1984, Wijesinghe, 1992,
Zabka & Kovac, 1996); 4 Old World genera are in the Cocalodes group (Wanless,
1982, 1985). We
will refer to these non-salticoid groups as "basal salticids", not
to imply that they have
predominantly primitive characteristics, merely to indicate that they fall
outside of the
speciose clade Salticoida.
Basal salticids offer special insight into early salticid evolution for two
reasons. First,
if the basal salticids do not form a single clade, but diverged successively
from the line
leading to the Salticoida, then they will have a strong influence over inference
of the
family's ancestral states, whether parsimony or likelihood methods are used.
Second, even
if basal salticids do form a single clade, which therefore would stand equal
to the
Salticoida as an indicator of ancestral states, each sampled basal species
would have more
influence on ancestral state inference than each sampled salticoid species.
For these
reasons the unusual predatory behaviour (Jackson & Pollard, 1996, Li,
2000) and eye
anatomy (Blest et al., 1990) of basal salticids are of particular interest
in understanding the
origins of salticid diversity. By recognizing what species are among the basal
groups and
how they are related, we will be able to characterize better the early evolution
of salticids.
In addition, we will have more complete outgroup information for reconstructing
phylogeny within the Salticoida.
In this paper we present molecular data to examine whether two little-studied
groups
of salticids might belong with the lyssomanines and spartaeines as basal salticids:
the
lapsiines (Maddison, 2006) and hisponines (Wanless, 1981, Prószy½ski
& òabka, 1983,
Weso»owska, 1993). Salticid systematists have made few comments on where
these groups
belong. Simon (1901) apparently considered lapsiines and hisponines relatively
primitive,
but he also considered various salticoids as equally primitive. Simon conferred
suggestive
names on one lapsiine ("Lapsias cyrboides") and one hisponine ("Tomocyrba")
that hint to
similarities with a spartaeine, Cyrba Simon. Prószynski & Zabka
(1983) placed
Tomocyrba Simon within the Euophryinae on the basis of the spiraled embolus.
The neotropical lapsiines include Lapsias Simon and two recently discovered
genera
(Maddison, 2006). Their basal phylogenetic placement is suggested by the presence
of a
tarsal claw on the female palpus, loss of which is considered a synapomorphy
of the
salticoids (Maddison & Hedin, 2003). In addition, the male palp has an
extra sclerite
associated with the tegulum, presumably homologous to the median apophysis
of
Cocalodes Pocock and Holcolaetis Simon (Wanless, 1982, 1985). The salticoids
and
spartaeines are characterized by the loss of this sclerite. If the lapsiines
are confirmed as
basal salticids, it would show that the New World has a previously unrecognized
radiation
of basal salticids.
Hisponines are Old World salticids primarily from Africa. The three extant
genera—
Hispo Simon (Wanless, 1981), Massagris Simon (Weso»owska, 1993) and
Tomocyrba
Simon (Prószy½ski & òabka, 1983)—are distinctive
for a constriction on the carapace just
behind the posterior median eyes. Our attention was drawn to hisponines by
two
observations. First, as in the lapsiines, the tegulum in many species has
a small sclerite that may be homologous to the median apophysis of other basal
salticids. Second, Baltic amber ZOOTAXA
salticids are dominated by two body forms, the first with the small eyes unusually
large
(placing them among the basal salticids), the second with a distinctive constriction
behind
the small eyes. As implied by Prószy½ski & òabka
(1983), this constriction suggests that
the Baltic amber genera Gorgopsina Petrunkevitch and Prolinus Petrunkevitch
are
hisponines. If Baltic amber has hisponines but not salticoids, hisponines
may have a
relatively ancient divergence from other salticids.
We therefore obtained molecular data from hisponines and lapsiines to determine
whether, as hinted by their morphology, they lie outside the Salticoida. We
also include
data from two nuclear genes not previously used in salticids (18S, Histone
3) and for
several other genera not previously studied.