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]
Morphological and molecular data have begun to resolve the basal
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.