BALLINAE
RELATIONSHIPS
Initial cladistic analysis of 29 equally weighted, informative characters produced three most parsimonious trees, with a consistency index (CI) of 0.66, a retention index (RI) of 0.68, and a length of 69 steps. The analysis was then repeated twice after successive character weighting ( Farris, 1969; Carpenter, 1988), using the maximum value of the rescaled consistency index. This analysis produced a single tree (length = 34 steps, CI = 0.90, RI = 0.90, rescaled CI = 0.81), identical in topology to one of the trees produced in the unweighted analysis. This tree is presented as the preferred hypothesis of
Ballinae
generic relationships ( Fig. 1
View Figure 1
).
The subfamily
Ballinae
is resolved at node 1, supported by the presence of ec (2-1), ec that coils at least once over 360∞ (4-1, 5-1, 5-2), dark band on both lateral sides of legs I–IV (27-1) and enlarged femur (30-1). Within
Ballinae
, the presence of an oval, bilobed tegulum (1-1), narrow epigynal septum (11-1) and the presence of a translucent septum (18-1) support node 2. Species that were formerly placed in
Marengo
(except for
Cynapes canosus
and
Sadies
) are united at node 4 as sister to
Colaxes
+
Ballus
(node 6). Node 4 is supported by the presence of a convex embolic coil (3-1), white patches on sides of opisthosoma (23-1) and lts (33-1); node 6 by the presence of 2/0/0/2 spines on tibia I (35-0). Nodes 4 and 6 are united at node 3, supported by the presence of a dorsoventrally flattened prosoma (20-1) and an enlarged tibia I (31-1).
Copocrossa Simon, 1901
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,
Mantisatta
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and
Padilla
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are included in
Ballinae
, although only
C. tenuilineata ( Simon, 1900)
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has been studied. The male genitalia of
C. tenuilineata
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are similar to those of other
Ballinae
. The same can be said for
Padilla
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, a genus of seven species, all from Madagascar. Judging from illustrations in Proszynski (2003) they may be included in
Ballinae
, as male genitalia show
Ballinae
synapomorphies: presence of ec (2-1) and spiralling embolus (4-1). Unfortunately, drawings are only available for
P. armata Peckham & Peckham, 1894
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, and
P. cornuta ( Peckham & Peckham, 1885)
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.
Mantisatta
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consists of two known species:
M. longicauda Cutler & Wanless, 1973
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from the Philippines and
M. trucidans Warburton, 1900
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from Borneo.
M. longicauda
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is the only species known from both sexes. Judging from illustrations of male genitalia in Proszynski (2003) and Cutler & Wanless (1973: figs 5–7) they are probably correctly placed in
Ballinae
. Their genitalia show
Ballinae
synapomorphies: presence of ec and a terminally coiled embolus. However, they possess enlarged femur I and tibia I, but lack lts.
Admestina
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,
Attidops
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and
Cheliferoides
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have been tentatively included in
Ballinae
by various authors. Maddison (1995, 1996) included
Admestina
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in
Ballinae
, based on male genital structure. Edwards (1999) tentatively included
Attidops
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in
Ballinae
, based mainly on male genital characters: presence of a pale longitudinal furrow, well-coiled embolus lying flat on the tegulum, unequally bilobed tegulum and presence of a flattened carapace. Platnick (1984) observed that
Cheliferoides
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possess a palp similar to that of
Marengo
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. However, I do not consider any of these genera to be correctly placed in
Ballinae
due to the absence of ec (2-1), a spiralled embolus that coils at least once (4-1, 5-1, 5-2).
Maddison (1995, 1996) used the following two characters to place
Admestina
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and
Cheliferoides
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in
Ballinae
: (1) a well-coiled embolus lying flat on the tegulum, and (2) a tegulum which is divided by a pale longitudinal furrow. Emboli of
Admestina
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and
Cheliferoides
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coil less than once around the tegulum, they are broad-based, stout and project outwards;
Attidops
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has a longer embolus ( Edwards, 1999). While the presence of an oval, bilobed tegulum in
Admestina
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and
Attidops
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suggests placement within
Ballinae
, the lack of an embolic coil means that their affinity to the subfamily remains unclear. These genera might form a taxon that is sister to
Ballinae
. The presence of an unequally bilobed tegulum (1-1) and flattened carapace (20-1) might turn out to be plesiomorphic for
Ballinae
. Thus, the phylogeny of
Ballinae
needs to be reassessed in future studies.
Salticidae
is the largest spider family, with 4834 described species and 531 genera ( Platnick, 2003). Many have been inadequately described and estab- lishing their evolutionary relationships is an enormous task. It is my hope that the present study will go some way towards achieving this goal and encourage other workers to examine scales and other morphological characters, together with cheliceral dentition and genitalia.